Chapter (non-refereed)

Welch, R. C.. 1986 What do we know about in Scottish woods? In: Jenkins, D., (ed.) Trees and wildlife in the Scottish uplands. NERC/ITE, 95-100. (ITE Symposium, 17).

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Contact CEH NORA team at [email protected] 95 What do we know about insects in Scottish woods?

R C WELCH Institute of Terrestrial Ecology, Huntingdon

1 Introduction distribution in Scotland. In the 1960s and early 1970s, Although this symposium is primarily concerned with teams of entomologists, funded by the Nature Conser- woodlands in the Scottish uplands, I have, by neces- vancy and Shell Chemical Company, surveyed several sity, had to take a wider overview of insects in Scottish areas of Scotland, paying particular attention to Nation- woodlands as a whole. However, information from the al Nature Reserves (NNRs). The resulting lengthy Forestry Commission (FC) (1985) and from Davies species lists are not suitable for publication and most (1985) shows that over the past 65 years the land now reside in various Nature Conservancy Council under trees in Scotland has risen from 3% to 14%, files. However, accounts of one such survey have mostly in the uplands, which is a loosely defined area been published for the Isle of Rhum (Steel & Wood- comprising about 42% of Scotland. Almost without roffe 1969; Wormell 1982). Despite the small size and exception, this planting has involved conifers. Indeed, recent origin of woodland of both deciduous and broadleaved woodlands probably occupy no more than coniferous species, the accounts contain reference to 10% of the 1 Mha of commercial and 'unproductive' many woodland insects. Woodroffe (1974) also com- woodland in Scotland. Clearly, in any examination of pared Hemiptera found on Rhum with those collected the insects occurring in Scottish woodlands, those of in other surveys based on Speyside, Deeside, and at the conifer plantations should take precedence. Invernaver, but interpretation of the species list Although rides, streamsides and cleafings within requires a previous knowledge of the biology of this forests are refuges for many sPecies, these group of insects. sites have been discussed elsewhere during this symposium (Young 1986), and I shall not consider Regrettably, most insects found in Scottish broad- them further. leaved woodlands are, with very few exceptions, much more abundant in the woods of England and Apart from the small band of dedicated FC staff, most Wales. Many species of insect appear to have spread entomologists visiting or resident in Scotland have into Scotland along the milder coastal regions, so that largely ignored the coniferous plantations. Instead, broadleaved trees in Caithness, Sutherland and even they seek out the relict areas of 'Caledonian' pine on the Orkneys have what can be regarded as a diluted forest or concentrate on the deciduous woodlands, southern fauna, with the addition of a few northern especially the birchwoods (Betula spp.) of the High- elements. However, our present knowledge of the lands. In both these ecotypes, their quest is for insect fauna of the western oak/birch woodlands is species of insects with distributions in Britain re- very limited, and many such woods are either totally stricted to the Highland forests. Such species may unknown entomologically or such information as does have an arctic—alpine distribution in continental exist resides in private collections and notebooks. It is Europe. Most entomologists give only a cursory mainly in the central Highlands where the insect fauna inspection of the outer margins of the dark serried differs markedly. I have previously described typical ranks of Sitka spruce (Picea sitchensis) and lodgepole examples from a wide range of habitats (Welch 1974, pine (Pinus contorta) plantations, whilst the attention 1981) and will not consider Scottish broadleaved of the FC entomologists is usually only attracted when woodlands further here. a particular insect species reaches epidemic propor- tions. It will, therefore, come as no surprise to learn 3 Coniferous woodlands that our knowledge of the entomology of Scottish In 1974, the Nature Conservancy set up a Native upland forests is somewhat scanty. That which does Pinewoods Discussion Group as a forum for owners, exist is largely scattered through the literature and is managers and scientists concerned with the conserva- difficult to find and abstract. My intention here is to tion and study of native pinewoods (Pinus spp.) in provide an introduction to the major sources of Scotland. This initiative culminated in a symposium information in this field, without too much specific volume in which Hunter (1977) provided an excellent detail. review of the 'ecology of pinewood beetles'. He lists 129 species of Coleoptera associated with pine, of 2 Broadleayed woodlands which 4 are of doubtful status. Of the remainder, 105 In Scotland, broadleaved woodlands are typical of the are known to occur in Scotland, and 44 of these have a central lowlands and the coastal fringe, especially on geographic range lying mainly within the relict Caledo- the west coast. Crowson (1962, 1964) made a nian forests. Hunter mentions an additional 6 species particular study of the Coleoptera of oakwoods (Quer- which occur both in pine and in association with other cus spp.) within easy reach from Glasgow, and trees. Although relying heavily on published records, recorded a number of species with a very restricted much of this account is based on his own experience 96 in the field. Not only is this the most comprehensive ing plantations of P. sylvestris)' (cf Welch 1974, 1981). record of any group of pine insects in Britain, but it is He further remarks that, although the saproxylic rare among published accounts in distinguishing the species associated with European pines are well Scottish element of the fauna. Speight (1985) argues represented in Britain, those species 'exclusive to one the absence of Caledonian pine insects in Ireland as or another of the introduced genera of conifers are evidence for the extinction there of Scots pine (Pinus generally lacking'. The inference is that colonization by sylvestris), and for its present fauna having been those insects which feed on the foliage and shoots is acquired since its reintroduction by man. much more rapid than colonization by species feeding on the woody tissues and associated saprophytic The of the Cairngorms NNR were sur- fungi. This hypothesis is supported by the fact that veyed by Mac Alpine (1979a, b) and most of his most of the 'Urwaldtiere', or ancient forest indicator trapping sites were in pine forest. His published lists species, fall into this latter group. One would expect, contain 393 species but, unfortunately, for most he therefore, that the colonization of introduced conifers provides only locality and date of collection. For only 4 will be by the true phytophages. The primary hindrance species is pine mentioned as the host plant, although to the accumulation of such data is the reluctance, or the knowledgeable reader will recognize a number of inability, of most entomologists to record the host tree other species associated with pine, and a few known beyond the generic level. This reluctance is true for to feed on species of spruce (Picea spp.). Many other both coniferous and broadleaved species, but is lists of Lepidoptera have been published for Scottish particularly true for introduced conifers. Not surpris- counties or localities but, often for brevity, data on ingly, such information as does exist stems almost habitat or host plant are lacking. Such lists usually entirely from publications by FC staff, not least presume that the reader possesses that information. It because they can identify the tree species concerned. will always be the case that the 'expert' can read more into a simple list than the novice, which is no help to Crooke (1957) listed 9 species of sawfly feeding on the latter. spruce and 7 on larch. He noted that, like their hosts, all were introductions into Britain. Furthermore, It can be seen that even for the 2 best studied insect Japanese larch (Larix leptolepis) and hybrid larch (L. groups, the Lepidoptera and Coleoptera, data are poor. eurolepis) appeared to be more attractive to the larch Information on the other Orders is even more sparse. sawfies than European larch (L. decidua), and Neodip- Crooke (1957) reviewed 27 species of conifer-feeding rion sertifer readily transferred from its usual host, sawflies (Hymenoptera: Symphyta), and, in recent Scots pine, to lodgepole pine. When present in high years, Liston has published on Scottish species but, densities, this insect was also capable of feeding on although several are pests of conifers, very few of his Sitka spruce. papers concern this element of the fauna (Liston 1981). Among a list of 66 conifer forest insects, Winter (1983) provides data for several species of Speight (1985) included 35 species of Symphyta. Of lepidopterous larvae feeding on both native and these, the larvae of 4 species of wood wasp in the introduced conifers. Under pine, he distinguishes family Siricidae are recorded feeding on a wide range Scots pine, lodgepole pine, Corsican pine, (P. nigra) .of tree hosts. Eleven species in the families Diprion- and Monterey pine (P. radiata), whilst under spruce he idae, Pamphilidae and Xyelidae have foliage-feeding includes Norway and Sitka spruce, together with 2 larvae which prefer pine as their plant host. A further records for oriental spruce (P. orientalis) and one for 11 species, almost exclusively Tenthredinidae, feed on Serbian spruce (P. omorika). In an FC Booklet, pres- spruce, but 9 of these also feed on fir (Abies spp.). ently in preparation, Bevan gives some information on Eight species are recorded from larch (Larix spp.). the preferred or alternative hosts of a number of insect Published accounts of other groups such as Homop- species. However, it is among the Homoptera, with tera and Diptera are few and far between. However, in their more intimate relationship with their host plant, his list of conifer insects, Speight includes 20 species that specific differences are best observed. Carter of hoverfly (Diptera: Syrphidae), 3 of which have (1983) provides a useful summary of the occurrence of saproxylic larvae, whilst the remainder feed on aphids 13 adelgid and 35 aphid species on 18 species of and related plant bugs. Eight species are listed conifer in Britain. He comments that only one adelgid exclusively from pine, but such predators generally is native to Britain, and among the aphids it is probable have less specific relationships with their host plants. that only 4 of the 22 species of Cinara (Lachnidae) are native. Parthogenetic, winged, female aphids are 3.1 Colonization of conifers by native and introduced insects among the first phytophagous insects to colonize Hunter (1977) comments that 'it would be difficult to exotic tree species. Most have spread to Britain from assemble separate British lists of beetles for species Europe, although 2 polyphagous species, one the of Pinus, and it is doubtful if such lists would differ'. ubiquitous bean aphid (Aphis fabae), now feed on However, Speight (1985) considers that 'in Great young seedling conifers throughout Britain. Adelgids Britain it is possible to differentiate between the are more catholic in their choice of host. Those of faunas of indigenous P. sylvestris.forest ("Caledonian" European origin may form galls on Asiatic and Am- pine forest) and commercial Pinus plantations (includ- erican spruces, but adelgids introduced from North Plate 1. Cultivation of deep peat moorland by double mouldboard plough at Naver Forest, Sutherland (Photograph Forestry Commission)

Plate 2. Shrub layer of juniper and well-developed ground layer of tussocky heather and bilberry in an opening of the old, native pine forest of Glen Tanar (Photograph N Picozzi) Plate 3. Fifth instar pine beauty larva (Photograph R Parks)

Plate 4. Defoliation caused by pine beauty moth larvae at Rimsdale, near Tongue, Sutherland (Photograph R Parks) Plate 5. Birchwood at Dunbeath, Caithness. Stable core on cliff, with some expansion over old pasture (Photograph G F Peterken)

Plate 6. Rassel Ashwood showing the dramatic regeneration within an enclosure. c1973 (Photograph G F Peterken) Plate 7. Management of native pinewoods must be carefully planned to benefit wildlife such as capercaillie and these golden eagles (Photograph N Picozzi)

.w • Plate 8. A family group of red deer in pre-thicket Sitka spruce. Red deer are now resident, often at high densities, in most forestry plantations in Scotland (Photograph M D C Hinge GROUPSOF LAND CLASSES

n

50 co°

Plate 9. Map showing the distribution in the UK of 32 land classes characterized by Bunce et al. (1982) (source: Bunce & Last 1981) Al B1

A2 B2 Plate 10. Fruitbodies ano mycorrhlzas formed by species of Laccaria (early state).and Leccinum (late-stage) with birch (Al Laccona proxirna and A2 Laccaria-type; B1 Leccinum roseofracta and B2 Leccinum-type) (Photograph Al A2, A Crossley: il31 02, V Flerno) 97

America almost exclusively gall only American species 8-toothed spruce bark beetle (Ips typographus). of spruce. Although it has been recorded regularly since 1849, breeding in Britain is not proven (Winter 1985). Winter Perhaps the most interesting tree colonists are those (1983) also lists some Scolytidae and Cerambycidae insects which have previously not had a conifer as a known only as non-breeding introductions in spruce host plant. As early as 1933, larvae of the bilberry and pine. tortrix ( viburnana) and the narrow-winged pug (Eupethecia nanata) had been recorded severely Many species of Lepidoptera are widespread and of damaging young plantations of various conifers after common occurrence in low numbers on pine in exhausting their normal food plants, bilberry (Vaccin- Britain, but only rarely reach pest proportions. How- ium), heath (Erica) and heather (Calluna vulgaris) in ever, lodgepole pine has proved to be unusually Cornwall and parts of Wales (Crystal 1937). Styles susceptible to attack by species such as the pine shoot (1959, 1961) was the first to compile lists of Lepidop- moth (Rhyacionia buoliana), larch bud Moth (Zeiraphera tera reared on unusual food plants based on FC field diniana), pine looper (Bupalus piniaria), and the pine survey notes and record cards. As more and more beauty moth (Panolis flammea). An intensive study of previously treeless upland areas were being planted, the pine beauty moth has revealed differences in larval mainly with introduced conifers, so such unusual behaviour from that reported for this species on the records increased. Several species of Lepidoptera continent (Bevan 1986). This is quite a common, appear to have found the young succulent shoots of although not well-documented, phenomenon. Many these conifers a readily acceptable alternative to the species of phytophagous insect may behave different- grasses, heather and other moorland plants on which ly on the same host plants in Britain and on the they normally fed. Between 1968 and 1970, Winter continent. Even pest species, which have been the (1974) investigated the causes of post-planting losses subjects of prolonged research in continental Europe, in recently afforested areas, including 18 forests in may behave quite differently in Britain. Hunter (1977) Scotland. He provided the first records for 24 species points out that 77 of the species of beetle which he of lepidopterous larvae feeding on conifers, of which lists for Scots pine are considered in Finland to be Sitka spruce was the most common new host. Some associated with spruce. In 17 of these species, this woodland species such as the winter moth (Operoph- association, is thought to be especially close. He tera brumata) and dotted border (Agriopis marginaria) mentions 6 pine species which have been found on were being increasingly found adapting to moorland spruce in Britain and states 'no doubt many others are conditions, feeding on heather (Wormell 1977). These capable of the same transition'. events were unusual enough in that both species have flightless females and rely for dispersal upon aeronaut- 3.2 Numbers of insect species on 4 conifer genera ing first instar larvae. However, once established in Southwood (1961) was the first person to assemble such areas, remote from deciduous woodland, the data concerning the number of insect species associ- winter moth \proved capable of a further change of ated with their host trees in Britain. Insect data were food plant and became a pest of young Sitka spruce. taken from only 3 Orders, Hemiptera, Lepidoptera and More recently in south-east Scotland, and in the Coleoptera. In addition to the native Scots pine and absence of heather and bog myrtle (Myrica gale), European larch, an introduction since 1629, South- winter moth is behaving as a primary pest of plant- wood included Norway spruce (Picea abies) and ations 8-12 years old (Stoakley 1985). Early plantings species of fir (Abies spp. (alba)) on the strength of of larch on areas of purple moor-grass (Molinia there having been native trees in Britain before the last caerulea) have suffered ring-barking by the clay- glacial phase, noting their reintroduction in the 16th coloured weevil (Otiorhynchus singularis). and 17th centuries respectively. However, in updating this information (Kennedy & Southwood 1984), he Foresters have always kept a watchful eye on the retains spruce but inexplicably drops fir; otherwise, dangers of introduced pests. The large larch bark the same tree hosts are used but coverage of insect beetle (Ips cembrae), Which was first recorded on larch orders is greatly improved, with the resulting increase in northern Scotland in 1955, has since spread south in numbers of associated insect species. as far as Peebles and has been found on windblown Sitka spruce. Another long-expected arrival from nor- Exactly what comprises an 'associated insect' is still a thern Europe was the great spruce bark beetle matter for considerable discussion, and even personal (Dendroctonus micans), albeit at present confined to interpretation. In Table 1, I have attempted to compile England and Wales. Although only discovered in 1982, lists of the numbers of insects attributed to 4 conifer by counting the annual rings of the tissues occluded genera by various authors. Comparison of the 2 lists by following attack, it was clear that this beetle had been Southwood and Kennedy shows how the data have in the UK since about 1973 (Bevan & King 1983). been improved since the much-quoted 1961 paper. In Bevan (1986), aware of such dangers, regards the each case, the records are referable to a single host white pine weevil (Pissodes strobi) and 3 species each species. I have included Hunter's comprehensive of the scolytids Dendroctonus and Ips as having a 'high Coleoptera list for Scots pine for comparison, and the quarantine hazard risk'. One of these species is the list prepared for Carlisle and Brown (1968) by 0 W 98

Table 1 Numbers of insect and mite species associated with 4 genera of conifers in Britain

Host Pinus Larix

LO oo Reference* • oo a) (author's initials co ro & date) 0 cT_ Acarina 0 1 Heteroptera 9 3 Homoptera 14 6 Thysanoptera 1 0 Lepidoptera 22 16 Diptera 3 1 Hymenoptera 10 5 Coleoptera 11 6 Total 70 38

*DB 1985 = D Bevan's MS for Forestry Commission Booklet PIDB = ITE's phytophagous insect data bank (see text)

Richards. When I compared the lists of Styles (1959, account so far abstracted (and possibly the only such 1961) with that of Winter (1974, 1983), I found reference) is from a continental reference. Table 1 numerous inconsistencies. In his 1983 FC Booklet, provides the non-specialist with some indication of the Winter includes only some, but not all, of Styles' range and accessibility of data on insects and their tree records and also omits some from his own earlier hosts. All numbers given in Table 1 relate to Britain as publication. I have, therefore, combined all these a whole, and many species included may not occur in records into a single column entry for all 4 tree hosts. I Scotland. have taken the liberty of abstracting details from Bevan's forthcoming Booklet as it is hoped that this 4 Conclusions and recommendations will become widely used by practising foresters when I hope that I have demonstrated the scattered nature available. His selection of insect species is based not of the available published data on woodland and forest only on those known to feed directly upon their host, insects. Much more information may exist in an but upon those species for which the Entomology unpublished form, so that the combined personal Branch of the FC had reCeived most enquiries over the knowledge of various specialist entomologists, forest- years. It is, therefore, biased towards pests and the ers and ecologists will vastly exceed this summary. In more obvious species. This and the remaining lists my opinion, the root of the problem lies in our lack of include the combined records of insects from a knowledge of the ecology of introduced tree species. number of different tree species within each genus. This is true for broadleaved as well as coniferous For example, of the 94 insect species listed from species. There appear to be 3 major areas where more spruce by Winter (1983), 19 were recorded from information is urgently needed and where future work Norway spruce and 29 from Sitka spruce. Among could be directed. These are as follows. those from non-specific spruces are 5 Coleoptera not yet known to breed in Britain. Similarly, a further 3 are 4.1 Mapping introduced trees included in his list for pines. I should remind you that Most county floras and plant mapping schemes ignore Bevan's list also includes 5 spruce and 2 pine insects introduced trees, although occasionally they are in- of similar status. cluded for counties where they regenerate naturally. We need to know more about their distribution in The last columns for each tree genus in Table 1 contain Britain. Jeffers (1972) provides some useful 10 km data from the Institute of Terrestrial Ecology's Phy- distribution maps for Scots pine, Corsican pine, lodge- tophagous Insect Data Bank (PIDB).The data included pole pine, Norway spruce and Douglas fir, but Sitka here are a very small part of the mammoth undertaking spruce is notable by its absence. If it has not already. by Dr Lena K Ward and D F Spalding to abstract been done, a start could be made by bringing together important published literature containing references to planting records for this (and other) species from all the relationships between insects in Britain and their the FC Conservancies. Hopefully, this compilation recorded plant hosts. Many of the records aie taken could include not only plantings on land owned by the from European sources which must be treated with FC, but also for grant-aided schemes in private caution when considering the British fauna. However, woodlands. This information, in turn, could possibly be in many cases, although an insect may be well known supplemented by data from the Kew Index of Living from a particular host in Britain, the only published Collections, which already contains the computerized 99 records of trees at Kew, Wakehurst Place, and 53 "attention is also paid to the increasing ability of certain National Trust properties. Once such base maps are insects to transfer to conifers from unrelated native available, there will be an incentive for others to add to host plants. them. Data on the numbers of insect species associated with 4.2 Recording insects on introduced trees 4 conifer genera, Pinus, Picea, Abies and Larix, are There is nothing better guaranteed to promote a flow presented, combining records from various sources. of information than the production of a 'definitive' list Caution is stressed when continental insect host data of insect species. We should consider publishing lists are used. of those insects which are currently known to occur on such trees as Sitka spruce. We should not then just sit Recommendations are made for the distributional back and await the anticipated flood of 'additional' mapping of introduced trees in Britain, and the records, but should consider implementing an active recording of the insects associated with them. It is survey programme in order to understand better which also suggested that the movement of insects between insects are able to utilize these vast new man-made Caledonian pinewood relicts and conifer plantations be habitats, and to encourage other people to do likewise. investigated.

4.3 Colonization from Caledonian pinewoods into plantations 6 Acknowledgements Many of the native pinewood areas identified by I am deeply grateful to my colleagues Dr Lena K Ward Steven and Carlisle (1959) were small relict pop- and D F Spalding for providing me with the available ulations which had survived at the heads of glens or in data in their Phytophagous Insect Data Bank, and for locations which became isolated by adjacent felling their time and assistance in ensuring that I understood and forest clearance. As a result, many of the rare the resulting coded printouts. It is a pleasure also to insects associated with them have survived in isolated thank D Bevan for access to themanuscript of his pockets throughout the Highlands. Increasingly, these Forestry Commission Booklet. areas have become caught up in the approaching legions of commercial plantations so that they may 7 References now be contiguous with, or relatively close to, young Bevan, D. 1986. Forest insects. (Forestry Commission booklet.) conifers of various species. I can only reiterate London: HMSO. In press. Hunter's (1977) comment that 'the ability of relict Bevan, D. & King, C.J. 1983. Dendroctonus micans Kug. - a new pinewood species to colonise plantations has (still) not pest of spruce in UK. Commonw. Forest. Rev., 62, 41-51. • yet received sufficient attention'. He also saw the Carlisle, A. & Brown, A.H.F. 1968. Pinus sylvestris L. (Biolc4cal need for monitoring the development of the beetle flora of the British Isles, no. 109.) J. Ecol., 56, 269-307. fauna in a number of new pine plantations within the Carter, C.l. 1983. Some new aphid arrivals to Britain's forests. Proc. Caledonian forest area. Ten years have passed since Trans. Br. ent. nat. Hist. Soc., 16, 81-87. that symposium, and the plea for more detailed Carter, C.l. & Nichols, J.F.A. 1985. Host plant susceptibility and information in this field is as real today as it was then. choice by conifer aphids. In: Site characteristics and population dynamics of lepidoteran and hymenopteran forest pests, edited by D. Bevan & J.T. Stoakley, 94-99. (Forestry Commission research 5 Summary and development paper no. 135.) London: HMSO. A brief background is given to our present state of Crooke, M. 1957. A brief review of the British conifer feeding knowledge of forest entomology in Scotland, which sawfl i es. Z. angew. Entomol., 41, 179-183. explains how entomologists have largely ignored Crowson, R.A. 1962. Observations on Coleoptera in Scottish oak recent conifer plantations in favour of relict 'Caled- woods. Glasg. Nat., 18, 177-195. onian' pinewoods and broadleaved woodlands. The Crowson, R.A. 1964. Additional records of Coleoptera from Scottish insect fauna of the latter is regarded as being oak woods. Glasg. Nat., 18, 371-375. essentially a diluted southern fauna with some add- Crystal, R.N. 1937. Insects of the British woodlands. L6ndon: itional northern elements. Warne. Davies, E.J.M. 1985. Forestry in Scotland. Edinburgh: Forestry Some examples are given of the types of insect Commission. species lists prepared by individuals and teams of Forestry Commission. 1985. Forestry facts apd figures 1984-85. entomologists. The acceptability • of such lists for Edinburgh: FC. publication and the interpretation of their contents are Hunter, F.A. 1977. Ecology of pinewood beetles. In: Native pinewoods of Scotland, edited by R.G.H. Bunce & J.N.R. Jeffers, discussed. 42-55. Cambridge: Institute of Terrestrial Ecology. Jeffers, J.N.R. 1972. Conifers and nature conservation. In: Conifers Introduced conifers are typically colonized by insects of the British Isles, 59-73. London: Royal Horticultural Society. which have either transferred from a native tree, or Kennedy, C.E.J. & Southwood, T.R.E. 1984. The numbers of have themselves become established by accidental species of insects associated with British trees: a re-analysis. J. introduction or natural immigration. Some immigrant Anim. Ecol., 53, 455-478. pests from continental Europe and North America are Liston, A.D. 1981. Notes on little known British Pristophora Lat. considered, and the susceptibility of certain introduced (Hym., Tenthredinidae) of the abietina group. Entomologist's mon. tree hosts to native insects is discussed. Particular Mag., 117, 73-75. 1 00

MacAlpine, E.A.M. 1979a. The Lepidoptera of the Cairngorms Welch, R.C. 1981. The modern story - insects. In: The Cairngorms, National Nature Reserve. Entomologist's Rec. J. Var., 91, 1-6, their natural history and scenery, edited by D. Nethersole-Thompson 65-70,213-216,241-244. & A. Watson, 281-283.2nd ed. Perth: Melvin Press. MacAlpine, E.A.M. 1979b. The Macrolepidoptera of Inverness-shire Winter, T.G. 1979. New host plant records of Lepidoptera associ- - Newtonmore District, Supplement 10. Entomologist's Rec. J. Var., ated with conifer afforestation in Britain. Entomologist's Gaz., 25, 91,157. 247-258. Southwood, T.R.E. 1961. The number of species of insect Winter, T.G. 1983. A catalogue of phytophagous insects and mites associated with various trees. J. Anim. Ecol., 30, 1-8. on trees in Great Britain. (Forestry Commission booklet no. 53.) Speight, M.C.D. 1985. The extinction of indigenous Pinus sylvestris Edinburgh: Forestry Commission. in Ireland: relevant faunal data. Ir. Nat. J., 21, 449-453. Winter, T.G. 1985. Is Ips typographus (Linnaeus) (Coleoptera: Steel, W.O. & Woodroffe, G.E. 1969. The entomology of the Isle of Scolytidae) a British insect? Entomologist's Gaz., 36, 153-160. Rhum National Nature Reserve. Trans. Soc. Br. Ent., 18, 91-167. Woodroffe, G.E. 1974. Hemiptera from Rhum, the Cairngorms, Steven, H.M. & Carlisle, A. 1959. The native pinewoods of Deeside and lnvernaver. Glasg. Nat., 19, 101-114. Scotland. Edinburgh: Oliver & Boyd. Wormell, P. 1977. Woodland insect population changes on the Isle Stoakley, J.T. 1985. Outbreaks of winter moth, Operophtera of Rhum in relation to forest history and woodland restoration. Scott. brumata L. (Lep., Geometridae) in young plantations of Sitka spruce For., 31, 13-36. in Scotland. Z. angew. Entomol., 99, 153-160. Wormell, P. 1982. The entomology of the Isle of Rhum National Styles, J.H. 1959. Notes on some Microlepidoptera. Entomologist's Nature Reserve. Biol. J. Linn. Soc., 18, 291-401. Gaz., 10, 43-44. Young, M.R. 1986. The effects of commercial forestry on woodland Styles, J.H. 1961. Additions to the list of foodplants of Lepidoptera Lepidoptera. In: Trees and wildlife in the Scottish uplands, edited by larvae, not recorded in 'Larval food plants' by P.B.M. Allan, and 'The D. Jenkins, 88-94. (ITE symposium no. 17.) Abbots Ripton: Institute caterpillars of the British ' by W.J. Stokoe. Entomologist, 94, of Terrestrial Ecology. 86-88. Welch, R.C. 1974. Insects and other invertebrates. In: The Cairngorms, their natural history and scenery, edited by D. Nethersole-Thompson & A. Watson, Appendix 2,257-285. London: Collins.