ThQ Glasgow NataraHst Volume 24 Part 1 2002 Journal of THE GLASGOW NATURAL HISTORY SOCIETY GLASGOW NATURAL HISTORY SOCIETY (formerly The Andersonian Naturalists of Glasgow) The object of the Society is the encouragement of the study of natural history in all its branches, by meeting for reading and discussing papers and exhibiting specimens and by excursions for field work. The Glasgow Natural History Society meets at least once a month except during July and August, in the University of Glasgow, the Glasgow Art Gallery and Museum, or Hillhead Library. The present rates of subscription per annum are: for Ordinary Members, £17: Family Members, £3 extra; Junior Members (under 21) £8; School Members, £1 . Payment by Direct Debit is encouraged. Further information regarding the Society's activities and membership application forms are obtainable from the General Secretary, c/o Natural History Department, Museum & Art Gallery, Kelvingrove, Glasgow G3 SAG. The Glasgow Natural History Society website is http://tftw.freeuk.com/gnhs/ , which also contains details of the Glasgow Naturalist. The Glasgow Naturalist The Glasgow Naturalist is published by the Glasgow Natural History Society. ISSN 0373-241X. Price £14.00 plus £1.50 p.&p. UK; £3.00 overseas. The Glasgow Naturalist is free to members. The Glasgow Naturalist is edited by Azra Meadows and Peter Meadows, Graham Kerr Building, University of Glasgow G12 8QQ (Tel: 0141 330 6622, Email: gbza3 1 @udcf.gla.ac.uk) with the help of J.R. Downie R.H. Dobson, A. McG. Stirling, and T.N. Tait. Contributions are invited, especially when they bear on the natural history of Scotland. Full details of how to contribute articles or short notes are given at the end of the volume. A limited number of advertisements can be accepted and enquiries should be sent to the Editors. This publication is included in the abstracting and indexing coverage of the Bioscience Information Service of Biological Abstracts and the Botanical Society of the British Isles Abstracts. The following back numbers are available for purchase in their separate parts: Vols.II - VII (1890-1918); Vols. XIII - XXIII (1937-1999). Of the earlier Journals the only parts available are: Proceedings and Transactions of the Natural History Society of Glasgow Vol. II pt. 2; Vol. VI pt. 1; Vol. Vll.pt. 3; Vol. VIILpts. 1 & 2. Enquiries regarding prices of and orders for any of the above, or for reprints or photocopies, should be addressed to the Librarian: Mrs Joan Chapman, 121 Randolph Road, Jordanhill, Glasgow G1 1 7DS. Publications of Glasgow Natural History Society Alien Species: friends or foes? Edited by J.R.Downie (2001). Proceedings of the GNHS ISO"" Anniversary Conference. Price £10.00 plus p & p £1.50. Bound copies of the following may be obtained from the Librarian at the address above and at the prices shown: The Flora of the Clyde Area (Original printing). J.R. LEE, Price £1 1.00 to members of GNHS and to the book trade; £13.50 to others (p. & p. £1 .00 extra). This is still the only work of its type and is in diminishing supply. A few unbound copies are available: £5 (p.&p. £1 extra). The Flora of Ailsa Craig. B. ZONFRILLO, 1994. Price £2.50 plus p.& p.50p. The Natural History of the Muck Islands, N. Ebudes: 1. Introduction and Vegetation with a List of Vascular Plants. R.H. DOBSON & R.M. DOBSON, 1985. Price £1.00 plus p. & p. 50p. 3. -Seabirds and Wildfowl. R.H. DOBSON & R.M. DOBSON, 1986. Price £1.00 plus p. & p. 50p. 5. Landbjrds: R.H, -E^BSON, 1988. Price £1 .00 plus p. & p. 50p. The follow'ing bound reprints from^he Glasgow Naturalist may be obtained from the librarian at the above address and at the prices shown. ; Additions to the Flora of the Clyde Atea. John R. Lee (1953). £1 (p.&p. 50p). Archives The Society's archives are in the hands of the City Archivist, Mr Andrew Jackson, and stored at the Mitchell Library, Ref. No. TD 1408. To access, telephone Mr Jackson 0141-287-2907 or Robert Urquhart 287-2914. The Glasgow Naturalist Volume 24 Part 1 2002 CONTENTS PRESIDENTIAL ADDRESS. Long-term natural history: some aspects of the environment and living things. 1-14 R.S.K.Gray EDITORIAL Climate change. Its history and future in relation to Scotland’s landscape, people and economy. 15-22 Azra Meadows and Peter Meadows FULL PAPERS Community structure and biometrics in microhabitats within mussel beds (Mytilus edulis) from 23 - 28 intertidal environments at Ardmore Bay, Clyde Estuary, Scotland. Noelia Carrasco, Azra Meadows & Peter Meadows. Scottish insect records for 2001 . 29-33 E. G. Hancock The natural history of the Glasgow Botanic Gardens. Plants growing in a wild state 1998-2001. 35 - 38 P. Macpherson Cotoneasters 1982-2001 in Lanarkshire. 39-41 P. Macpherson and E.L.S. Lindsay Barbel barbus barbus in the River Clyde: a new fish species established in Scotland. 43 - 46 Peter S. Maitland and William W. Miller Nest-site competition with blue tits and great tits as a possible cause of declines in willow tit 47 - 50 numbers: observations in the Clyde area. James Maxwell The Scottish Lynx: Is reintroduction a possibility? 51-58 Katie McDonald A former water-meadow in the upper Carron Valley, Stirlingshire. 59 - 63 John Mitchell Loeh Lomondside depicted and described 5. Early natural historians. 65 - 68 John Mitchell Environmental impact on the sea bed caused by trawling for the Norway Lobster, Nephrops 69 - 82 Norvegicus in the Clyde Sea area. A geotechnical assessment. Azra Meadows, Peter S. Meadows and John M. H. Murray A new species of parasitic copepod, Sphaeronella keppelensis n.sp. (Siphonostomatoida: 83-91 Nicothoidae), from the amphipod Orchomene nanus (Kroyer, 1846) in the Firth of Clyde, Scotland. Myles O’Reilly I SHORT NOTES Compiled by A. McG. Stirling Pill millipede Glomeris marginata on Jura. 93 Glyn M. Collis and V. Dawn Collis Platyartrus hoffmanseggi in Kirkcudbright (VC 73). 93 V. Dawn Collis and Glyn M. Collis Lime tree {Tilia spp.) regeneration 200 1 . 93 R.K.S. Gray Orange ladybird Halyzia 16-guttata in Hyndland. 94 Norman R. Grist Summertime swifts. 94 Norman R. Grist Urban foxes in Hyndland, Glasgow. 95 Norman R. Grist A rich botanical site at Leadhills. 96 P. Macpherson Lesser hairy-brome at the Falls of Clyde. 96 P. & L.M.D Macpherson and J. Waddell Predator/prey relationships in an urban environment. 97 Azra Meadows, Peter S. Meadows and Willie On- Great Spotted Woodpeckers feeding on the nectar of Red-Hot Pokers. 98 John Mitchell Haeckelian radiolarian material and the Microscopical Society of Glasgow. 98 - 101 P. Geoffrey Moore and Richard Sutcliffe Lily beetle {Lilioceris lilii), in Glasgow. 101 Richard Sutcliffe Rabbit Calicivirus Disease on Ailsa Craig, Ayrshire in 2003 102 - 103 B Zonfrillo, H. Thomson & R A Stewart BOOK REVIEWS Compiled by Ruth M. Dobson (reviewers names in brackets) 105 - 117 A Guide to Bird Watching in the Clyde Area (2001) Ed. by Cliff Baister and Marion Osier. (Ian C. McCallum) Amphibians and Reptiles: A Natural History of the British Herpetofauna (2000) by Trevor J.C. Beebee and Richard A. Griffiths. (Roger Downie) Minding Animals: Awareness, Emotions and Heart (2002) by Marc Bekoff. (Felicity Huntingford) Bird Migration: General Survey (2001) by Peter Berthold. (David Houston) Collins Field Guide: Caterpillars of Britain and Europe (2001) by David J. Carter, illustrated by Brian Hargreaves. (Ronald M. Dobson) Global Warming, The Science of Climate Change (2000) by Frances Drake. (Peter Meadows and Azra Meadows) The New Encyclopedia of Reptiles and Amphibians (2002) Ed. by Tim Halliday and Kraig Adler (Roger Downie) Wetland Ecology: Principles and Conservation (2000) by Paul Keddy (Colin Adams) Fossils and Evolution (1999) by T.S Kemp (Alastair Gunning) Coral Reef Fishes of the Indo-Pacific and Caribbean (2001) by Ewald Lieske and Robert Myers (Rupert Ormond) Moths (2002) by Michael E,N. Majerus (E.G. Hancock) The Shielding and Drove Ways of Loch Lomondside (2000) by John Mitchell (Ruth H. Dobson) Trees of Britain and Northern Europe (2001) by Alan Mitchell, illustrated by John Wilkinson (Ruth H. Dobson) The Broads (2001) by Brian Moss (John Mitchell) Collins Bird Guide (2000) by Killian Mullarney, Lars Svensson, Dan ZetterstrOm, Peter J. Grant (Bernard Zonfrillo) Small Freshwater Creatures (2001) by Olsen, Sunesen and Pedersen (Ian C. McCallum) Small Woodland Creatures (2001) by Olsen, Sunesen and Pedersen (Ian C. McCallum) Animal Tracks and Signs (2001) by Bang Preben and Dahlstrom Preben (Margaret M.H.Lyth) Conservation Biology (2002) by Andrew Pullin (Roger Downie) Colour Identification Guide to the Grasses, Sedges, Rushes and Ferns of the British Isles and North Western Europe (2000) by Francis Rose (Peter MacPherson) Trees: their Natural History (2000) by Peter Thomas (Bob Gray) Photographic Guide to the Butterflies of Britain and Europe by Tom Tolman (2001) (Richard Sutcliffe) The Variety of Life (2002) by Colin Tudge (Ronald M. Dobson) Seabird Numbers and Breeding Success in Britain and Ireland, 1999 (2000) by A.J. Upton, G. Pickerell and M. Heubeck (Bernard Zonfrillo) An Essential Guide to Bird Photography (2001) by Steve Young (T. Norman Tait) OBITUARY 119 Dick Hunter (1906 - 2002) By Ian C. McCallum and James H. Dickson Proceedings 2001 Officers and Council, Session MMI 2001 Advice to Contributors III Glasgow Naturalist 2002. Volume 24. Part 1. Pages 1-14. Presidential Address, 26"' February, 2002 LONG-TERM NATURAL HISTORY SOME ASPECTS OF THE ENVIRONMENT AND LIVING THINGS R.Gray President Glasgow Natural History Society 6, Prince Albert Road Glasgow G12 9JX Our new president, Roger Downie, in his earlier guise as Editor of the most recent issue of ‘The Glasgow Naturalist’ commented in his editorial that the Society rarely engages with major modern advances in the biological sciences and he posed the question: ‘Is our Society taking too narrow a view of what natural history is?’ Whilst not proposing to answer this question in full I propose to take a brief look at some controversial aspects of the subject that draws us all to this Society. Natural history has been defined as the study of natural phenomena including inanimate phenomena, such as rocks, soils and climate, but commonly confined to living things, animals and plants in the wild. (Fitter, 1967). Ecology is defined as the study of the relationship between plants and animals and their environment. (Haeckel, biologist and philosopher, 1866). I should like to look at the connection between these two in the light of present day knowledge. This means that I shall look at the inter- relationships between the environment, plants (mainly trees) and animals (mainly birds and mammals) with particular but not exclusive reference to a Scottish context. My interest in atmospheric CO2 concentrations goes back to science lab experiments carried out to compare the O2, C02and N2 volumes of inhaled air with exhaled air. The CO2 content of inhaled air, 370 ppm, is so small as to be immeasurable in the context of an ordinary laboratory. There is some 40 times more water vapour in the air than CO2 and it is a matter of common observation that cloudy evenings are warmer than clear evenings. So simple logic suggests that small variations in atmospheric CO2 concentration will have little effect on global temperature. This is very much at odds with what we come across in the media. EARTH’S CLIMATIC HISTORY: THE LAST 1,000,000 YEARS Some 10 major ice ages have occurred over the past 1 million years (Figure 1). They recur at approximately 100,000 year intervals, persist for about 90,000 years, after which they have been followed by approximately 10,000 year interglacials. This periodicity has been attributed to the Croll - Milankovitch cycle (Figure 2), whereby regular changes in the distance of the earth from the sun affect the amount of solar radiation reaching the earth’s surface (Farrow, 2001). The glacial epoch about which we know the most is the most recent one which was at its peak about 20,000 years ago. Land plants suffered as the air’s CO2 content fell to about 180 ppm. This fall was caused by: a) the increased ability of colder water to hold more dissolved CO2 and b) larger growth rates of phytoplankton caused by the amount of iron rich dust carried by the stronger winds of the period. Had the CO2 concentration dropped much lower it is likely that several plant extinctions would have occurred, since many plants find it difficult to survive at CO2 concentrations of the order of 50 to 100 ppm (Idso, 1989; Salisbury and Ross, 1978). Large and rapid shifts in climate have been detected in Greenland and Antarctica from deep sediment cores, ice cores, lake sediments and pollen series, e.g. in Greenland rapid warming of some 7°C in a few decades was observed about 1 1 ,500 years ago (Dansgaard et al., 1989; Johnsen et al., 1992; Grootes et al., 1993). Rapid warming, followed by periods of slower cooling and then rapid freezing are typical of interstadial events (as well as interglacials), of which about 20 occurred during the last glacial period. They lasted between 500 and 2000 years (Dansgaard et al., 1993). CO2 AND TEMPERATURE: ICE CORE CORRELATIONS (Eischer et al, 1999) Evidence from Antarctic ice cores through the last three ice ages showed that the earth warmed up well before there was any increase in the air’s CO2 content. The relationship between temperature and CO2 is just the reverse of what is assumed in all the climate model studies that warn of dramatic warming in response to the ongoing rise in the air’s CO2 content: temperature rises first, and then comes an increase in atmospheric CO2. NEARLY HALF A MILLION YEARS OF CLIMATE AND CO2 (Petit et al., 1999) Petit et al. (1999) showed from the Vostok ice core going back 420,000 years that the 4 interglacials preceding the present one (the Holocene) were warmer by an average temperature of > 2°C. Hence the current interglacial is by far the coolest of the five most recent such periods (Figure 1). Also “during glacial inceptio the CO 2 decrease lags the temperature decrease by several thousand years.”. Since the current interglacial is by far the longest stable warm period of the past 420,000 years we are probably overdue for the next ice age. AIR TEMPERATURE OF THE PAST DECADE The air temperature of the last decade of the 20th century in terms of typical interglacial temperatures is clearly unusually cool, even if the temperature is warmer than it has been over the past 100 years (Figure 1). The air’s CO2 concentration today 1 Global mean temperature change (“C) Figure 1. Temperature change vs Time solder ^1 I I I I 1000 750 500 250 0 Thousands of years ago Source: Budiansky, 1995 Hislarical Isotopic Temperature Record from the Vostok Ice Core +0.5 -0.5 I I I I 1880 1900 1940 1980 Year Source: Budiansky, 1995 Regular patterns appear only on time scales of 1 00,000 years or more. Top: Temperatures inferred from oxygen isotope fluctuations in ocean sediments. Mid: From oxygen isotopes in the Vostok ice core. Bottom: Global mean temperature from direct measurement. Figure 2. Croll-Milankovitch Cycle Earth's orbit varies in three ways: a), b) & c). These affect amount of radiation and so climate Source: Lamb & Sington, 1999 stands at nearly 370 ppm whereas in the 4 prior interglacials it never rose above 290 ppm. So the higher temperatures of previous interglacials cannot be attributed to either COj concentrations (they were lower) or to human interference (too few humans). So what is the most likely cause of climate change? A 1000 YEAR HISTORY OF SUNSPOT NUMBERS (Rigozo et al., 2001) Both the Mediaeval and Modem maxima in sunspot numbers and solar radiation output are far above all other periods of the past thousand years. On this basis current temperatures are expected to be higher than at any other time during the past millennium. Studies of recent data from the European Space Agency’s sun watching Soho satellite indicate a solar energy surge and a particularly big increase in UV light. This has coincided with a doubling in the strength of the sun’s magnetic field, which blocks cloud-forming cosmic rays. Fewer clouds would mean that more heat reaches the Earth’s surface, although it could be argued that warming of the Arctic Ocean has resulted in increased cloudiness at high northern latitudes. There is however no need to invoke variations in the air’s COj content as a cause of temperature variation. ICE SHEETS IN THE NORTHERN HEMISPHERE (Figure 3) When the ice of the last glaciation was at its greatest extent it reached south to the Chiltern hills, to the south and west of which was glacial outwash and tundra (Figure 4). The area south and west of Cornwall was dry land and may have been a refuge for trees such as the Strawberry tree {Arbutus unedo) which possibly migrated to southwest Ireland as the ice melted (Mitchell & Coombes, 1998). The ice of the last glaciation began to melt about 17,900 years ago (Huntley et al., 1997) when the exposed land was colonised firstly by open ground taxa. These were then replaced by a dwarf shrub community that included Juniperus, Salix and Betula nana and occasional tree birches (Ramsay & Dickson, 1997). By about 14,600 years ago the ice had almost disappeared completely but by 12,900 year ago a substantial ice sheet, the Loch Lomond readvance, centred on Rannoch Moor had accumulated. About 11,200 years ago extremely rapid climatic warming caused the ice to melt finally thus allowing dwarf shrubs to migrate again into the area behind the melting ice. These were gradually followed by the major tree taxa. After late-glacial times the first tree pioneers were the downy and silver birches {Betula pubescens and B. pendula) and Scots pine {Pinus sylvestris). The pollen record suggests that birch spread from the east across land now forming the bed of the North Sea. The genetic evidence is strong for the colonisation of Scotland by Scots pine by migration from north-central Europe mainly and from southern Europe (Ennos et al., 1997). The expansion of pine from sheltered pockets in north west Scotland where it may have survived the last glacial period remains a matter of debate. In Scotland the tree birches were followed soon by hazel (Figure 5) then elm. Oak tends to arrive either at the same time as elm or slightly later (Ramsay & Dickson, 1997). Pollen analysis cannot distinguish between the two native oaks but the present distribution, with the sessile oak predominant in the north and west and at higher altitudes, suggest that the sessile preceded the common oak, which is found mainly in lowland areas and more in the south and east (Mitchell & Coombes, 1998). Also early to arrive were aspen and rowan. The total complement of native trees was about 35, about 25 in Scotland (Table 1). Europe was unfortunate during the Ice Ages because the great mountain systems run from east to west. The flora and fauna were trapped between the northern ice and the mountain ice. The Mediterranean prevented access to Africa. In Canada and the northern USA, which experienced the same ice ages, the mountains run north to south. Trees migrated southwards along mountains or valleys at their preferred climate, and afterwards migrated back again. Hence there is an immense wealth of species, broadleaved and coniferous, in North America. The National Park of the Appalachians, for example, possesses 131 native trees whereas the European figure is 85. Britain has two species of oak; the USA has 80. Britain was recolonised by species hardy enough to have survived on the European plains and sufficiently fast moving to migrate back in the 6000 years before the Channel was created. ISOCHRONE CONTOUR MAPS (FIGURES 6A AND 6B) The maps show the history of the spread of some trees through these islands. During a warm early period the small-leaved lime {Tilia cordata) was dominant over large areas and the wych elm {Ulmus glabra) and bird cherry {Prunus padus) were other early arrivals. Ash {Fraxinus excelsior), field maple, yew and hawthorn arrived with time to spare, probably with wild cherry {Prunus avium) and crack willow. The land bridge was of chalk, and apart from holly and hornbeam, the last trees to cross it were those that thrive on chalk and are often now found wild only near chalk hills - the wild service tree, whitebeam, beech and, probably last, box. (Mitchell & Coombes, 1998.) BRITAIN’S TREES C.6500 YEARS AGO (Figure 7) There is little evidence for Scots pine being a major component of central Scottish woodlands (Dickson et al., 2000). It was however the principal tree in the Highlands with tree birches and hazel dominating the Outer Hebrides and Northern Isles. By the time of maximum woodland expansion low lying, mainland Scotland north to the Great Glen was in the zone of oak dominance, with wych elm, alder and ash. Soil conditions determined woodland composition. Small amounts of Scots pine grew on drier areas of peat bog. 3 Figure 3. Ice sheets in the northern hemisphere I I Ice sheets I I Sea ice Source: Lamb & Sington, 1998 Figure 4. European vegetation at last ice glaciation KEY — Glacial shore line Limit of land ice Limit of ice at maximum Remainder was covered by ice or tundra Source: Simms, 1990 Figure 5. Source; Dickson & Dickson, 2000 Drawings of pollen grains and spores frequently recognised in samples from British peats, lake muds and archaeological layers, a. Scots Pine, b. Tree Birch, c. Dwarf Birch, d. Hazel, e. Yew, f funiper, g. Wych Elm, h. Poaceae (Grasses), i. Oak, J. Ash, k. Alder, 1. Willow, m. Heather, n. Cyperaceae (Sedges and related plants), 0. Ribwort Plantain, p. Small-leaved Lime, q. Mugwort, r Dandelion-type, s. Pondweed, Hogweed- type, u. Common Rock- rose, V. Bogmoss, w. Lesser Clubmoss. 1 micron = one thousandth of a mm. From Pigott and Pigott (1959) I Table 1: British Native Tree Species Bold =15 species of natural woodlands. Non-bold = species found in non-woodland habitats. (except Arbutus, wild now in Ireland, not in Britain) *= not native in Scotland (Mitchell, 1974) Alder (Alnus glutinosa) Crab Apple {Malus sylvestris) Ash (Fraxinus excelsior) Aspen (Populus tremula) *Beech (Fagus sylvatica) Silver Birch (Betula pendula) Downy Birch (Betula pubescens) Blackthorn (Prunus spinosa) *Box (Buxus sempervirens) Bird Cherry (Prunus padus) Wild Cherry, Gean (Prunus avium) Wych Elm (Ulmus glabra) Hazel (Corylus avellana) Hawthorn (Crataegus monogyna) Holly (Ilex aquifolium) ^Hornbeam (Carpinus betulus) Juniper (Juniperus communis) *Small-leaved Lime (Tilia cordata) Broad-leaved Lime (Tilia platyphyllos) *Field Maple (Acer campestre) Pedunculate Oak (Quercus robur) Sessile Oak (Quercus petraea) *Wild Pear (Pyrus communis) Scots Pine (Pinus sylvestris) Black Poplar (Populus nigra) Rowan (Sorbus aucuparia) *Wild Service Tree (Sorbus torminalis) *Strawberry Tree (Arbutus unedo) *Whitebeam (Sorbus aria) Almond Willow (Salix triandra) (probably not native to Scotland, Stace, 1997) Bay Willow (Salix pentandra) Crack Willow (Salix fragilis) Goat Willow (Salix caprea) White Willow (Salix alba) Grey Sallow (Salix cinerea) Yew (Taxus baccata) Source: Miles, 1999 5 Figure 6a. Isochrone Contour Maps Spread of tree species Data from Prof. Birks of Bergen using radiocarbon dated pollen samples Birch (Betula pubescens and B. pendula) Pine (Pinus sylvestris) Hazel (Corylus avellana) Alder (Alnus glutinosa) Source: Milner, 1992 6 Figure 6b. Isochrone Contour Maps Arrows give direction of spread. Lines represent limit of spread at the date before present. Elm (Ulmus speaes) Lime (Tilia cordata and T. platyphyllos) Oak (Quercus species) Ash (Fagus sylvatica) Source: Milner, 1992 Figure 7. Woodland cover of Scotland several thousands of years ago, before woodland clearance had begun. Source: Dickson & Dickson, 2000 Within a thousand years of the maximum a dramatic reduction in elm occurred, primarily a result of disease but possibly enhanced by human activity. Pollen analyses from Lenzie Moss, Lochend Loch bog and Drumpellier show that extensive woodland clearance took place before 2000 years ago by Iron Age pastoralists. During the last two thousand years woodlands have grown and declined as a result of variations in both climate and human activity. NATURALISED TREES (Table 2) The native tree flora conspicuously lacks a spruce, fir, larch, beech or maple (Scott, 2002); but examples of all of these have been introduced to Scotland and have shown themselves capable of natural regeneration. Table 2: Naturalised Trees Naturalised refers to an alien plant that has become self perpetuating (Stace, 1997) a) Most common naturalised trees. Source: Milner, 1992 Several common trees that were introduced to these islands over the past few hundred years have since become naturalised. For instance, the Sycamore {Acer pseudoplatanus), which originated in central Europe, is now one of our most common trees, invading much of our scrub and woodland and growing vigorously as far north as the Shetlands and at altitudes of up to 460m. in mountain areas. The following is a list of the most common naturalised species, with their approximate dates of introduction: Norway spruce Picea abies Native in last glacial period Reintroduced early 14“’ centurySweet chestnut Castanea sativa Introduced c. 100 AD SycamoreAc^r pseudoplatanus Introduced c. 1250 AD Walnut Juglans regia Introduced before 1000 AD White poplar Populus alba Early 14“’ century Plane Platanus orientali 1350 Holm oak Quercus ilex 1580 SWy ex fix Abies alba 1603 Horse chestnut Aesculus hippocastanum 1616 European larch Larix deciduas 1620 False acacia Robinia pseudoacacia 1630 b) referred to as regenerating in ‘The Changing Flora of Glasgow’, 2000: h^mxmxxi Laburnum anagyroides 1560 Norway maple Acer platanoides 1638 Turkey oak Quercus cerris 1735 Grey aldex Alnus incana 1780 Sitka spruce Picea sitchensis 1831 Lawson’s cypress Chamaecyparis lawsoniana 1854 Balsam poplar Populus trichocarpa 1880 Ontario poplar Populus candicans (‘Aurora’) 1925 22 x willows + beech, hornbeam, field maple and lime (not native to Scotland but naturalised). The dates of introduction are derived mainly from Wilkinson, 1981; also Campbell-Culver, 2001 and J. Dickson, pers. comm. THE LAST 200 YEARS The Gingko, a ‘fossil’ tree unaltered for 200 million years native to China, and Chile pine are examples of trees introduced in the 18“’ century. However the previous two centuries have seen an explosion in the number of tree species introduced to these islands as a result of the activities of the great plant hunters such as Menzies, Douglas, Fortune, Lobb, Forrest and Wilson. Five of the more remarkable examples of conifer introductions are as follows. Three conifers from the Pacific North-west have grown to more than 200 feet in Britain: Sitka spruce, Douglas fir and grand fir. Sitka Spruce. Sitka spruce is regarded by foresters as the ‘designer tree’. It has an extraordinary natural range of 1500 miles with uniform features. It is widely planted in poor, upland, acidic peat soils, where it can grow rapidly to produce a marketable stand of high quality timber from areas that are otherwise non-productive. It forms the most productive woods in the temperate world. Forest policy has introduced a random planting pattern of alternative species to replace the “ serried ranks of conifers” so derided by conservationists. Douglas fir. The tallest 50 trees in the UK are Douglas firs and the tallest two are located in Glendaruel, Argyll and Moniack Glen, Beauly (Alderman, 2002). The Grand Fir The grand fir at Cairndow, Argyll was the first in the country to achieve 200 feet and for a while held the title of tallest tree in GB . The Coastal Redwood. The coastal redwood. Sequoia sempervirens, is the tallest tree in the world, found growing in the coastal fog belt of N. California. It survives up to 2000 years. The Giant Sequoia The giant Sequoia, Sequoiadendron giganteum, from the Sierra Nevada, has the biggest volume of any tree in the world. Within 80 years of its introduction in 1853 it was the biggest tree by volume in every county of GB. The largest are C.4000 years old. Both of these trees grow exceptionally well in our climate. Many broadleaved trees have been introduced during this period but in height none of them remotely approaches that of champion conifers. Aesthetically however many of them surpass native species. Bright autumn tints distinguish introduced trees such as Japanese maples from the less bright natives. Furthermore the bark of species such as the Snake barked maple and Tibetan cherry is something to be marvelled at. The 20* century has seen a reduction in the number of foreign trees being introduced but two may be mentioned: 9 The Dawn redwood. Metasequoia glyptostroboides , was known from Pliocene fossils when it was found in Hupeh province in 1941. Now there is one in most arboreta. The Bristlecone pine, Pinus aristata, from the Rockies is closely related to P. longaeva, the oldest living tree at 5000 years. THE CURRENT STATUS OF BRITISH NATIVE TREES Most tree habitats in Britain have a long history of human interference and so it could be argued that there are no natural habitats left. Introduced specimens of a particular species bring alien pollen into the landscape and so pollute the native species. Many populations are only vaguely related to our original post-glacial trees. Small-leaved lime is the most researched species in this category. Isolated coppice stools are known that date back 2000 years. For hundreds of years the climate has not suited small-leaved lime particularly well but recent warm summers have encouraged the species to produce fertile seeds again (Gray & Grist, 2000). The native black poplar is very rare on account of the introduction of more productive hybrid timber trees (Miles, 1999). Willows are often adulterated by promiscuity, e.g. the grey willow {Salix cinered) forms with other species three hybrids named in The Changing Flora of Glasgow’ and five in the British Isles (Stace, 1997). Furthermore as many as 18 species of willow (including shrubs) may be found in Britain (Lusby, 2001). Imported beech and oak have caused considerable genetic pollution of existing trees. It has been argued that since common oak was better for shipbuilding purposes it was introduced to Scotland in preference to sessile oak with the result that much hybridisation with the local sessile species occurred. MAN & NEW ECOSYSTEMS Man has created entirely new ecosystems and some examples of these are as follows: Heather Moorlands Heather moorlands are largely an artifact (Gimingham, 1975). Along exposed coasts and at high altitude where trees compete poorly moors are natural. However the treeless tracts of heath that cover the Scottish highlands only appeared with the arrival of man and sheep grazing. Burning and grazing have prevented the heaths from returning to forest for thousands of years. With the decline in profitability of sheep grazing heath is being replanted with more profitable forests over much of its range. It is debatable whether this is a return to a natural state. Britain was heavily forested before the arrival of pastoralists. However burning and grazing for such a continuous length of time has arguably changed what is natural for these lands. On many heaths tree seed is so scarce that even when burning is halted and sheep removed forests do not immediately return. Many wild species have grown dependent upon these artificial ecosystems, including a number of endangered Arctic birds. Chalk Fauna of the Downs The unique chalk flora of the downs is an assemblage of light demanding plants that could never have flourished in the pre-settlement forests. This chalk and limestone grassland habitat possesses a wide variety of alkaline tolerant plants on the thin, dry and nutrient poor soils. The large blue butterfly is a species that requires wild thyme and ant grubs, Myrmica sabuleti, which rely on such close-cropped grasslands (Asher et al., 2001). Since the decimation of rabbits in the 1950’s by the introduction of myxomatosis the grass has grown taller, the wild thyme and ants scarcer, and the butterfly is in trouble. Yet the rabbit was itself an alien introduction of mediaeval times; it was imported from the continent and kept for food and fur. So what is natural is debatable. Perhaps a landscape with alien rabbits and native blue butterflies or one without either. Linear Features Studies have found that many artificial linear features in the landscape - hedges, roadside verges and ditches, for example - often support a greater diversity of life than is found in the open countryside. Roadside vegetation serves as a breeding habitat for 20 of 50 native mammals, 40 of 200 birds, all 6 reptiles, 5 of 6 amphibians and 25 of 60 butterflies. Such vegetation also serves as corridors linking larger areas of wildlife habitat and so increase the chances of a species survival. Agriculture Agriculture has been practised for thousands of years. All the recent gains in production have come through technological improvement rather than clearing more land. Timber Some 75% of the total world production of commercial timber comes from temperate forests, almost entirely from lands that have been managed for sustained timber production for more than half a century. These plantations could meet the entire world’s timber needs on a mere 5% of the area of all the world’s existing forests. ANIMAL BIODIVERSITY (KITCHENER, 1998) There were two main causes of extinction of mammals since the end of the last Ice Age: 1 . Climate change created new habitats where cold adapted species could no longer survive, e.g. reindeer 2. Human activity by a) hunting, e.g. polecat b) habitat destruction for agriculture e.g. lynx and c) hybridisation between native and introduced species. Table 3 gives examples of other mammals that have become extinct since the last Ice Age. Several large species were dependent on forest habitats, e.g. beaver, wolf, brown bear, lynx, wild pig and moose, but by the 18* century Scotland 10 Table 3: Extinction of land mammals in Scotland since the end of the last Ice Age. Species Date/Time when lost Causes of extinction Giant deer {Megaloceros giganteus) ?Mesolithic C Wild horse (Equus ferus) ?Mesolithic C, ?X, ?K Reindeer {Rangifer tarandus) ?Mesolithic C Auroch (Bos primigenius) Bronze age (c. 3500 years ago) ?H,?K Moose (Alces alces) Bronze age (c. 3500 years ago) ?H, 7P/K Brown bear (Ursus arctos) Roman times (or 710'*’ C. AD) H,P/K Beaver {Castor fiber) c. 1550 AD ?K,?H Wild boar {Sus scrqfa) c. 1600 AD ?K, ?H, ?X Wolf (Canis lupus) 1743 AD P,?H Lynx {Felis lynx) 1770 AD ?H, 7P/K Red Squirrel (Sciurus vulgaris) c. 1800 AD H,?D Polecat {Mustela putorius) c. 1912 AD P,?K Key: C - climate change; D - disease; H - habitat loss; K- hunting; P - persecution; X - hybridisation with domestic species. Sources: Kitchener, 1998, 2001 and Humphrey & Quine, 2001 was amongst the most deforested countries in Europe. The large mammals had become extinct except for red deer which adapted to open moorland. Even the red squirrel may have become extinct. Most of today’s population derives from introductions in the 18'*’ and 19“’ centuries from England and Scandinavia. The red squirrel today may be declining because of its inability to digest acorns (Thomas, 2000). The acorns contain a digestive inhibitor that greys can ameliorate but reds cannot. Hence reds are more successful in conifer plantations where they feed on more nutritious pine seeds and where there are no oaks to give greys a competitive edge. Bird extinctions have three main causes (Table 4): Habitat loss. Wetland drainage has resulted in the loss of larger birds such as crane, bittern and white stork. The capercaillie became extinct as the Caledonian pine forest declined. It breeds in coniferous woodland from Norway to Siberia, with glacial relics in the Alps & Pyrenees. Birds of Swedish stock were re-introduced to Perthshire in 1837. From there they have colonised the E. Highlands and Loch Lomondside, where the islands contain amongst the highest densities in Scotland (Mitchell, 2001). Hunting. The great auk, for example, was hunted to extinction. The last known British specimen was killed on Stac an Armin, St. Kilda, in 1840. Persecution. Birds of prey suffered severe persecution from gamekeepers, sportsmen and farmers, e.g. osprey (returned to Scotland in 1954 to Loch Garten, eyrie built in Scots pine), goshawk, white-tailed sea eagle and red kite. The overall temperature drop of each glaciation drove the bird population E, SE & S with the retreating vegetation. The birds returned N & W in a process that continues today. In the last 100 years > 20 species have thus extended their range. Yet > 50 species nest regularly in Britain but not in Ireland, a phenomenon not easily explained. Mammal introductions: There are three main reasons for which mammals have been introduced (Table 5): 1 . Aesthetic e.g. fallow deer 2. Accidental e.g. ship stowaways, such as rats; escapees such as rabbits 3. Sport e.g. sika deer, brown hares Invertebrates and trees. The number of invertebrates found on native trees is much greater than that found on introductions, although non-native conifer species appear to provide suitable habitat for a wide range of native fauna (and flora) (Humphrey & Quine, 2001). Some 500 species are associated with oak and 450 with the genus Salix. Most of these are insects, particularly moths and beetles. Of course rarity can be as important as sheer numbers, e.g. endangered species found on aspen are the aspen hover fly and the rare dark bordered beauty moth. The SWT hopes to create aspen corridors to keep these rare insects in existence. Table 6 summarises gains and losses of mammals and birds since the last ice age. The number of birds and mammals lost or gained is in itself less important than the effect of losses or introductions on the indigenous fauna. The absence of top mammalian carnivores causes much ecosystem imbalance such as an excess of deer leading to a lack of young trees. It has to be said that animal introductions have proved to be far more devastating than plant introductions. Mink and water voles The American mink was first brought to Britain in the late 1920’s to be farmed for its fur. However it escaped into the wild in 1938, colonised successfully and is now implicated in the severe decline of the water vole (Kitchener, 1998). 11 Table 4. Extinction of birds in Scotland since the end of the last Ice Age. Table 5. Introductions of land mammals to Scotland since the end of the last Ice Age. Species Date of Extinction in Scotland (Date of Global Extinction) Cause of Extinction. Species Date of arrival AD Purpose Success (last record) Bittern 1830 AD P Red-necked Wallaby [Botaurus {Macropus 1975 AD A Y stellaris) rufogriseus) Osprey 1916 AD P American Mink 1938 AD E Y (Pandion {Mustela vison) haliaetus) Fallow Deer {Dama 900 AD K. A Y Sea Eagle 1918 AD P dama) (Haliaeetus albicilla) Sika Deer (Cervus 1870 AD K, A Y nippon) Red Kite 1884 AD P (Milvus milvus) 71917 AD Wapiti {Cervus 1819 AD K N canadensis) Goshawk 1883 AD P {Accipiter White-tailed Deer 1832 AD ?K,? A N(I872) gentilis) {Odocoelius virginianus) Capercaillie 1785 AD H.K {Tetrao Grey Squirrel {Sciurus 1892 AD A Y itrogallus) carolinensis) Spotted Crake 1912 AD H Canadian Beaver 1875 AD A N (1903) {Porzana {Castor canadensis) porzana) Muskrat {Ondatra 1927 AD E N(I937) White Stork 1416 AD* ?H zibethicus) {Ciconia ciconia) Orkney Vole 3700 BC - E Y {Microtus arvalis) 3400 BC Great Auk 1840 AD K (Pinguinus (1844 AD) House Mouse Iron Age E Y impennis) {Mus domesticus) Crane ? K,H Black Rat Iron Age or E N* {Grus grus) {Rattus rattus) H'C AD Great Bustard lO'^C AD ?H,?K Brown Rat ? I730’s; E Y {Otis tarda) {Rattus norvegicus) 1744-1754 Great Spotted 1840-1850 AD H Rabbit {Oryctolagus I3thC E Y Woodpecker cuniculus) {Dendrocopos major) Brown Hare {Lepus ? IstC K Y europaeus) H = Habitat loss. K = Hunting. P = Persecution. ^ - Aesthetic. K - Hunting. E - Accidental Source: Kitchener, 1998. N = No.Y= Yes. Source: Kitchener, 1998. * excluding population on Shiants. 12 Table 6: Birds & Land Mammals: Numbers of Species Gained and Lost Since End of the Last Ice Age in Scotland Native Mammals extinct = 12 Re-introductions = 1 Net loss = 1 1 Native Birds extinct = 12 Re-colonisation & re-introduction = 7 Net loss = 5 Alien colonisations & introductions: Mammals = 15 (4 unsuccessful) = 1 1 Birds = 44 (4 unsuccessful) = 40 Totals =51 Net gains/(losses): Mammals Birds Net gains 1 1 40 Net losses 1 1 5 Totals: 0 35 Sources; Summary of Tables 3, 4, 5 and, for bird introductions. Kitchener, 1998 The powan The powan is a relict cold water fish confined to a few mountain lochs. In Loch Lomond the introduced ruffe is a significant predator of its eggs. So reserve stocks of this legally protected fish have been introduced to Loch Sloy and the Carron Reservoir (Mitchell, 2001). The introduced NZ flatworm The introduced NZ flatworm has a negative impact on native worms and is spreading rapidly. Crayfish The aggressive and invasive signal crayfish {Pacifastacus leniusculus) has almost wiped out the native white-clawed species {Austropotamobhis pallipes) (Maitland et al., 2001). Anyone wishing to discover more about the controversial topic of ‘Alien species: friends or foes?’, especially in a Scottish context, need look no further than the publication of the proceedings of the symposium about this topic held in 2001 to celebrate the Society’s 150* anniversary. Although this has been a wide ranging address, the world of micro-organisms has not been mentioned, but I hope to have demonstrated that whereas plant introductions have overall provided a welcome increase in this country’s biodiversity animal introductions are proving to be far more controversial. ACKNOWLEDGEMENTS For providing me with slides to illustrate this address I thank Ewen Donaldson of Glasgow Botanic Gardens, Mogens Hansen of the Society, Frank Morris of St. Aloysius’ College, Glasgow and Richard Sutcliffe of Kelvingrove Museum. For helpfully commenting on a draft of the manuscript I thank Prof. Jim Dickson and Dr. Petra Mudie, as well as Keith Watson and Bernard Zonfrillo of the Society. REFERENCES Alderman, D. (2002). Registrar’s Report. The Tree Register Newsletter No.ll. The Tree Register of the British Isles, Bedford. Asher, A. et al. (2001). The Millennium Atlas of Butterflies in Britain and Ireland. Oxford University Press, Oxford. Budiansky, S. (1995). Nature’s Keepers. Phoenix, London. Campbell-Culver M. (2001). The Origin of Plants. Hodder Headline, London. Dansgaard, W., White. J.W.C. and Johnsen, S.J. (1989). The abrupt termination of the Younger Dryas. Nature 339,532-534. Dickson, C. & Dickson, J.H. (2000). Plants & People in Ancient Scotland. Tempus, Stroud. Dickson, J.H., Macpherson P. and Watson K. (2000). The Changing Flora of Glasgow. Edinburgh University Press, Edinburgh. Ennos, R.A., Sinclair, W.T. and Perks, M.T. (1997). Genetic Insights into the Evolution of Scots pine, Finns sylvestris L., in Scotland. Botanical Journal of Scotland 49 (2), 257-265. Farrow, G.E. (2001). James Croll: 19"’ Century Pioneer of Climate Change. The Glasgow Naturalist 23 (6), 9-18. Fischer, H., Wahlen, M., Smith, J., Mastroianni, D. and Deck. (1999). Ice core records of atmospheric CO, around the last three glacial terminations. Science 283, 1712-1714. Fitter, R. (1967), The Penguin Dictionary of British Natural History. Penguin Books, Middlesex. Gimingham, C.H. (1975). An Introduction to Heathland Ecology. Oliver & Boyd, Edinburgh. Gootes, P.M., Stuiver, M.. White, J.W.C. , Johnsen, S. and Jonzel, J. (1993). Comparison of oxygen isotope records from the GISP2 and GRIP Greenland ice cores. Nature 336, 552-554. Gray, R.K.S. and Grist, N.R. (2000). Natural regeneration of limes {Tilia spp.) in Scotland: locally widespread and more numerous in 1999. The Glasgow Naturalist 23 (5), 13-16. Gribben, J. and M. (2001). Ice Age. Penguin Books, London. Hadfield, M. (1996). Pioneers in Gardening. Bloomsbury. Haeckel, E. (1866[1988]). Generelle Morphologie des Organismus, Bd.2: Allgemeine Entwicklungsgeschichte . Reprint. Berlin: de Gruyter. Humphrey, J. and Quine, C. (2001). Sitka spruce plantations in Scotland: friend or foe to biodiversity? The Glasgow Naturalist 23 Supplement 66-76. Huntley, B., Daniell, J.R.G. and Allen, J.R.M. (1997). Scottish vegetation history: the Highlands. Botanical Journal of Scotland 49 (2), 163-175. Idso, S.B. (1989). CO,, soil moisture and future crop production. Soil Science 147, 305-307. Johnsen, S.J., Clausen, H., Dansgaard, W., Fuhrer, K., Gunderstrup, N.S., Hammer, C.U., Iverssen, P., Jouzel, J., Stauffer, B. and Steffensen, J.P. (1992). Irregular glacial interstadials recorded in a new Greenland ice core. Nature 359, 3\\A13. Johnson. H. (1993). The International Book of Trees. Mitchell Beazley, London. Kitchener, A.C. (1998). Extinctions, Introductions and Colonisations of Scottish Mammals and Birds since the Last Ice Age. Pp. 63-92 in Lambert, R.A. (editor) Species History in Scotland. Scottish Cultural Press, Edinburgh. Lamb, S. and Sington, D. (1999). Earth Story. BBC Books, London. 13 Lambert, R. A. ed. (1998). Species History in Scotland. Scottish Cultural Press, Edinburgh. Lusby, P. and Wright, J. (2001). Scottish Wild Plants. Mercat Press, Edinburgh. Maitland, P.S., Sinclair, C. and Doughty, C.R. (2001). The status of freshwater crayfish in Scotland in the year 2000. The Glasgow Naturalist 23 (6), 26-32. Marren, P. (1990). Woodland Heritage. David & Charles, Devon. Miles, A. (1999). Silva. Ebury Press, London. Milner, J.E. (1992). The Tree Book. Collins & Brown, London. Mitchell, A. & Coombes, A. (1998). The Garden Tree. Weidenfeld & Nicolson, London. Mitchell, A. (1974). A Field Guide to the Trees of Britain and Northern Europe. Collins, Glasgow. Mitchell, A. (1996). Trees of Britain. HarperCollins, London. Mitchell, J. (2001). Loch Lomondside. HarperCollins, London. Morton, A. (1998). Tree Heritage of Britain and Ireland. Swan Hill Press, Shrewsbury. Petit, J.R., Jouzel, J., Raynaud, D., Barkov, N.L., Barnola, J.-M., Basile, L, Bender, M., Chappellaz, J., Davis, M., Delaygue, G., Delmotte, M., Kotlyskov., V. M., Legrand, M., Lipenkov, V.Y., Lorius, C., Pepin, L., Ritz, C., Saltzman, E. and Stievenard, M. (1999). Climate and atmospheric history of the past 420,000 years from the Vostok ice core, Antarctica. Nature 399, 429-436. Ramsay, S. and Dickson, J.H. (1997). Vegetational history of central Scotland. Botanical Journal of Scotland A9 (2), 141-150 Rigozo, N.R., Echer, E., Vieira, L.E.A. and Nordemann, D.J.R. 2001. Reconstruction of Wolf sunspot numbers on the basis of spectral characteristics and estimates of associated radio flux and solar wind parameters for the last millennium. Solar Physics 203, 179-191. Salisbury, F.B. and Ross, C.W. (1978). Plant Physiology. Woodsworth Publishing Co., Belmont CA. Scott, A. (2002). A Pleasure in Scottish Trees. Mainstream, Edinburgh. Simms, E. (1990). Woodland Birds. Collins, London. Smith, A. (1997). All for a Handful of Seed. Smith A.K.. Perth. Smout, T.C. ed. (1997). Scottish Woodland History. Scottish Cultural Press, Edinburgh. Stace, C. (1997). New Flora of the British Isles. Cambridge University Press, Cambridge. Thomas, P. (2000). Trees: Their Natural //Afo/7.Cambridge University Press, Cambridge. Wilkinson, G. (1981). A History of Britain’s Trees. Hutchinson, London. 14 Glasgow Naturalist 2002. Volume 24 Part 1 . Pages 15-22. Climate Change. Its History and Future in Relation to Scotland's Landscape, People, and Economy Editorial. Azra Meadows and Peter Meadows Division of Environmental and Evolutionary, Biology, Graham Kerr Building, Institute of Biomedical and Life Sciences, University of Glasgow G12 8QQ INTRODUCTION 'Flavit et dissipati sunt' (It blew and they were scattered) Unusually changeable weather conditions occurred around Britain from 1500 to 1800, ending with the mini ice age in the late 18th century. Major North Sea storms developed in 1530, 1570, 1634, and 1697, and 1694 the Culbin sands, just west of Nairn on the Moray Firth coast, shifted significantly during very high winds. A less dramatic storm occurred in September 1588. However this one changed history as it destroyed many of the ships of the Spanish Armada, and as a result Queen Elizabeth I of England issued a commemorative medal inscribed 'Flavit et dissipati sunt' - referring to the Spanish Armada. So climate change of one sort or another is a normal feature of UK weather, and we can expect more of it during the coming decades. There are a number of excellent general and more technical accounts (Harding, 1982; Libby, 1983; Shackleton et al, 1988; Leggett, 1990; Ribot et al., 1996; Curran, S. 1998; Department of the Environment, n.d.; Drake, 2000 - reviewed by the authors in this volume; Cookson, 2002). We address this very topical subject in our article, focussing on the importance of climate change for Scotland. James Croll was also very much aware of this in the last century, as Farrow points out in a recent and most elegant paper in the Glasgow Naturalist (Farrow, 2001). We begin by emphasising the importance of the interaction between mountains, rivers and the coastal zone in relation to climate change. This is followed by a consideration of historical evidence for climate change in Scotland and England. We then consider current views on climate change and the importance of greenhouse gases in determining weather patterns and global warming. Predicted climate and weather change in Scotland has been the subject of an exhaustive recent study by Kerr, Shackley, Milne and Allen (1999) entitled 'Climate Change : Scottish Implications Scoping Study'. In the last part of our article we discuss and comment on some of the major points raised by this study, which was commissioned by the Scottish Executive Central Research Unit. As is often the case in government reports, there are some very specific points raised, but a great deal of generality. Mountains, rivers and the coastal zone as AN interacting ENVIRONMENT Mountains rivers and the coastal zone are extremely important interrelated ecosystems on a global scale that act together in terms of responding to and often influencing global climate change. Until very recently few scientists realised their integrated nature, and the central part they play in climate change and in controlling the living conditions of human populations - including the resources of land and water (Dobby 1962; Groombridge 1992). This is especially relevant for Scotland, in terms of predictions of climate change and global warming (Manley, 1975; Ford, 1982; lUCN 1990; Houghton et al. 1996; Watson, Zinyowera & Moss 1996; Batterbury, Forsyth & Thompson 1997). Scotland is made up of mountains, hills and rivers, and is surrounded by a coastline that is often indented by extensive marine loch systems. It also contains magnificent freshwater lochs, vast tracts of uninhabited highlands in its centre, and habitats that contain many rare and endangered species of plants and animals. Its weather is relatively dry on the east coast, wet on the west coast, and in the central highlands alternately very cold in winter and warm in summer. Its large cities, Glasgow, Edinburgh, and Aberdeen, perhaps belie the essentially rural nature of the country. The existence of much of the human population is in scattered villages and small towns around the coast connected by coastal road systems, together with a few highland villages joined by small roads that cross inhospitable but very beautiful country. Scotland has been shaped in the past by large glaciers during the most recent ice age that only ended approximately 12,000 years ago. Its whole landscape of mountains, glens, rivers, lochs and coast have been shaped by these events. How will climate change alter this? Table 1 summarises some of the more important aspects of how climate change will affect natural and human impacts on these systems. Many of these aspects are of direct relevance to Scotland, as an inspection of the asterisked items show. They include decreased snow cover, deforestation, erosion, flooding, sea level rise, storms, and sea defences. In Scotland as elsewhere, the erosion of hillsides and mountains is a natural process. Rain and snow feed mountain streams and cause periodic flash floods. The rain, snow, streams and flash floods carry water down hillsides and at the same time cause erosion. This erosion consists of sedimentary material - small rocks, stones and soil particles - carried along by the water. The whole process feeds water and sediment into the main branches of a river that lie at lower levels. Periodic river floods in the lower reaches of rivers 15 distribute water and fine sediment onto the flatter country there, thus producing good agricultural land. It is not often appreciated that the soil cultivated by present day man has been transported there from mountainous areas in past geological periods. This soil has been transported by the continuous action of melting glaciers in the past the present day action of rivers, and by the floods. Eventually rivers discharge both their water and fine suspended sediment into the coastal marine environment. The huge mud and sand banks in as a habitat for the small invertebrates that migrating birds feed on. The mountains, rivers and the coastal zone of Scotland have therefore to be considered as a series of contiguous interacting ecosystems, in which water and sediment are transported from high mountain environments, onto river basins to form rich agricultural land, and then to the coastal zone and into the oceans. The implications of this, and of future changes in the climate over Scotland in terms of human habitation, resource potential and biodiversity cannot be overemphasised. On a global stage this has been recognised by the 1992 Rio Earth Summit on Biodiversity, the 1997 Earth Summit II in New York, the Kyoto Climate Summit in December 1997, together with international scientific conferences such as the Indus River meeting in 1994 (Meadows & Meadows 1999b). These initiatives show that at last political and scientific communities are becoming aware of the future dangers and threats. Table 1 . Global climate change and Natural and human impacts on mountains, rivers and coastal zones. *Major impact of climate change likely. (Modified from Meadows & Meadows 1999a). TYPE OF IMPACT PROCESS OUTPUT ENVIRONMENTAL ENVIRONMENT MANAGEMENT Natural impact. Melting of Water discharge. Prevention of build-up glaciers*. soil erosion. of greenhouse gases*. Hills decreasing and snow cover* , Mountains weathering of rocks. Soil erosion. Reforestation*, Human impact. Deforestation*, erosion*. silting. managed land use. Natural impact. Flooding*. Soil, Flood control particulates. defences*, dams. salts, water discharge. barrages, dykes. Rivers and Human impact. Water logging. Abandoned Construction of outfall Flood plains desertification* agricultural drains. salinisation. land. Industrial, Pollutant domestic discharge. Pollution control. pollution. water quality. Natural impact. Sea level rise.* Flooding of low Coastal defences*. Storms*. lying Hurricanes. coastlines*. cyclones. Loss of human Weather predictive life, habitat. models, satellites. Coastal Zones Estuaries Human impact. Industrial and Decline in Pollution monitoring. and domestic fisheries*. Firths pollution. Eutrophication*. Over- Depletion of Managed resource exploitation of natural exploitation*. natural resources. resources. Habitat Loss of nursery Conservation, degradation. grounds for fish. protection. 16 HISTORICAL CLIMATE CHANGE IN SCOTLAND AND ENGLAND Climate change is not new, as we have seen. The following contemporary quotations from Baker (n.d. In fact c. 1883 - editors) describe unusual and often catastrophic weather changes over Scotland and England. Many of them still have an immediacy. 764 AD: 'There fell such a marvellous great snow, and therewith so extreme a frost, as the like had not been heard of, continuing from the beginning of the winter almost till the midst of spring, with the rigour whereof trees and fruits withered away, and not only feathered fowls, but also beasts on the land and fishes in the sea died in great numbers Holenshed; Roger de Hoveden'. 793 AD: 'Dreadful prodigies alarmed the wretched nation of the English; for terrific lightnings and dragons in the air and strokes of fire were seen hovering on high, and shooting to and fro, which were ominous sighs of the great famine and the frightful and ineffable slaughter of multitudes of men which ensued (Roger de Hoveden) 800 AD: 'On the ninth day before the calends of January, the day before the Nativity of our Lord, a mighty wind, blowing either from the south or south-west, by its indescribable force destroyed many cities, houses, and towns in various places; innumerable trees were also torn up from the roots. In the same year an inundation took place, the sea flowing beyond its ordinary limits (Roger de Hoveden; M. of W.).' 1092 AD: 'By the high spring tides many towns, castles, and woods were drowned, as well in Scotland as in England. After the ceasing of the tempest, the lands that sometime were Earl Goodwin's, by violent force and drift of the sea were made a sandbed, and ever since have been called Goodwin's Sands. Such dreadful thunder happened also at the same time, that men and beasts were slain in the fields, and houses overturned even from their foundations. In Lothian, Life, and Angus, trees and corn were burnt up by fire kindled no man knew how (Holenshed).' 1210 AD: 'Inundation at Perth about the time of the Feast of St Michael, which carried off much of the harvest crops from the fields. The waters of Tay and Almond so swelled that the large bridge of St John was overthrown (Fordun and Major).' 1333. AD. 'November 23. At night through a marvellous inundation and rising of the sea all along by the coasts of this realm but especially about the Thames, the sea banks or walls were broken down with the violence of the water, and infinite numbers of beasts and cattle drowned fruitful grounds and pastures were made salt marshes, so as there was no hope that in long time they should recover again their former fruitfulness (Holinshed).' 1771 AD: 'Dr Johnson says that the season was so severe in the Island of Skye that it is remembered by the name of the Black Spring. The snow, which seldom lies at all, covered the ground for eight weeks, many cattle died, and those that survived were so emaciated that they require no male at the usual season. The case was the same in the south; never were so many barren cows known as in the spring following that dreadful period. At the end of March the face of the earth was naked to a surprising degree. Wheat hardly to be seen, and no signs of any grass; turnips all gone, and sheep in a starving way. All provisions rising in price. Farmers cannot sow for want of rain (Gilbert White). May 6. By letters from Gloucester we learn that the late rains have produced such an alteration that everything promises a plentiful crop, though a late one.' It is even possible to construct historical weather patterns from contemporary records. Kington's classic book on 'The Weather of the 1780's over Europe' (1988) shows what can be done by attention to detail and an inspection of the records of learned and scientific societies. These also show that climate change was happening the whole time, with unexpected storms, very hot summers or cold winters, and flooding. These records of meteorology began early in the European scientific and cultural renaissance of the 17th and 18th centures. In 1653, Ferdinand II, the Grand Duke of Tuscany and a member of the Medici family, arranged for a network of permanent meteorological stations to be set up in Italy .There were about 12 stations mainly in northern Italy, and data was entered on standard forms concerning pressure, humidity, temperature, wind direction and sky state. The data was collected largely using standard instruments, and then sent to Florence to the Accademia del Cimento (the Academy of Experiments). Unfortunately the system broke down when the Academy was disbanded in 1667. However Kinton (1988) reminds us that Robert Hooke, the first Curator of the Royal Society, proposed in 1667 'A Method for Making a History of the Weather'. He also designed instruments that included a barometer for use on land and at sea, an anemometer for measuring wind source base on a pressure plate, and a thermometer graduated with zero based on the freezing point of water. By 1723, the Royal Society of London was recording weather data in Weather Journals, with details of temperature, rain, wind, and sky state. In France things were also happening. The Societe Royale de Medicine was established under Louis XVI , and by the mid 1780s there was a network of over 70 weather stations across France which was extended to receive information from America and Asia. These and similar reports taken together show that climate change is happening the whole time. Perhaps the relative stability over the past half century may be an exception. But even here, many readers will remember the very cold winter of 1947 and 1 963 - when the sea froze along parts of the 17 Scottish coastline. Temperatures of about -25^C occurred in the Kirklee area of Glasgow during one winter in the 1980's, -18^C over the Christmas and New Year period during one winter in the 1990's, and there was an extremely hot and dry summer in 1987. CURRENT VIEWS ON CLIMATE CHANGE. THE IMPORTANCE OF GREENHOUSE GASES Current views of potential climate change are focusing on temperature change, alterations in the amount and distribution of rainfall, change in sea level, and frequency and location of storms. However there is no universally accepted consensus in spite of the clear evidence that world temperatures have increased significantly since the beginning of the industrial revolution in the early 19th century, and that sea levels are rising on some coastlines. Much of the recent change in temperatures, often referred to as global warming, is related to the increase in release of greenhouse gases. A quick explanation of the effect will help. The sun's energy warms the earth. This energy is absorbed by the earth and some of it is irradiated back into space as long wavelength infra red radiation. Greenhouse gases in the upper atmosphere stop some of this long wavelength infra red radiation leaving the earth's atmosphere and reaching space. As a result this radiation is trapped and the earth gets fractionally hotter - global warming.. The difficulty is the lack of precision in predictions. It is not possible to obtain an accurate prediction of the amount and distribution of global warming that is likely to occur. It is also very difficult to identify how much of the global warming that has taken place during the last 150 years is caused by man's activities as compared to natural long term perturbations in weather patterns. Atmospheric carbon dioxide, a major greenhouse gas, has certainly been rising since the beginning of the industrial revolution in the early nineteenth century. Currently, the burning of fossil fuels contributes 80% of this and is rising at about 0.5% per year. World temperatures have broadly increased over the same period. Other greenhouse gases such as methane, nitrous oxide, chlorofluorocarbons (CFCs) and hydrochlorofluorocarbons (HCFCs) are being added to the atmosphere by man's activities (Table AAA). All of these gases will certainly add to global warming by acting as a blanket in the upper atmosphere. It is particularly worrying that CFCs and HCFCs have effects that are two or more orders of magnitude greater than methane or carbon dioxide (Table 2, column 2). For example HCFC 22 has 1600 times the effect of carbon dioxide, CFC 1 1 has 3400 times the effect of carbon dioxide, and CFC 12 has an amazing 7 100 times the effect of carbon dioxide. Table 2. Greenhouse Gases. Their approximate relative greenhouse effect (Global Warming Potential GWP). concentrations in the atmosphere (ppmv: part per million by volume), current rate of change (% per year), and atmospheric lifetime. Global warming potential is the predicted reduction in the amount of heat radiated from the earth produced by a particular gas, in relation to the reduction produced by carbon dioxide over the same time period. It is only a very approximate measure of the relative effect of a greenhouse gas, as errors of up to 35% are recognised (Houghton et al., 1996, p. 21). * carbon dioxide does not have a single lifetime because the uptake rates differ for different processes. CFC = chlorofluorocarbons. HCFC = hydrochlorofluorocarbons. Greenhouse Approximate Current average Cun-ent rate of Atmospheric Gases relative atmospheric change lifetime in years Greenhouse effect concentration (%pa) G.W.P. (ppmv) (1992) Carbon dioxide 1 355 1.8 50 - 200* Methane 11 1.72 0.8 12 Nitrous Oxide 270 0.31 0.25 120 CFC 11 3400 0.00026 4 50 CFC 12 7100 0.00045 4 - HCFC 22 1600 0.0001 12 (Source: The Department of the Environment ND; Houghton et al.. 1996) 18 Predictions suggest an increase in global temperatures of 0.5 °C to 2.5°C by 2030, which is likely to lead to a rise in sea level of between 10 cm and 20 cm. At first sight this does not seem very much, but in very low lying areas such as on the east coast of England and in some low lying areas of Scotland, the effects are likely to be catastrophic. These latter include all the major estuaries in Scotland, the area around Elgin, and some parts of Glasgow through which the Cathcart flows. Floods in these coastal regions caused by high river flows and by abnormally high sea levels associated with low atmospheric pressures - storm surges, can already cause large scale economic problems. There may also be a change in weather patterns and increased frequency of severe storms. Even if worst case scenarios do not occur, it is undoubtedly true that within the last decade there have been significant changes in coastal weather. The southern parts of England have become much drier during summer. These changes are obvious even to the ordinary person, and are already having an Impact on the economy of the areas affected. Conventional crops are suffering, and new cultivars are being assessed that can withstand drier, hotter conditions. These changes are obvious even to the ordinary person, and are already having an impact on the economy of the areas affected. Conventional crops are suffering, and new cultivars are being assessed that can withstand drier, hotter conditions. Coastal areas short of drinking water are being forced to introduce water rationing, and desalination plants - once only seen in the more arid parts of the tropics. PREDICTED CLIMATE AND WEATHER CHANGE IN SCOTLAND THE RECENT CCSIS STUDY A recent 75 page study on climate change in Scotland, 'Climate Change : Scottish Implications Scoping Study' (the CCSIS study) published by the Scottish Executive Central Research Unit makes fascinating reading (Kerr, Shackley, Milne, and Allen, 1999). In some ways it is very specific, but in others it is somewhat vague. There are likely to be a number of quite specific changes in climate and weather in Scotland during the 21st century. The weather will become warmer by about 1.2 to 2.6° centigrade, with more warming occurring in winter than in summer. Rainfall intensity will increase, most noticeably in winter which will lead to higher risks of flooding. Severe gales will increase in number, and short- wave radiation will become reduced because of increased cloud cover. The report identifies two possible megascale phenomena which although unlikely to occur would have profound implications for the climate, and hence for all biodiversity - ourselves included, during the present century. The first is a major alteration of the Gulf Stream and North Atlantic Drift oceanic circulation in the North Atlantic, which brings warm water to the western coasts if Ireland, Scotland and England. This would affect the climate of the whole of Europe and would probably lead to a mini-ice age in the region. The second is the collapse of the West Antarctic ice sheet which in turn would lead to an increase in the rate and degree of the rise in sea level, climate change, but surprisingly little on means of adapting to climate change. Means of adapting to climate change can be natural or planned. Natural adaptations consist of biological change such as potential changes in bird migration patterns, human sociological changes, and potential changes in human outdoor activities. The authors define planned adaptations as those planned by man which may include such items as revisions of building codes and improved air conditioning systems. They also include consideration of the aims of a biodiversity policy for Scotland, stating that 'a static view of natural conservation is not viable'. The second aspect concerns policies that are intended to reduce human-induced climate change, and the impact of these policies on Scottish society. Specifically, the UK government is required to meet international obligations on reducing greenhouse gas emissions. It must reduce its emissions of the six most important greenhouse gases from a 1990 level of 216 million tonnes carbon equivalent, to 189 million tonnes carbon equivalent averaged over the period 2008 to 2012. The UK government has made an additional domestic target of reducing carbon dioxide emissions to 20% below international baselines over the same period. Neither the first aspect nor the second aspect is in any way specific to Scotland, in our view, although the points are worthwhile making on a broader canvas. The CCSIS study goes on to report on two recent workshops organised by the Scottish Office in 1998. Workshops The first workshop focused on 'Climate Change Impacts in Scotland'. The conclusions of the workshop identified a need for higher resolution climate data for Scotland. There were also specific concerns about the impact of climate change on the following resources and infrastructure. 1. Salmon, marine and freshwater fisheries. 2. Oil and gas infrastructure. 3. Hydroelectric plant. 4. Roads, coastal defences and other public infrastructure. 5. Water resources. Highland and subarctic ecosystems. 6. Agriculture and forestry. 7. Skiing industry. 8. Health. 9. Environmental indicators determining critical levels of damage.. The second workshop focused on a 'Climate Change Mitigation Strategy for Scotland'. Its objectives were to raise awareness of the main issues involved in the development of a Scottish Climate Strategy. The workshop considered actions that might be needed and gathered views from sources regarding the development of a Scottish Climate Strategy. It received expert input from over 70 government organisations, public 19 companies, research institutes and universities. The subjects discussed were under five headings. These are worth listing, as if acted on they are likely to have an impact on everyone in Scotland in due course, and also to have a potentially large effect on our environment in towns and villages on the one hand and in the countryside on the other. 1. Business. Fiscal measures to raise investment funds and provide incentives. Regulation and integrated pollution control. Negotiated and voluntary agreements. Emissions trading. Targeted advice and assistance to small and medium enterprises. 2. Energy Supply. Investment in renewable energy and other non- fossil generation. Combined heat and power. Energy services for customers. Green tariffs. Utility regulation. 3. Households. Energy efficiency in the home - lights, appliances, central heating, insulation, advice and information. Financial incentives. Local authorities under the Home Energy Conservation Act. Electricity Standards of Performance. Market transformation strategy, appliance suppliers and government. 4. Land use and forestry. Expansion of woodland areas. Energy crops (biomass). Animal husbandry. Reduced fertiliser use. 5. Transport. Increasing fuel duties. Voluntary agreements on car efficiency. Alternative fuels, fuel cells, electricity/hybrid vehicles. Best practice techniques for fleet operators. Demand management and model shifts. User charges and traffic management. The Economy in Scotland The economy of Scotland will be affected by climate change to different degrees depending on the area of business operation. Transport including the commercial operation of ships, trains, road transport and associated infrastructure will all be affected by climate change. The CCSIS study states that the existing infrastructure for shipping and fen-y operations in Scotland is robust. However we doubt this. Past experience shows clearly that major storms in the Firth of Clyde and Firth of Forth would do untold damage. Commercial trains in Scotland will need rolling stock and track updating to take into account the worsening rain and gale predictions. The same is true to an even greater degree for roads, and public and private transport, particularly in the Highlands. Service industries in Scotland, retail and finance, will be less affected by climate change. This is important, because the service industries contribute two thirds of Scotland's Gross Domestic Product and three quarters of its workforce. The tourism industry is also very important, especially in the Highlands. Clearly warmer climates will mean more tourists unless it is too wet. But rain is not expected to increase in the summer. Winter sports will suffer if snow fall is reduced much below its current level. The short break holiday of two to three days is usually taken in Scottish cities or towns, and so is unlikely to be affected dramatically by climate change unless it is very dramatic. The Natural Environment in Scotland The natural environment in Scotland is particularly important for agriculture, forestry and fisheries, and the CCSIS study is aware of and comments on these areas. 'Fishing, forestry and agriculture are valued sources of employment and income in rural parts of Scotland, and are therefore an essential element of a sustainable rural economy'. It identifies salmon and trout as being central to sports fishing for example. It will therefore pay us to consider the potential effects of climate change on each of these three areas in turn. 1. Fisheries and Aquaculture. The most economically important Scottish based fisheries are the freshwater salmon and sea- trout fisheries in Scottish rivers, loch aquaculture of salmon and to a lesser degree of trout and shell fish, and the fisheries of the North Atlantic and North Sea. Sea trout and salmon catches have fallen significantly during recent years in west coast Scottish rivers. Possible causes are overfishing, pollution, acid run-off from forestry and changes in predator pressure. It is also possible that salmon farming in cages in sea lochs may have had an impact. These farmed salmon may be a source of parasitic sea lice infecting wild fish. They may also cross breed with wild fish. Climate change is very likely to affect all of the above fisheries, but the precise nature of the effect is difficult to define at present. Any change in water temperature, salinity and oxygen content will have a significant influence on the fish themselves as well as on their food sources. Changes in ocean currents will also disturb larval behaviour and dispersal. Increased rainfall would interfere with the salmon eggs on river beds, and increase in the number or intensity of storms would affect river flow and certainly reduce the efficiency of marine fishing vessels. A significant increase in water temperature in freshwaters and the sea may lead to faster growth rates in commercial species of fish, but it may also cause southern species to migrate northwards. Southern species may need different fishing methods and equipment, and there are cost implications here. 2. Forestry Forestry in Scotland is an important industry as identified by the CCSIS report and also has a significant role to play in attracting tourists to the country. There are also major governmental and farming interests involved. The Forestry Commission is responsible for advice to the UK Government on forestry policy and also the implementation of that policy. In July 1999 the Forestry Commission became answerable to the Scottish Executive for most its activities in Scotland. A Scottish Forest Strategy was launched in November 2001 which will "guide the 20 development of Scotland's expanding forest and woodland area into the 21st century and beyond". Climate change is most likely to influence Scotland's forests through changes in temperature. Increased temperature usually means an increase in growth rate. However changes in the variation of temperature between summer and winter are also likely to be important. Commercial forestry may benefit by being able to grow a larger number of species than at present if the temperature increases significantly, although this effect will take several decades to be noticeable because of the long growth period of most trees. Increased rainfall will have a significant influence in south east Scotland - where it is relatively dry at present, and major storms are likely to cause wind damage by uprooting trees and also limiting the height of growth in exposed sites. All of these effects of climate change will have an important influence on biodiversity. This is as yet not properly quantified. 3. Agriculture Agriculture in its broadest sense is practised on almost 80% of land in Scotland, and most of this is grazing by cattle or more frequently sheep as the weather and environmental conditions are do not allow of much else. Profitable farming is difficult because of this, and also because of the long road distances to the industry's main markets. There have been very few studies of the potential influence of climate change on agricultural practice. However it has been suggested that the predicted changes in climate will only have a marginal effect on agriculture at least for the next century. We doubt this, and think that the effects of increased rainfall, more large storms, and increasing temperatures will together have a significant influence during the next 30 years or so. It is certainly true, however, that a longer growing season caused by higher temperatures would tend to increase the diversity of species that can be grown. These would include fodder maize, sugar beet, and oil seed rape. CONCLUSIONS There is nothing new about climate change. It is clear from the sedimentary and fossil record that climate change has taken place since the formation of the planet Earth. It has also been a driving force in the whole process of evolution. Historically, records from Scotland and England show that unexpected dry or wet, hot or cold, and windy or calm conditions are phenomena which were common and yet continue to surprise. 1771 AD: 'Dr Johnson says that the season was so severe in the Island of Skye that it is remembered by the name of the Black Spring. The snow, which seldom lies at all, covered the ground for eight weeks, many cattle died, and those that survived were so emaciated that they require no male at the usual season.'. It really must have been quite bad. Our current increased awareness of the whole process of climate change in all its aspects must surely come from our increased abilities to predict. This predictive ability is based on highly complex modelling scenarios that would not have been possible before the development of computers. The interactive nature of different parts of the environment is also very important. Mountains rivers and the coastal zone all influence each other and in turn are influenced by changing patterns of climate. This is obvious in Scotland. Global warming together with increasing rainfall - especially in the eastern parts of Scotland, means more flooding, rising sea levels and changing patterns of species distribution. Man has much to answer for here. The totally unacceptable annual increase in the production of Greenhouse gases has to be halted. Carbon dioxide from the burning of fossil fuels, and the increasing release of such gases as Methane, Nitrous Oxide, chlorofluorocarbons and hydrochluorofluorocarbons, will eventually set in motion events that cannot easily be reversed. A change in the North Atlantic Drift current, or the collapse of the West Antarctic Ice sheet are examples. Temperatures are likely to rise 0.5®C to 2.5°C by 2030, which will lead to an average global rise in sea level of 10 cm to 20 cm. This does not sound a great deal, but it will have major impacts on many low lying areas, particularly where land is close to sea level in estuaries around Scotland. Parts of Glasgow and other Scottish cities are also vulnerable. The main problem here is likely to be the onset of storms when tides are very high in Autumn and Spring. To the authors knowledge, there are no contingency plans for major sea level rise in Scotland, although the Scottish Environmental Agency SEPA issues regular flood warnings and advice through its web site and on radio and TV. A recent study published by the Scottish Executive Central Research Unit 'Climate Change : Scottish Implications Scoping Study' (the CCSIS study) (Kerr, Shackley, Milne, and Allen, 1999) has begun the process of officially addressing the medium and long term implications of global warming for Scotland. The study reports on two important workshops held in 1999, and then goes on to predict the effects of global warming on the Scottish climate and environment, on the economy and business, and on fisheries agriculture and forestry. It is clear that within the next 30 to 50 years everyone will be affected. As a selection, winters will get warmer, the east coast will get wetter, and severe storms will increase in frequency. Southern species of animals and plants will move north, and tourism will be affected by warmer weather with in some areas more rainfall. Transport including the operation of ships, trains and road transport will all be affected by the higher level of rainfall and the increased frequency of severe storms. The freshwater and marine fisheries 21 of Scotland will almost certainly be affected, although there is not enough information currently available to allow specific predictions. Effects on forestry will include the introduction of new tree species from warmer climates, and the trees themselves becoming shorter - tall ones may well be torn down by high winds. All of these influences are bound to have major and long lasting effects on the biodiversity of natural and farmed land over the coming years. All in all, it is an interesting time to be living in environmentally. Homo sapiens has lived through it before, but been less aware of it. One only has to look at the snow scenes in Dutch 18th century landscape paintings to realise that we have been here before. REFERENCES Bakler, T.H. n.d. (In fact c. 1883 - editors). Records of the Seasons, Prices of Agricultural Produce, and Phenomena Observed in the British Isles. Simpkin, Marshall & Co., London. Batterbury, S., T. Forsyth and K. Thomson (1997) “Environmental transformations in developing countries; hybrid research and democratic policy”. The Journal of Geography, vo\. 163 : 125-132. Cookson, C. 2002. 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Maskell (eds) (1996) Climate change 1995. Contribution of working group 1 to the second assessment report of the intergovernmental panel on climate change. Cambridge : Cambridge University Press. lUCN (1990) lUCN directory of South Asian protected areas. Gland, Switzerland and Cambridge, U.K. ; lUCN. Kerr, A. Shackley, S. Milne, & Allen, S. 1999. Climate Change: Scottish Implications Scoping Study. Scottish Executive Central Research Unit. Kinton, J. 1988. The Weather of the 1780's over Europe. Cambridge University Press, Cambridge. Leggett, J. (Ed.) 1990. Global Warming. The Greenpeace Report. Oxford University Press. New York. Libbe, L.M. 1983. Past Climates. Tree Thermometers, Commodities, and People. University of Texas Press. Austin, Texas, U.S.A. Manley, G. 1975. 1684: the coldest winter in the English instrumental record. Weather 30, 382 - 388. Meadows, A, & Meadows, P.S. 1999a. Mountains, rivers and coastal zones in Aseia: environmental management and community integration into the 21st century, pp. 49 - 76. In Integrated Environmental Management. Development, Information, and Education in the Asian- Pacific Region. Y. Itakura, J.S. Eades, F.M. D'ltri, M. Kawashima, S. Endoh, & H. Kitamura. Lewis Publishers. Boca Raton, U.S.A, Meadows, A. & P.S. Meadows (eds) (1999b). The Indus River: Biodiversity, Resources, Humankind. Karachi : Oxford University Press. Ribot, J.C., Magalhaes, A.R., & Pangides, S.S. (Eds.) 1996. Climate Variability, Climate Change and Social Vulnerability in the Semi-arid Tropics. Cambridge University Press. Cambridge. Shackleton, N.J. West, R.G. & Bowen, D.Q. (Eds.) 1988. The Past Three Million Years: Evolution of Climatice Variability in the North Atlantic Region. The Royal Society. London. Watson, R.T., M.C. Zinyowera and R.H. Moss (eds.) (1996) Climate change 1995. Impacts, adaptations, and mitigation of climate change: scientific-technical analyses. Cambridge : Cambridge University Press. 22 Glasgow Naturalist 2002. Volume 24. Part 1. Pages 23-28. COMMUNITY STRUCTURE AND BIOMETRICS IN MICROHABITATIS WITHIN MUSSEL BEDS {MYTILUS EDULIS) FROM INTERTIDAL ENVIRONMENTS AT ARDMORE BAY, CLYDE ESTUARY, SCOTLAND. Noelia Carrasco, Azra Meadows, and Peter S. Meadows Biosedimentology Unit, Graham Kerr Building, Division of Environmental and Evolutionary Biology, Institute of Biomedical and Life Sciences, University of Glasgow, G12 8QQ. ABSTRACT The edible mussel, Mytilus edulis, is very common in the Firth of Clyde and Clyde Estuary, Scotland, living intertidally on rocks and mud where it often forms extensive beds. The beds are bound together by byssus threads secreted by the species. Two packing structures occur in the beds and provide microhabitats for other invertebrate species. These are towers of mussels forming upwards from the general bed to 20 cm, and the rings of mussels forming on the surface of the sediment at the edge of beds. Samples (clumps of mussels) from the towers and rings at an exposed and a sheltered site at Ardmore Bay, Clyde Estuary were collected and analysed in two ways. Firstly, biometric data of the mussels (length, breadth, wet weight) in the samples were recorded. There were no differences between the biometry of the mussels in the towers and rings, or between the sheltered and exposed sites, except that the mussels at the exposed site were slightly broader. This suggests a greater tissue weight, which may be caused by more abundant food, at the exposed site. Secondly, the number of species and number of individuals/species of invertebrates in the sampled clumps of mussels were recorded. Sixteen species were recorded, belonging to the crustaceans, the annelid polychaetes and oligochaetes, and the molluscan bivalves and gastropods. When found outside the mussel clumps, these species are either infaunal, epifaunal, or surface dwelling species able to migrate into the overlying water. This suggests that the mussel clumps provide an unusual mix of microhabitats. Cluster analysis was used to cluster the towers and rings at the exposed and sheltered sites, and to cluster the species. There was a tight clustering of the rings and towers with much less significant clustering of the exposed and sheltered sites. There was no obvious ecological clustering of species by similarity of habitat, but there was an unexpected taxonomic clustering of the molluscan species. The results are discussed in terms of the microhabitats that may exist within the towers and rings, and the need for a comparison of these with microhabitats where the invertebrate species are more usually found. These latter include intertidal sedimentary and rocky environments. INTRODUCTION The coastal zone is an area of dynamic change and spatial heterogeneity. This is particularly marked in the intertidal zone where sedimentary environments are alternately exposed to air and water as the tide falls and rises. Wind, water currents, changes of salinity and fresh water flux, also produce temporal and spatial heterogeneity. This heterogeneity then imposes temporal and spatial heterogeneity on the distribution and abundance of intertidal species. The intertidal zone, therefore, is one that offers a unique opportunity to investigate the some of the central questions in marine ecology about the presence of and factors governing spatial and temporal heterogeneity at different scales (Reise, 1985; Tufail et al., 1989; Meadows et al., 1998; Armonies & Reise, 2000). The west coast of Scotland is particularly rich important here in terms of the range and diversity of intertidal habitats, and it contains some of the finest examples of rocky, sandy and muddy shores in Europe. Spatial heterogeneity is everywhere on these intertidal shores (Meadows & Anderson, 1978). Typical examples are the vertical zonation of organisms on rocky shores, the biodiversity of rock pools, and the different localised communities associated with muddy and sandy shores. Sometimes organisms themselves provide a substrate or habitat for different communities. These are of special interest because they provide examples of animals and plants interacting with other animals and plants in the intertidal zone to produce spatial heterogeneity, rather than distributional heterogeneity being controlled mainly by physical and chemical parameters in the intertidal zone (Fenchel & Reidl, 1970; Anderson & Meadows, 1978; Mckindsay & Bourget, 2000; McQuaid et al., 2000). Well known examples include species of tube worm, such as Spirorbis borealis and Spirorbis spirillum living on selected species of seaweeds, and the red seaweed Polysiphonia lanosa living on the bladder wrack, Ascophyllum nodosum. Mussel beds formed by the edible mussel Mytilus edulis (Field 1922; White, 1937; Theisen, 1968; Mason, 1976) also offer habitats for many species of rock dwelling and sedimentary organisms and these are common in the Clyde estuary and Clyde Sea area. This paper is concerned with the biodiversity of unusual structures formed in some of these beds in the Clyde Estuary by clumps of mussels. Mytilus edulis is a common intertidal species in the Clyde. It occurs naturally here and elsewhere from mid-tide level into the shallow sublittoral, on rocky sandy and muddy substrates, and often forms dense beds (Scott, 1896; Maas Gesteranus, 1942; Kuenen, 1942; Verwey, 1952; Theisen, 1968; Mason, 1976 Hayward 1986). The beds consist of complex multilayered structures produced by the mussels, with density-dependent effects on survival, growth and self-thinning (Meadows & Shand, 1989; Guinez and Castilla 1999). The whole structure is 23 bound together by byssus threads produced by the mussels themselves. We have often observed two characteristically different packing structures of mussels in the beds. The first are towers of mussels forming upwards from the general level of the bed to a height of about 20 cm. The second are rings of mussels that sometimes form on the surface of the sediment at the edge of a bed. The present study, conducted at Ardmore Bay in the Clyde Estuary, compares the biometry of the mussels in the towers and rings, and the biodiversity of the benthic infauna within the towers and rings. Comparisons are made on towers and rings from two sites - a sheltered part of the bay and an exposed part of the bay. MATERIALS AND METHODS The study site was a bay at Ardmore Point in the Clyde Estuary (latitude 55° 58’ 32” N. Longitude 4° 41’ 29” W) (British National Grid Reference NS322792) (Meadows et al.l998). Ardmore Point is a S.S.I. (Sites of Special Scientific Interest) and a Coastal Nature Reserve (Boyd 1986). The field sampling was undertaken during October, November and December 1999 and the frequency of sampling was once a week. Samples were collected at approximately the time of low tide. Two sampling sites were established, an exposed site on the southerly headland of the bay and a sheltered site within the bay on its north-westerly side. Samples consisting of clumps of mussels from towers and rings were taken at the two sites, which were then transported to the laboratory. Two laboratory investigations were conducted on the samples. The first was a biometric study of the length, width, whole weight and shell weight of 30 mussels from the towers and rings at the sheltered and exposed site. Hence 120 mussels in total were measured. This was analysed by ANOVA. The second study consisted of an investigation of the biodiversity of benthic infaunal invertebrates within towers and rings at the exposed and sheltered sites. This involved isolating the invertebrates from the mussel clumps, and then identifying and counting the number of individuals in each species. Cluster analyses were also conducted on the data. This was done in two ways. Eirstly, the towers and rings from the exposed and sheltered sites were used to cluster the species. Secondly, the species were used to cluster the towers and rings at the sheltered and exposed sites. In both cases the data used were the abundance of the species in the towers and rings at the exposed and sheltered sites. RESULTS Biometric analysis of Mytilus edulis in towers and rings from the exposed site and the sheltered site. Statistical analyses by ANOVA of the biometric measurements of Mytilus edulis length, width, and wet weight gave the following results. There was no significant difference in the length, width and wet weight of the mussels between towers and rings at the exposed site and between towers and rings at the sheltered site. There was also no difference between the towers at the sheltered site and the towers at the exposed site, and between the rings at the sheltered site and the rings at the exposed site. There was one statistically significant difference. The length/width ratio of Mytilus edulis forming rings at the exposed site was lower than at the sheltered site (F: 9.246; 0.005>P>0.001; D.F: 1). Community structure of benthic infauna within towers and rings at the exposed and sheltered site. Benthic infaunal invertebrates Table 1 gives a list of the benthic infauna found within the towers and rings at the exposed and sheltered sites. Sixteen species were found in the towers and rings, of which seven species are molluscs including four species of Littorina, five species are Crustacea, three species are polychaetes, and one species is an oligochaete annelid. All the species of molluscs, polychaetes and the annelid are true benthic forms (Table 1). Six of these benthic species are normally considered as infaunal burrowing organisms that either plough through the sediment or construct tubes or burrows within it (Table 1, species labelled A). The species labelled B in table 1 are all epifaunal species which in a normal sedimentary environment live at the sediment-water interface. The species labelled C in table 1 are all Crustacea. These latter species, although living on the sea bed either on sediments on rocks or under stones, are able to make excursions into the overlying water column when covered by the tide. Clustering of rings and towers at the exposed and sheltered sites, and clustering of benthic infaunal invertebrates. The clustering of rings and towers, and of the benthic infaunal invertebrates is shown in table 1 . The lower cluster diagram in figure 1 shows the clustering of the towers and rings and the exposed and sheltered sites. There are three distinct clusters (Figure 1, lower cluster diagram, clusters 1,2, and 3). The clustering indicates that the towers and rings are clustered more than are the sheltered and exposed sites when assessed by the numbers of individuals in the different species living in them. Cluster one contains seven towers and one ring, distributed equally between four exposed sites and four sheltered sites. It can therefore be considered as a tower cluster. Cluster two contains seven rings, five of which are from sheltered sites and two from exposed sites. This cluster can therefore be considered as a ring cluster made up largely of rings in sheltered sites. Cluster three, an outlying cluster, contains two towers both from sheltered sites. The upper cluster diagram in figure 1 shows the clustering of the sixteen benthic invertebrates (table 1 ) isolated from the towers and rings at the exposed and sheltered sites. There are two major clusters (Figure 1, upper cluster diagram, clusters 1 and 2), the first cluster being divided into two smaller clusters (3 and 4 in figure 1), and the second containing a smaller subcluster (5 in figure 1). These clusters are derived from, and hence 24 Table 1. List of invertebrate species found in towers and rings of Mytilus edulis at the exposed and sheltered sites, showing their taxonomic group and habitat. Common names given after scientific names, where known. Normal Habitat = usual habitat that the species is found in, defined thus: A, infaunal benthic invertebrates that either plough through the sediment or construct burrows or tubes in it; B, epifaunal benthic invertebrates that live at the sediment/water interface; C, benthic invertebrates that live on the surface of rocks or sediments, and make regular forays into the overlying water. Species Taxonomic Group Normal Habitat Carcinus maenas (Lineaus) Common shore crab. Crustacea C **Cerastoderma edulis (Linneaus) Edible cockle. Mollusca, Bivalvia A Chaetogammarus marinas Leach Crustacea C **Hediste (Nereis) diversicolor (O.F .Muller) Rag Worm Annelida, Polychaeta A *Hydrobia ulvae (Pennant) Laver Spire Shell Mollusca, Gastropoda B Jaera albifrons Leach Crustacea C **Lanice conchilega (Pallas) Sand Mason Annelida, Polychaeta A *Littorina littorea (Linneaus) Edible Periwinkle Mollusca, Gastropoda B *Littorina mariae Sacchi and Rastelli Mollusca, Gastropoda B *Littorina obtusata (Linneaus) Flat Periwinkle Mollusca, Gastropoda B *Littorina saxatilis (Olivi) Rough Periwinkle Mollusca, Gastropoda B **Macoma balthica (Linneaus) Baltic Tellin Mollusca, Bivalvia A Orchestia gammarellus (Pallas) Crustacea C **Oweniafusiformis (Delle Chiaje) Annelida, Polychaeta A Talitrus saltator (Montagu) Sand Hopper Crustacea C **Tubificoides spp. Annelida, Oligochaeta A considered molluscan clusters. However cluster one is very mixed taxonomically, containing three species of Crustacea, two species of polychaetes, and one species of mollusc. Subclusters three and four within cluster one are also taxonomically mixed. There are two pairs of species that cluster at a very high similarity, and therefore are very closely related in terms of their distribution and abundance between the towers and rings at the exposed and sheltered sites. The first pair consists of the two infaunal molluscs Macoma balthica and Hydrobia ulvae in cluster 2 and subcluster 5. The second pair consists of two very disparate species, the polychaete Hediste diversicolor and the isopod crustacean Jaera albifrons. measured by, the towers and rings and at the exposed and sheltered sites. A careful inspection of the species making up the different clusters shows little commonality amongst the species within each cluster. There is a random distribution between the clusters of the infaunal and epifaunal species and the species that migrate into the water column (Table 1 , species labelled A, B, and C) The clustering is therefore not based on infaunal or epifaunal species ecological requirements. There is some indication of a taxonomic clustering. Cluster two contains seven molluscan and one crustacean species. Within cluster two, subcluster five contains five molluscan species and one crustacean species. So both cluster two and cluster five within it can be 25 Similarity Species Similarity Site ES SEEESSSSSSSE ESS S Towers (T). Rings (R) and Exposed (E), Sheltered fs) Figure 1. TOP CLUSTER DIAGRAM species. Of Owenia fusiformis, Lc Lanica conchilega, Lo Littorina obtusata, Hd Hediste(Neries) diversicolor, Ja Jaera albifrons. Cm Chaetogammarus marinas, Ts Talitrus saltator, Tb Tubificoides bem Mb Macoma balthica, Hu Hydrobia ulvae, Lm Littorina mariae, LI Littorina littorea, Ce Cerastoderma edulis. Cm Carcinus maenas, Ls Littorina saxatilis, Og Orchestia gammarellus. Figure 1. BOTTOM CLUSTER DIAGRAM. Structure (R rings and T towers), and site (S sheltered and E exposed) 26 DISCUSSION There are three aspects of the results reported in this paper that require comment. The first concerns the relative biometries of the mussels collected from the towers and rings at the exposed and sheltered sites. The second concerns the ecological and biological characteristics of the invertebrates found in the towers and rings. The third concerns possible interpretations of the clusters that are shown by the towers and rings and the exposed and sheltered sites, and of the clusters that are shown by the invertebrate species found within the towers and rings. We will consider these in turn. The biometric measurements taken on Mytilus edulis consisted of length, width, and wet weight. The towers and rings exist in different microenvironments, and the exposed and sheltered environments have very different energy regimes in terms of wave power. One might expect, therefore, that factors determining length, width and wet weight might vary considerably between these environments. This does not appear to be the case however, as there were no statistically significant differences in the biometric parameters between Mytilus edulis collected from the towers and rings at the exposed and sheltered sites. There was only one exception to this. The length/width ratio was lower in Mytilus edulis from the rings collected at the exposed site than in Mytilus edulis collected from rings at the sheltered site. This means that at the exposed site the shell of Mytilus edulis was significantly wider when compared with its length than at the sheltered site. In other words, the shells of Mytilus edulis at the exposed site are broader in relation to their length than those at the sheltered site. This implies that the soft parts of the body are larger in the rings at the exposed site. It is possible that this may be associated with a higher food supply and hence energy in terms of the planktonic and detrital material that Mytilus edulis filters and uses as a food source. However it does beg the question as to why the same effect was not apparent between Mytilus edulis from the towers at the exposed site when compared with the towers at the sheltered site. The ecology of the sixteen species of benthic invertebrates found in the towers and rings formed by Mytilus edulis at the exposed and sheltered sites are interesting (Table 1). They are a mixture of infaunal species, epifaunal species and species that are known to move between the seabed and the overlying water column when covered by the tide. The presence of such a wide range of invertebrates within the towers and rings shows that the microhabitat provided by the towers and rings is an interesting one. It is clearly suitable for species that normally live in very different local environments. The towers and rings may have biological, physical and chemical properties that provide a mixture of characteristics shown by the water column, the sediment/water interface, and the surficial sedimentary column below the sediment/water interface. It would be interesting to know what these characteristics are, and to compare them in detail with the same characteristics within mussel beds proper, and with the three more usual habitats. They may include differential water circulation within the towers and rings, localised trapping of sediment, localised differences in redox potential, and changing predator/prey relationships. We suggest, therefore, that microscale variation in habitats might explain the diverse range if invertebrate species found in the towers and rings. In this context would be interesting to compare the overall biodiversity and species composition of invertebrates living in mussel beds with those of the ring and tower structures, and to dissect microscale differences in biodiversity and species composition within single rings and single towers. The clustering of the towers and rings at the exposed and sheltered sites, and of the invertebrate species found in these structures (Figure 1), provides some insight into the ecological phenomena and species interactions and that occur there. It also raises some questions. The cluster analysis is based on a two dimensional matrix of data, consisting of the numbers of individuals in each species in each tower and each ring at the exposed and sheltered sites. The row headings of the matrix are the species, and the column headings are successive towers and rings that have been sampled at the exposed and sheltered sites. The numbers in the body of the matrix are the numbers of individuals in each species. Firstly the species were used to group and hence cluster the towers and rings and the exposed and sheltered sites. This produced the bottom cluster diagram in figure 1. Then the towers and rings at the exposed and sheltered sites were used to group and hence cluster the individual species. This produced the top cluster diagram in figure 1 . The clustering of the towers and rings and the exposed and sheltered sites shown in the bottom cluster diagram in figure 1 indicates a tight grouping of the rings and towers by the abundance of the individual species, with much less influence on the clustering of the exposed and sheltered sites. Specifically, cluster one is a tower cluster and cluster two is a ring cluster (Figure 1, bottom cluster diagram). Hence the relative abundance of the species is correlated closely with and accurately distinguishes between the towers and the rings. This is not so of the exposed and sheltered sites. It is not clear why this difference should be so. One can only conclude - following the argument outlined above, that the micro-environments within the towers are significantly different from the micro environments within the rings, and that these are appreciably greater than any differences imposed on the data by the exposed and the sheltered site - at least as far as the invertebrate species inhabiting them are concerned. A comparison the environmental parameters governing the characteristics of the microhabitats within the towers and within the rings would be very worthwhile. 27 The clustering of the sixteen species (Figure 1 upper cluster diagram) is more difficult to interpret. There is no obvious ecological clustering of species by similarity of habitat (Table 1). The random distribution between the clusters of the infaunal and epifaunal species and of the species that migrate into the water column, is puzzling. This random distribution is emphasised by the unexpected taxonomic clustering. Here, cluster two and subcluster five (Figure 1 upper cluster diagram) are almost entirely made up of molluscs. There is no obvious explanation of this taxonomic clustering, and it becomes more inexplicable in view of the lack of taxonomic similarity in cluster one and subclusters three and four (Figure 1 upper cluster diagram), which contain crustacean, polychaete and mollusc species. Two pairs of species, Macoma balthica and Hydrobia ulvae (within subcluster 4, figure 1 upper cluster diagram), and Hediste diversicolor and Jaera albifrons (within subcluster 5, figure 1 upper cluster diagram) require comment. In both cases the pairs cluster at very high similarity levels. The first pair, Macoma balthica and Hydrobia ulvae, are both molluscs, and both are deposit feeders. In a normal sedimentary environment Macoma balthica lives just below the sediment/water interface and Hydrobia ulvae lives at the sediment/water interface. Both are found at Ardmore Bay in the same part of the upper intertidal sedimentary ecosystem, co-existing in essentially the same habitat. There is considerable similarity, therefore, between their usually observed ecological niches on the shore. The second pair contains two very disparate species taxonomically and ecologically, consisting of the polychaete Hediste diversicolor and the isopod crustacean Jaera albifrons. Jaera albifrons is normally recorded from the upper intertidal region in estuaries, and lives under stones. It is not a species that is normally associated with sedimentary ecosystems, and our own field observations suggest that it is not particularly common at Ardmore Bay. On the other hand, Hediste diversicolor is an infaunal polychaete very characteristic of muddy sediments in estuaries, where it constructs burrows to a depth of 10 cm or more. This species is very common at Ardmore Bay. Clearly, the towers and rings at the exposed and sheltered sites have between them have combinations of characteristics that are specifically required by both pairs of species in different ways. We have no indication of the ecological microhabitats within the towers and rings and how they might select these two pairs of species. It is particularly interesting because the first pair of species are closely related in an ecological sense, while the exact opposite is true of the second pair of species. Here again, further investigation of their microhabitats within the towers and rings would be very worthwhile. REFERENCES Anderson, J.G. & Meadows, P.S. 1978. Microenvironments in marine sediments. Proceedings of the Royal Society of Edinburgh 76B 1-16. Armonies, W. & Reise, K., 2000. Faunal diversity across a sandy shore. Marine Ecology Progress Series, 196, 49- 57. Fenchel, T. & Reidl, R,J, 1970. The sulphide system: a new biotic community underneath the oxidised layer of marine sandy sediments. Marine Biology, 7, 255-268. Field, J.A. 1922. Biology and economic importance of the seal mussel, Mytilus edulis. Bulletin of the Bureau of Eisheries, Washington 38, 127-260. Guinez, R & Castilla, J.C. 1999. A tridimensional self- thinning model for multilayered intertidal mussels. American Naturalist 154, 341-457. Kuenen, D.J. 1942. On the distribution of mussels on the intertidal sand flats near Den Helder. Archs neerl Zool 6, 8-158. Mason, J. 1976. Cultivation. Pp 385-410. In Bayne, B.L. (ed). Marine Mussels. Cambridge University Press. Cambridge UK. Maas Gesteranus, R.A. 1942. On the formation of banks by Mytilus edulis. L. Archs neerl Zool, 6 283-326. Mckindsay, C.W. & Bourget, E. 2000. Explaining mesoscale variation in intertidal mussel community structure. Marine Ecology Progress Series, 205, 155-170. McQuaid, C.D., Linsay, J.R. & Linsay T.L. 2000. Interactive effects of wave exposure and tidal height on population structure of the mussel Perna perna. Marine Biology, 137,925-932. Meadows, A., Meadows, P.S., & McLaughlin. P. 1998. Spatial heterogeneity in an intertidal sedimentary environment and its macrobenthic community. In: Black, K.S., Paterson, D.M. & Cramp, A. (eds) Sedimentary Processes in the Intertidal Zone, Geological Society, London, Special Publications 139, 367-388. Meadows, A., & Shand, P. 1989. Experimental analysis of byssus thread production by Mytilus edulis and Modiolus modiolus in sediments. Marine Biology, 101, 219-226. Reise. K.. 1985. Tidal Plat Ecology. An Experimental Approach to Species Interactions, Springer-Verlag. Berlin Theisen. B.F. 1968. Growth and mortality of culture mussels in the Danish Wadden Sea. Meddr Danm Pisk. ~og. Havunders. (N.S.) 6: 47-78. Scott, A. 1896. Mussels and mussel beds. Reports of the Lancashire Sea-Pishery Laboratories, 58-87. Tufail, A., Meadows, P.S. & McLaughlin. P. 1989. Meso- and micro-scale heterogeneity in benthic community structure and the sedimentary environment on an intertidal muddy-sand beach . Proceedings of the 22'"' European Marine Biology Symposium. Barcelona. Scientia Marina 53: 319-327. Verwey. J. 1952. On the ecology of distribution of cockle and mussel in the Dutch Waddensea. their role in sedimentation and source of their food supply. Archs Neerl Zool. 10, 171-239. White. K.M. 1931 . Mytilus. L.M.B.C. Memoir. 3, 1-177. 28 Glasgow Naturalist 2002. Volume 24. Part 1. Pages 29-33. SCOTTISH INSECT RECORDS FOR 2001 Compiled by E.G. Hancock Zoology Museum, Graham Kerr Building, University of Glasgow, Glasgow, G12 8QQ. INTRODUCTION The aim of these compilations, begun by Christie (1984), is to bring together records of interest that have come to the attention of field workers active in Scotland during each year. In the list, specific names of Lepidoptera and the reference numbers are as in Bradley (1998). As these entries are numbered, following the well-established system initiated by Bradley & Fletcher (1979) family names have been omitted. The new check list of the British Diptera (Chandler, 1998) has been used for that order. Other orders of insects follow the names that are given in Kloet & Hincks (1964, 1977, 1978) with some recent additions and amendments. Much of Scotland still lacks basic faunistic information in its more isolated parts for insects that may elsewhere be unremarkable and so not only ‘rare’ species are treated. ODONATA LIBELLULIDAE Sympetrum striolatum (Charpentier). Common darter. SWT Falls of Clyde reserve. South Lanarkshire, male and female in tandem settled on post of pond-dipping platform (NS882425), Vc77, 01/8/01, SWG. HEMIPTERA CIMICIDAE Cimex lectularius L. Bedbug. In tenement flat, Dumbarton Rd, Glasgow, examples with different instars sent in by local General Practioner (NS5566), Vc77, 1/3/01, EGH. LEPIDOPTERA 15 Hepialus sy Ivina (L.). Orange swift. Lynachlaggan, Insh Marshes, Kingussie, Inverness- shire, mv trap in birchwood (NH8 17017), Vc96, 28/07/01, TB. 18 H. fusconebulosa (DeGeer). Map-winged swift. Dumbarton, in garden (NS386752) Vc99, 1/7/01, K&SF. 56 Stigmella dryadella (Hofmann). Caenlochan Crags, mines in Dryas (N01776), Vc90, 27/7/01, KPB. New vice county record. 64 S. continuella (Stainton). Allt Conait Gorge, single mine on birch, Betula pendula (NN5146), Vc88, 9/8/0 1,JC. 145 Nemophora minimella (Denis & Schiffermiiller). Tailend Moss, adult swept (NT0067), Vc84, 15/7/01, KPB. New to vice county. 154 Heliozela sericella (Haworth). Riechip, mine in oak (NO0647), Vc89, 22/9/01, KPB. New to vice county. 169 Zygaena filipendulae (L.). Six-spot burnet. Ailsa Craig (NX0199), Vc75, 8/8/01 , BZ. 257 Leucoptera orobi (Stainton). Isle of Rum, mines in Trifolium pratense (NG3900), Vcl04, 31/8/00, KPB. The adult emerged this year from mines collected in 2000 and it is included as being new to the vice county. 285 Caloptilia azaleella (Brants). Blackford, adult to light (NT2571), Vc83, 1-2/7/01, KPB. New to vice county. 409a Argyresthia trifasciata Staudinger. Broughty Ferry, Dundee, adults by day (N04631), Vc90, 24/6/01 & 15/7/01, JC. 564 Coleophora obscenella Herrich-Schafer. Bemersyde Hill, cases on Solidago (NT5934), Vc81, 27/10/01 , KPB. New to vice county. 722 Ethmia pyrausta (Pallas). The Caimwell, adult at 810m. alt. (N01278), Vc92, 28/5/01, KPB. New to vice county. 784 Biyotropha galbanella (Zeller). Adderstone Moss, adult on wing (NT5312), Vc80, 2/7/01, KPB. New to vice county. 789 B. domestica (Haworth). Arthur’s Seat, Edinburgh, larvae reared from moss (NT2773), Vc83, 17/2/01, KPB. New to vice county. 822 Scrobipalpa acuminatella (Sircom). Dunhog Hill, larvae in Cirsium palustre (NT4624), Vc80, 1/7/01, KPB. New to vice county. 814 Blastobasis decolorella (Wollaston). Stenton, adults collected (NT6274), Vc82, 28/5/01, AEW (det. KPB). New to vice county. 884 Mompha miscella (Denis & Schiffermiiller). St Mary’s Loch, mines in rockrose (NT2423), Vc79, 26/5/01, KPB. New to vice county. 896 Cosmopteryx orichalcea Stainton. Isle of Rum, reared from mines in Anthoxanthum (NG3503 & NM4099), Vcl04, 27-30/8/00, KPB. This record is confirmed as a Scottish resident by this rearing record, emerging from larvae collected in the previous year. 898 Limnaecia phragmitella Stainton. Duddingston Loch, infesting Typha heads (NT2872), Vc83, 4/7/01; Uphall, in Typha heads (NT0670), Vc84, 9/6/01, KPB. New to vice county. 985 Cacoecimorpha pronubana (Hiibner). Dumbarton, in garden (NS386752) Vc99, 5/8/01, K&SF. An introduced species well established in the south of Britain and spreading slowly into the north of England. It was found as a pupa attached to a Buddleia davidii purchased from a local garden centre. New record for Scotland. 998 Epiphyas postvittana (Walker). Broughty Ferry, Dundee, at light (N04631), Vc90, 10/6/01, JC. An introduced species, originally native to Australia and spreading throughout Britain. This appears to be the first Scottish record. 1030 Earn incanana (Stephens). Killiecrankie Wood, to light (NN9161), Vc88, 14/7/01 & 3/8/01; Ardrostan Wood, at light (NN6923), Vc88, 10/7/01 , JC. 29 1130 Epinotia pygmaeana (Hiibner). Mincrieffe Hill, one adult (N01319), Vc88, 12/5/01, KPB. New to vice county. 1316 Catoptria falsella (Denis & Schiffermiiller). Killiecrankie Wood, to light (NN9161), Vc88, 13/8/01, JC. 1428 Aphomia sociella (L.). Bee moth. Dumbarton, in garden (NS386752) Vc99, 31/7/01, K&SF. 1551 Pieris napi (L.). Green-veined white. Ailsa Craig, male and female seen (NX0199), Vc75, 8/8/01, BZ. 1553 Anthocharis cardamines (L.). Orange tip. Camasserie Farm, Loch Fraphorm (NM8304), Vc98, 6/5/01, RC; nr Oban (NM890300); Glen Lonan(NM9128),Vc98, 12/5/01, JPB; SWT Falls of Clyde reserve. South Lanarkshire, nectaring on cuckoo flower (NS882425), Vc77, 08/05/01 , SWG. 1555 Callophrys rubi (L.). Green hairstreak. South end of Shuna (NM70), Vc98, 13/5/01, DH; road to Sherrifmuir Inn (NN8202), Vc87, 2001, DB; LFplandway, Helensburgh (NS299840), Vc99, 13/5/01 K&SF; Pappert Muir (NS412802), Vc99, 20/5/01, JM. 1597 Inachis io (L.). Peacock. Falls of Clyde reserve. South Lanarkshire, nectaring on daisies and dandelion (NS882425), Vc77, 1/5/01, SWG; Moy Castle, Mull (NM599247), Vcl03, 11/5/01, D&AW; Glasdmm (NN001454), Vc98, 12/5/01, JPB; Craigens, Loch Gruinart, Islay (NR294667), Vcl02, 22/12/01, LK; King's Cave, Arran (NR884310) VclOO, 18/9/01, RT; on A841 roadside, Arran, adults and larvae (NR898313), VclOO, 2/7/01, GW; Fallen Rocks, Laggan, Arran (NR9850, NR9949 & NS0048), VclOO, 28/8/01, RT; Dunchraigaig Cairn (NR9783), Vc98, 31/5/01, RD. 1607 Argynnis aglaja (L.). Dark green fritillary. South Kiloran Bay (NR3998); track to Balnahard Bay, Colonsay (NR4098), Vcl02, 14/7/01, DB. 1614 Pararge aegeria (L.). Speckled wood. Machrie Moor road, Arran (NR9133), VclOO, 22/7/01; fallen Rocks, Laggan, Arran (NR9949) VclOO, 28/8/01, RT. 1618 Erebia aethiops (Esper). Scotch argus. Loch Fyne smokehouse (NN1812). Vc98, 27/7/01, AH; Loch Sloy (NN277089-287089), Vc99, 2001, JM; Daer Reservoir (NS9702), Vc77, 9/7/01 , JW; Ailsa Craig, details to be published separately (NX0199), Vc75, 8/8/01, BZ. 1626 Maniola jurtina. (L.). Meadow brown. SWT Falls of Clyde reserve. South Lanarkshire. Netted in long grass in overgrown meadow. (NS882425), Well, 15/08/01, SWG. 1628 Coenonympha tullia (Muller). Large heath. Reservoir at Scalasaig Hotel, Colonsay (NR3894) Vcl02, 12/7/01, DB. 1629 Aphantopus hyperantus (L.). Ringlet. Waste ground at Airdrie Road, Carluke (NS844510), Vc77, 2-7/7/01, EY; near Forth (NS91 1554), Vc77, 3/9/01, DM ; Daer Reservoir (NS9708) Vc77, 29/07/01 , JW; SWT Falls of Clyde reserve, South Lanarkshire, netted in long grass in overgrown meadow (NS882425), Vc77, 02/07/01, SWG; Balloch Park, Dumbarton (NS390836), Vc99, 2/7/01, K&SF. 1722 Xanthorhoe designata (Hufnagel). Flame carpet. SWT Falls of Clyde reserve. South Lanarkshire, netted flying over dense vegetation in woodland (NS882425), Vc77, 01/8/01 , SWG. 1732 Scotopteryx chenopodiata (Linnaeus). Shaded broad-bar. SWT Falls of Clyde reserve. South Lanarkshire, netted flying over dense vegetation in woodland (NS882425), Vc77, 01/8/01, SWG. 1770 Them cognata (Thunberg). Chestnut- coloured carpet. Insh Marshes, Kingussie, Inverness-shire, mv trap in birchwood (NH817017), Vc96, 28/07/01, TB. Red Data Book status ‘Notable B’. 1801 Perizoma taeniata (Stephens). Barred carpet. Killiecrankie Wood, to light (NN9161), Vc88, 13/8/01, JC. 1866 Carsia sororiata (Hiibner). Manchester treble-bar. Insh Marshes, Kingussie, Inverness- shire, mv trap, herb-rich meadow (NN787996), Vc96, 28/07/01, TB. Red Data Book status ‘Notable B'. 1870 Odezia atrata (L.). Chimney sweeper. Insh Marshes, Kingussie, Inverness-shire, mv trap, herb- rich meadow (NN787996), Vc96, 5/07/01, TB; SWT Falls of Clyde reserve. South Lanarkshire, netted in long grass in overgrown meadow. (NS882425), Vc77, 20/6/01, SWG; Balloch Park, Dumbarton, over 1000 individuals flying together (NS388833), Vc99, 2/7/01, K&SF. 1874 Euchoeca nebulata (Scopoli). Dingy shell. Ardrostan Wood, at light (NN6923), Vc88, 10/7/01 , JC. 1941 Aids repandata (Linnaeus). Mottled beauty. SWT Falls of Clyde reserve. South Lanarkshire, netted flying over dense vegetation in woodland (NS882425), Vc77, 01/8/01, SWG. 1991 Deilephila elpenor (L.). Elephant hawkmoth. Craigmarloch, North Lanarkshire, attracted to house light (NS739757), Vc77, 06/07/01, SWG. 1995 Cemra vinula (L.). Puss moth. Insh Marshes, Kingussie, Inverness-shire, mv trap, herb- rich meadow (NN787996), Vc96, 5/07/01 , TB. 2103 Eugnorisma depimcta (L.). Plain clay. Insh Marshes, Kingussie, Inverness-shire, mv trap, herb- rich meadow (NN787996), Vc96, 28/07/01, TB; Killiecrankie Wood, to light (NN9161), Vc88, 13/8/01 & 6/9/01, JC. Red Data Book status ‘Notable B’. 2116 Paradiarsia sobrina (Duponchel). Cousin German. Insh Marshes, Kingussie, Inverness-shire, MV trap, herb-rich meadow (NN787996), Vc96, 28/07/01, TB. Red Data Book status ‘Notable A’ (BAP prority). 30 2137 Envois occulta (L.). Great brocade. Lynachlaggan, Insh Marshes, Kingussie, Inverness- shire MV trap in birchwood (NH8 17017), Vc96, 17/07/01 , TB. Red Data Book status ‘Notable A’ . 2138 Anaplectoides prasina (Denis & Schiffermiiller). Green Arches. Insh Marshes, Kingussie, Inverness-shire, mv trap, herb-rich meadow (NN787996), Vc96, 25/07/01 , TB. 2150 Folia nebulosa (Hufnagel). Grey arches. Killiecrankie Wood, to light (NN9161), Vc88, 3/8/01, JC. 2162 Papestra biren (Goeze). Glaucous shears. Blackhill Mire, Helensburgh (NS307837), Vc99, 20/5/01, K&SF. 2216 Cucullia umbratica (L.). Shark. Dumbarton, in garden (NS386752) Vc99, 30/6/01, K&SF. 2248 Dryobotodes eremita (Fabr.). Brindled green. Killiecrankie Wood, to light (NN9161), Vc88, 13/8/01, JC. 2313 Enargia paleacea (Esper). Angle-striped sallow. Killiecrankie Wood, to light (NN9161), Vc88, 1/9/01, JC. 2440 Plusia putnami gracilis (Lempke). Lempke’s gold spot. Insh Marshes, Kingussie, Inverness-shire, mv trap, herb-rich meadow (NN787996), Vc96, 28/07/01, TB. HYMENOPTERA SIRICIDAE Urocerus gigas (L.). Giant wood wasp. SWT Falls of Clyde reserve. South Lanarkshire, adult caught in spiders web on stonework of the SWT Visitor Centre (NS882425), Veil, 15/8/01, SWG. COLEOPTERA CARABIDAE Carabus glabratus Paykull. Loch Einich, E. Inverness (NN9198, NN9199), Vc96, 22/6-24/7/01, SB. C. problematicus Herbst. Loch Einich, E. Inverness (NN9198, NN9199), Vc96, 22/6-24/7/01, SB. C. violaceus L. Loch Einich, E. Inverness (NN9198, NN9199), Vc96, 221 6-2411 101, SB. Nebria salina Eairmaire & Laboulbene. Loch Einich, E. Inverness (NN9198, NN9199), Vc96, 22/6-24/7/01, SB. N. rufescens (Strom) (= gyllenhali (Schroeder). Loch Einich, E. Inverness (NN9198), Vc96, 22/6- 24/7/01, SB. Notiophilus aquaticus (L.). Loch Einich, E. Inverness (NN9198), Vc96, 22/6-24/7/01, SB. N. biguttatus (Fabr.). Loch Einich, E. Inverness (NN9198), Vc96, 22/6-24/7/01, SB. N. germinyi Eauvel. Loch Einich, E. Inverness (NN9198), Vc96, 22/6-24/7/01, SB. Clivina fossor (L.). Loch Einich, E. Inverness (NN9198), Vc96, 22/6-24/7/01, SB. Trechus obtusus Erichson. Loch Einich, E. Inverness (NN9198, NN9199), Vc96, 22/6-24/7/01, SB. Patrobus assimilis Chaudoir. Loch Einich, E. Inverness (NN9198, NN9199), Vc96, 22/6-24/7/01, SB. Calathus melanocephalus (L.). Loch Einich, E. Inverness (NN9198), Vc96, 22/6-24/7/0 1 , SB. C. micropterus (Duftschmid). Loch Einich, E. Inverness (NN9198, NN9199), Vc96, 22/6-24/7/01, SB. Pterostichus diligens (Sturm). Loch Einich, E. Inverness (NN9198), Vc96, 22/6-24/7/01, SB. Olisthopus rotundatus (Paykull). Loch Einich, E. Inverness (NN9198, NN9199), Vc96, 22/6-24/7/01, SB. DYTISCIDAE Coelambus confluens (Fabr.). Murton Farm, Forfar, Angus (N04951), Vc90, 16/5/01, GNF. Hydroporus longicornis Sharp. Old fishpond. West Perthshire (NN357283), Vc87, 25/9/01; nr Aricastlich, Glen Orchy (NN242314), Vc98, 9/10/01, GNF. H. rufifrons (Muller). Strath Orchy, Argyll (NN1427), Vc98, 23/6/01, GNF. Oreodytes alpinus (Paykull). Loch Brora (NC852084), Vcl07, 14/7/01, GNF. Recently added to British list (see Hodge & Jones, 1995). Agabus biguttatus (Olivier). Doonbank Cottage stream, Ayrshire (NS3218), Vc75, 30/9/01, GNF. A. chalconatus (Panzer). River Brora, Balnaco (NC794106), Vcl07, 15/7/01, GNF. HYDROPHILIDAE Hydrochus brevis (Herbst). Pool beside Loch Vaa, Moray (NH911 177), Vc95, 14/7/01, GNF. Helophorus granularis (L.). Loch Brora (NC852084), Vcl07, 14/7/01, GNF. HYDRAENIDAE Hydraena rufipes Curtis. Loch Yarrows, Caithness (ND306447), Vcl09, 14/7/01, GNF. STAPHYLINIDAE Carpelimus bilineatus Stephens. Stenness, Orkney, 1 male in remains of long-disused midden (H Y299 1 1 3) , Vc 1 1 1 , 30/5/0 1 , MS . C. pusillus (Gravenhorst). Stenness, Orkney, plentiful in remains of long-disused midden (H Y299 1 1 3) , Vc 1 1 1 , 30/5/0 1 , DL . Philonthus politus (L.). Stenness, Orkney, 1 male in remains of long-disused midden (HY2991 13), Vcl 1 1 , 30/5/01, DL. P. rectangulatus Sharp. Stenness, Orkney, 1 male in hay in disused byre (HY299113), Vcl 11, 30/5/01, MS (det. J.A. Owen). Falagria caesa Erichson. Stenness, Orkney, several in disused midden (HY299113), Vcl 11, 30/5/01, MS. Atheta (Datomicra) celata (Erichson). Stenness, Orkney, abundant in heap of mown grass (HY2991 12), Vcl 1 1 , 30/5/01 , MS. SCIRTIDAE Cyphon pubescens (Fabr.). Loch Baile a' Ghobhainn, Lismore (NM861426), Vc98, 23/6/01, GNF. 31 C. laevipennis Tournier. Wester Duncanstone, Aberdeenshire, pondside vegetation (NJ5626) Vc93, 1/7/01, AWE. C. laevipennis is the now regarded as the correct name for phragmiteticola Nyholm (see Klausnitzer, 1998). C. kongsbergensis Munster. Insh Marshes, Inverness-shire, vegetation in marsh (NH7800), Vc96, 2/7/01, AWE; Rhilochan, Knockarth (NC742071), Vcl07, 15/7/01, GNE. This species added to British list (Hodge & Jones, 1995) since Kloet & Hincks (1977). ELMIDAE Riolus cupreus (Muller). Kilcheran Loch, Lismore (NM829393), Vc98, 23/6/01, GNE. PTINIDAE Mezium affine Boieldieu. Maryhill, Glasgow, in old upholstered furniture (NS5669), Vc77, 12/01, EGH. Trigonogenius globulus Solier. Maryhill, Glasgow, in old upholstered furniture (NS5669), Vc77, 3/01, EGH. CRYPTOPHAGIDAE Atomaria lewisi Reitter. Stenness, Orkney, fairly plentiful in heap of mown grass (HY299112), Vein, 30/5/01, MS. A. apicalis Erichson. Stenness, Orkney, very plentiful in heap of mown grass (HY299112), Vein, 30/5/01, MS. COCCINELLIDAE Coccinella undecimpunctata L. 11 -spot ladybird. Pinbain Bum, nr Girvan, about 20 individuals in company of 2-spot ladybirds (NX138915), Vc75, 23/6/01, K&SF. LATHRIDIIDAE Eniemus histrio Joy & Tomlin. Stenness, Orkney, 2 males and 1 female in heap of mown grass (H Y299 11 2) , Vc 11 1 , 30/5/0 1 , MS . RHIPIPHORIDAE Metoecus paradoxus (L.). Battleby, Redgorton, Perth, 1 male (N00530), Vc88, 11/9/01, IMcG. DERODONTIDAE Laricobius erichsoni Rosenhauer. Castle Fraser, Aberdeenshire, under storey in mixed woodland (NJ724134), Vc92, 6/9/01, AWE. Added to British list by Hammond & Barham (1982). In addition to those in Peacock (1993) there appear to be two earlier unpublished records represented by specimens in the Hunterian Museum (Zoology) insect collections, viz-, on spruce, Penicuik House, Midlothian (NT2458), Vc83, 29/9/86, R.A. Crowson; Stanley, Perthshire, River Tay flood drift (NOl 133), Vc88, 10/2/89. K.H. Lockey. CHRYSOMELIDAE Donacia clavipes (Fabr.). Kilcheran Loch, Lismore (NM829393), Vc98, 23/6/01, GNE. CURCULIONIDAE Otiorhynchus armadillo (Rossi). Dairy, Edinburgh, in open scrubby area (NT237726), Vc83, 22/7/2000, BS. Although this record is for the previous year it warrants mention as being the 3rd British record of an introduced but not established species (det. Max Barclay, Natural History Museum, London). Polydrusus splendidus (Herbst). Dairy, Edinburgh, 2 females on birch foliage (NT237726), Vc83, 22-24/7/2000, BS (det. confirmed by J.A. Owen). New to Scotland. Eubrychius velutus (Beck). Insh Marshes, Inverness-shire, in lochan (NH7800), Vc96, 22/7/01, AWE. DIPTERA DOLICHOPODIDAE Medetera inspissata Collin. Insh Marshes, Kingussie, Inverness-shire, on dead aspen (NH779003), Vc96, 24/06/01, TB. Red Data Book status ' 3'. HYBOTIDAE Tachypeza nubila (Meigen). Insh Marshes, Kingussie, Inverness-shire, on dead aspen (NH779003), Vc96, 15/06/01, TB. SYRPHIDAE Xanthandrus comtus (Harris). Craigiebum, nr Moffat, Dumfries-shire, spmee plantation on hogweed flowerhead (NT125063), Vc72, 14/9/01, KW. A very rare Scottish species with previous records detailed by Watt (2002); British RDB status ‘Notable’. Hammerschmidtia ferruginea (Fallen). Insh Marshes, Kingussie, Inverness-shire, on dead aspen (NH7700), Vc96, 3/07/01, TB; Dulicht, nr Granton-on-Spey, larvae under bark of fallen aspen (NJ0327), 30/4/01, EGH. Red Data Book status ‘ r. Xylota tarda Meigen. Insh Marshes, Kingussie, Inverness-shire on dead aspen (NN776994), Vc96, 4/07/01, TB. Red Data Book status 'Notable'. Eerdinandea cuprea (Scopoli). Inverfarigaig, Inverness-shire on Achillea millefolium (NH522238), Vc96, 10/08/01, TB. Sericomyia silentis (Harris). Inverfarigaig, Inverness-shire On Succisa pratensis (NH522237), Vc96, 10/08/01, TB. Cheilosia illustrata (Harris). Inverfarigaig, Inverness-shire, on Achillea millefolium (NH522238), Vc96, 10/08/01, TB. Leucozona latemaria (Muller). Inverfarigaig, Inverness-shire, on Achillea millefolium (NH522238), Vc96, 10/08/01, TB. Volucella pellucens (L.). Glenborrodale, Ardnamurchan, swept from roadside vegetation at entrance to RSPB car park (NM6061), Vc97, 27/07/01, TB. ULIDIIDAE (=Otitidae s.l.) Homalocephala biumbratum (Wahlberg). Insh Marshes, Kingussie, Inverness-shire, on dead aspen (NN776994), Vc96, 24/06/01, TB. Red Data Book status ' 1'. 32 LONCHAEIDAE Lonchaea hackmani Kovalev. Insh Marshes, Kingussie, Inverness-shire, on dead aspen (NH779033), Vc96, 20/06/01, TB. Added to British list by McGowan & Rotheray (2000). L. fugax Becker. Insh Marshes, Kingussie, Inverness-shire, on dead aspen (NH779003), Vc96, 27/06/01, TB. CLUSIDAE Clusiodes apicalis (Zetterstedt). Insh Marshes, Kingussie, Inverness-shire, on dead aspen (NN776994), Vc96, 28/06/01, TB. Red Data Book status ‘Notable’. CALLIPHORIDAE Cynomya mortuorum (L.). Invertromie Meadow, Kingussie, Inverness-shire, swept from vegetation in herb-rich meadow (NN776996), Vc96, 22/07/01, TB. CONTRIBUTORS Tracey Begg (TB); Shona Blake (SB); Jim Black (JPB); Keith Bland (KPB); David Bowker (DB); R. Campbell (per L. Farrell) (RC); John Clayton (JC); Ruth Dobson (RD); Arthur Ewing (AWE); Stuart Glen (SWG); G.N. Foster (GNF); Keith & Susan Futter (K&SF); Allister Hamilton (AH); Geoff Hancock (EGH); David Hayes (DH); Lawrie King (LK); Derek Lott (DL); Iain McGowan (IMcG); David Mellor (DM); John Mitchell (JM); Bob Saville (BS); Magnus Sinclair (MS); Ruth Thompson (RT); Gordon Watson (GW); Kenneth Watt (KW); David & Anne Welham (D&AW); AE Whittington (AEW); Jim Wilson (JW); Elizabeth Young (EY); Bemie Zonfrillo (BZ). REFERENCES Bradley, J.D. (1998). Checklist of Lepidoptera recorded from the British Isles. Fordingbridge, Hants. Bradley, J.D. & Fletcher, D.S. (1979). A Recorder's Log Book or Label List of British Butterflies and Moths. Curwen Books, London. Chandler. P. J. (1998). Checklists of British Insects (New Series) Diptera, 12. Royal Entomological Society of London, London. Christie, I.C. (1984). Lepidoptera in Strathclyde, 1983. Glasgow Naturalistic, 435-438. Hammond, P.M. & Barham, C.S. (1982). Laricobius erichsoni Rosenhauer (Coleoptera: Derodontidae), a species new to Britain. Entomologist’s Gazette 33, 35-40. Hodge, P.J. & Jones, R.A. (1995). New British beetles species not in Joy’s practical handbook. British Entomological & Natural History Society, Reading. Klausnitzer, B. (1998). Uber die Cyphon- Arien Henri Tourniers. Beitrdge zur Entomologie 48, 411-415. Kloet, G.S. & Hincks, W.D. (1964). A Checklist of British Insects (2nd ed.), part 1, Small Orders and Hemiptera. Royal Entomological Society of London, London. Kloet, G.S. & Hincks, W.D. (1977). A Checklist of British Insects (2nd ed.), part 3, Coleoptera & Strepsiptera. Royal Entomological Society of London, London. Kloet, G.S. & Hincks, W.D. (1978). A Checklist of British Insects (2nd ed.), part 4, Hyrnenoptera. Royal Entomological Society of London, London. McGowan, I & Rotheray, G.E. (2000). New species, additions and possible deletions to British Lonchaea Fallen (Diptera, Lonchaeidae). Dipterists Digest 7, 37- 49. Peacock, E.R. (1993). Adults and larvae of hide, larder and carpet beetles and their relatives (Coleoptera: Dermestidae and of derodontid beetles (Coleoptera: Derodontidae). Handbooks for the Identification of British Insects 5(3), 1-144. Watt, K. (2002). Scotland’s hoverfly recording sheme: Xanthandrus comtus back from the edge of extinction. Hoverfly Newsletter 33, 11. 33 Glasgow Naturalist 2002. Volume 24. Part 1. Pages 35-38. THE NATURAL HISTORY OF THE GLASGOW BOTANIC GARDENS PLANTS GROWING IN A WILD STATE 1998-2001 P. Macpherson 1 5 Lubnaig Road, Glasgow G43 2RY INTRODUCTION During the years 1994-1997 members of the Glasgow Natural History Society visited the Glasgow Botanic Gardens on a regular basis to record the wildlife. The results were published in a series of articles in the Glasgow Naturalist of 1998, including one on the 279 plants considered to be growing “in a wild state” (Macpherson 1998). 1 have continued to record in the Gardens on at least two occasions per year and, in addition, had a number of plants drawn to my attention by Keith Watson and Jean Millar. For recording purposes the Gardens are in Lanarkshire (VC 77). A further 85 flowering plants have been recorded. Of these 30 (35.3%) are regarded as being native and 55 (64.7%) aliens. In the latter category, 16 are presumed to have arrived on site by natural dispersal, 11 to be accidental introductions and 28 to have been planted originally in the Gardens but subsequently spread to another area (Fig. 1). As before, in a number of cases it was difficult to be sure of the status but all have been allocated to that considered to be the most likely, rather than having a doubtful category. This difficulty arose particularly in the Natural Dispersal and Accidental Introduction categories. The lists of plants in the various categories are given as an appendix. The 55 aliens have been subdivided further according to their persistence state: 18 are established, 7 are surviving and 30 of casual occurrence. The definitions used are those recommended by the Botanical Society of the British Isles (Macpherson, et a/. 1996). SITE SURVEYS The previous report highlighted the presence of cotoneasters at the long abandoned railway station near the main entrance to the Gardens. In recent years the area has been surrounded by an impenetrable fence, but looking through, I have not noticed any new species. However, Cotoneaster lomahuensis can be added to the list. It was first seen in 1995 but only identified recently (as sp. nov.). C. divaricatus has now been recorded as well established along a river bank fence. Fallow beds have again been the source of interesting records. Of particular note are Common Amaranth {Amaranthus retroflexus). Rough Bent (Agrostis scabra). Black-bindweed {Fallopia convolvulus), a Bird’s-eye {Gilia clivorum). Scarlet Pimpernel {Anagallis arvensis)- both in its scarlet and blue forms, Bristly-fruited Mallow {Modiola caroliniana) and Lesser Swine-cress {Coronopus didymus) all considered to be native or accidental and Beet {Beta vulgaris si), Red-maids {Calandrinia ciliata), Skunkweed {Navarretia squarrosa), and Vervain {Verbena officinalis) all recorded as having been grown elsewhere in the Gardens. Path sides in this area have been sites for Hoary Cinquefoil {Potentilla argentea) and Trailing Tormentil {Potentilla anglica) presumed to be native or of accidental occurrence and Chamomile {Chamaemelum nobile) which has been grown in the Gardens. At a path side in a more remote area there is a colony of Great Tussock-sedge {Carex paniculata) for which there is no planted record and Flax {Linum usitatissimum) has been seen, presumably the result of birdseed scattered by visitors. On the river bank there is a tiny colony of Water- pepper {Persicaria hydropiper) and a larger one of Bithynian Squill {Scilla bithynica) which presumably migrated from elsewhere downstream to the site. Strictly speaking, it is not actually in the Gardens, but is above the weir which we have taken before to be the southern extremity. Elsewhere there are established colonies of Narcissus taxa. On a rough grassy bank there is a colony of Cyclamen {Cyclamen spp.) and a single plant of Stinking Hellebore {Helleborus foetidus). I have ascertained that C. hederifolium and C. count were planted here in 1996, but not the hellebore. In the neighbourhood of the dump area at the north end of the Garden there have again been some interesting casuals, including Green Alkanet (Pentaglottis sempervirens) and Balm-leaved Figwort {Scrophularia scorodonia). FAMILY RECORDS In the previous report it was noted that one alien and 1 6 native grasses had been recorded in the Gardens. An additional four have been seen: the natives Black Bent {Agrostis gigantea), Hairy- brome {Bromus ramosus) and Tall Fescue {Festuca arundinaceae), and Rough Bent {Agrostis scabra) an accidental introduction from North America to Glasgow, first recorded in 1979 (Macpherson & Stirling 1980). Three extra ferns have been noted: Maidenhair Fern {Adiantum capillus-veneris) and Brittle Bladder - fern {Cystopteris fragilis) on walls, and Krauss’s Clubmoss {Selaginella kraussiana) which had “escaped” from a glasshouse. There are three new daffodil {Narcissus) species or cultivars. There have been no additions to the wood-rush {Luzula) or rush {Juncus) lists, and just one new sedge-Great Tussock-sedge {Carex paniculata). DISCUSSION In the report for the 1994-1997 period, 279 plants were recorded in a ‘wild state’ in the Glasgow Botanic Gardens. Of these 58.4% were regarded as native and 41.6% alien. In the recent survey, a 35 Status of Plants 1998-2001 N=85 Figure 1 . The native and alien status of the plants recorded in ‘a wild state’ in the Glasgow Botanic Gardens 1998-2001. Status of Plants 1994-2001 N=364 Native 53.0% Alien 47.0% Figure 2. The native and alien status of the plants recorded in ‘a wild state’ in the Glasgow Botanic Gardens 1994-2001. 36 further 85 taxa were noted, but, as would be expected during continuous recording in such an area, the proportion of native to alien plants had reduced, there being 35.3% native and 64.7% alien. In both survey periods, in the alien categories, approximately numbers occured in the combined groups arriving either by natural dispersal or accidental introduction and those presumed to have spread from material planted elsewhere in the Gardens. However, whereas previously 70% of aliens were either established or surviving, this has been reduced to 45.5%. Again, in such an area, this can be attributed to the relatively high turnover of hortal casuals. As on the previous occasion, a number of national rarities have been recorded. The combined totals and plant status for the entire 1994-2001 period are given in Figure 2. Native Plants on Site Adiantum capillus-veneris Maidenhair Fern Agrostis gigantea Black Bent Angelica sylvestris Wild Angelica Bromus ramosus Hairy-brome Centaurea nigra Common Knapweed Chrysanthemum segeturn Corn Marigold Cystopteris fragilis Brittle Bladder-fern Cytisus scoparius Broom Epilobium hirsutum Great Willowherb Euphorbia helioscopia Sun Spurge Euphorbia peplus Petty Spurge Festuca arundinaceae Tall Fescue Galeopsis bifida Lesser Hemp-nettle Hypericum x desetangsii Des Etangs’ St John’s-wort Lamium purpureum Red Dead-nettle Papaver dubium Long-headed Poppy Persicaria hydropiper Water-pepper Polygonum arenastrum Equal-leaved Knotgrass Potentilla anglica Trailing Tormentil Potentilla sterilis Barren Strawberry Ranunculus ficaria ssp. bulbilifer Lesser Celandine (bulbous) Rorippa sylvestris Creeping Yellow-cress Solanum dulcamara Bittersweet Stachys x ambigua Hybrid Woundwort Taraxacum hamatum a Dandelion Taraxacum laticordatum a Dandelion Valerianella locusta Common Comsalad Veronica beccabunga Brooklime Vida sepium Bush Vetch Viola tricolor Wild Pansy Natural Dispersal to Site Agrostis scabra Rough Bent Calystegia silvatica Large Bindweed Carex paniculata Great Tussock-sedge Cotoneaster divaricatus a Cotoneaster Cotoneaster lomahuensis a Cotoneaster Crocosmia paniculata Aunt Eliza Helleborus foetidus Stinking Hellebore Lamiastrum galeobdon ssp. montanum ‘Variegatum’ Yellow Archangel cultiver Lotus corniculatus var. sativa Common Bird’s-foot-trefoil variant Mentha x villosa Apple mint Picea abies Norway Spruce Raphanus raphanistrum Wild Radish Reseda luteola Weld Senecio squalidus Oxford Ragwort Scilla bithynica Bithynian Squill Trifolium hybridiim Alsike Clover Accidental Introduction to Site Anagallis arvensis Scarlet Pimpernel Anagallis arvensis Scarlet Pimpernel (blue form) Amaranthus retroflexus Common Amaranth Coronopus didymus Lesser Swine-cress Fallopia convolvulus Black -bindweed Gilia clivorum a Bird’s-eye Linum usitatissimum Flax Macleaya x kewensis Hybrid Plume-poppy Modiola caroliniana Bristly-fruited Mallow Potentilla argentea Hoary Cinquefoil Veronica persica Common Field-speedwell Originally Planted in the Gardens but spread to Site Aucuba japonica Spotted-laurel Beta vulgaris si Beet Calandrinia ciliata Red-maids Chamaemelum nobile Chamomile Chenopodium ficifolium Fig-leaved Goosefoot Eruca vesicaria ssp. sativa Garden Rocket Forsythia suspensa Golden-ball Fuchsia magellanica Fuchsia Heuchera cylindrica a Coralbell Hypericum androsaemum Tutsan Lamiastrum galeobdolon ssp. argentatum Yellow Archangel subspecies Lonicera nitida Wilson’s Honeysuckle Mahonia aquifolium Oregon -grape Navarretia squarrosa Skunkweed Narcissus Div 7 W-W Trumpet Daffodil Narcissus Div 3Y-YO Short-cupped Daffodil Narcissus tazetta Bunch-flowered Daffodil Pastinaca sativa Wild Parsnip Pentaglottis sempervirens Green Alkanet Primula x polyantha Polyanthus Raphanus sativus Garden Radish Ribes alpinum Mountain Currant Scrophularia scorodonia Balm-leaved Figwort Selaginella kraussiana Krauss’s Clubmoss Silybum marianum Milk Thistle Symphytum tuberosum Tuberous Comfrey Tilia x europaea -suckering Lime Verbena officinalis Vervain ACKNOWLEDGEMENTS I am grateful to the following for help with identification: EJ Clement, AA Dudman, J Fryer, DR McKean, AJ Richards, AMcG Stirling & K Watson. Information regarding what had been planted in the Gardens was kindly provided by E Donaldson, J Logan & P Matthews. 37 REFERENCES Macpherson, P (1998). The Natural History of the Glasgow Botanic Gardens. Plants Growing in a Wild State. Glasgow Naturalist 23,44-49. Macpherson, P, Dickson, JH, Ellis, RG, Kent, D & Stace, CA (1996). Plant Status Nomenclature. BSBI News 12, 13-16. Macpherson, P & Stirling, AMcG (1980). Clyde Dock Aliens. Glasgow Naturalist 20, 75-76. 38 Glasgow Naturalist 2002. Volume 24. Part 1. Pages 39-41. COTONEASTERS 1982- 2001 IN LANARKSHIRE P. Macpherson‘ & E.L.S. Lindsay^ 15 Lubnaig Road, Glasgow G43 2RY ‘ 18 Monreith Road, Glasgow G43 2NY ^ INTRODUCTION In a series of articles we have reported on the 26 taxa of cotoneasters known to us to have been recorded in the wild in Lanarkshire (VC 77) (Macpherson & Lindsay 1992, 1993 & 1996). For each taxon we provided a drawing made from live local material. With permission these were reproduced in The Changing Flora of Glasgow (Dickson et al 2000). Since the last report, a further seven species have been identified. In addition to constituting a new vice-county record, each was usually the first record for a much wider area. These occurrences are described in detail and, as before, a drawing provided of a typical leaf in each case. We report also on the gains and loses of the plants commented on in the previous reports. ADDITIONAL SPECIES Cotoneaster ascendens (Fig. 1. a) Despite its name, the single plant of this species, recorded in 1998, hangs from the stonework of a bridge over the River Nethan at Lesmahagow. C. ignescens (Fig. 1. b) A number of strong plants are present on an abandoned railway viaduct at Dalmamock, Glasgow. These were first seen in 2001 and constitute the first record for the plant in the British Isles. C lornahuensis sp. nov. (Fig. 1. c) A single plant is present in the long abandoned railway station at the south east corner of the Glasgow Botanic Gardens. It was first seen in 1995, but not reported by us before as at that time it had not been published, being a species nova. C mairei (Fig, 1. d) The first sighting was that of a single plant at the side of a minor road at Chapelhall in 2000 and a second on the abandoned railway viaduct at Dalmamock in 2001. C marginatus (Fig 1. e) A number of plants grow in relation to a pathside railing at Firhill, Glasgow, where the plant is well established. They were first seen in 1998. C. microphyllm (Fig. 1. f) This name was initially used for plants now regarded as being C. integrifolius. True C. microphyllus was confirmed in 2001 from the Dalmamock viaduct. This constitutes a second British record. C nitens (Fig. 1. g) A single plant, first seen in 1998, grows in roadside scmb at Bothwell. In addition to the above, we report on a colony of strong plants growing on the bank of the River Kelvin adjacent to the railings and bridge on Kelvin Way. It was thought originally that at least one might have been C. henryanus and further material was requested. In 2000 flowering and then fmiting specimens were taken from each plant. The verdict has been that all have a basis of C. frigidus, either alone, or in combination with C. salicifolius (C. x watereri) , and that in one case C. henryanus is involved. UPDATES OF THE TAXA PREVIOUSLY REPORTED The totals given relate to IKm square records over the entire survey period. In a number of cases, there is more than one site within the square. When there are from one to five new records the site locations are given. We consider it appropriate to point out that there are four separate abandoned industrial sites at Shettleston. In a few instances we refer to the Glasgow Rectangle. This is more-or- less Greater Glasgow, and the study area for The Changing Flora of Glasgow. C. adpressus Total 3. In 1986 a plant presumed to be this species was seen on a bing complex, but the site was cleared for development before further material could be obtained for positive identification. There are now two verified records. In 1999 seedlings were seen in an abandoned industrial site at Shettleston, Glasgow, and on waste ground in Dmmpellier Country Park C. astrophoros Total 1. This has not been refound, despite a search in 2001. C. atropurpureus Total 6. Only one of the three plants reported in 1992 is extant, but during 1999 three further records were made: sloping woodland at Glenboig, spontaneous appearance in a garden at Sandford and in abandoned industrial ground at Shettleston. C boisianm Total 1. No change. C. bullatus Total 29. In our last report we noted that there had been 18 records with only two losses. Since then there have been 11 additional sightings. As the plant is so widespread, no attempt has been made to check that the plant is still present at the other old sites. Six of the total have been recorded outwith the Glasgow Rectangle. C. cashmiriensis Total 1. The extermination of the the single plant was previously reported. C. conspicuus Total 8. At the time of the last report, the plant was still present in three of its four sites. At one of these, Kingston, the plant has been eradicated, but we have made four additional records. On grassy waste ground at Cessnock, Glasgow in 1996 (now gone). 39 Figure 1 . Leaves of Cotoneaster species, (a) C. ascendens\ (b) C. ignescens\ (c) C. lomahuensis', (d) C. mairer, (e) C. marginatus; (f) C. microphyllus: (g) C. nitens. barish waste ground at Drumpellier in 1999, abandoned industrial estate at Shettleston in 2000 and scrubby grassland at Forgewood, north of Motherwell in 2001 . C. dammeri Total 4. Of our three previous records, only that on waste ground at Dalmamock is extant and we a±l the occurance on a soil heap at the Bothwell Riding School in 2000. C. dielsianus Total 9. Added to our three previous records are six in such habitats as adjacent to a hedge and in a wood. C. divaricatus Total 5. In 2000 plants were noted growing on the walls of a mined castle at Highouse, north of Biggar, abuntly along a railing by the River Kelvin iind in an abandoned industrial site at Shettleston. These, plus the two previous sites where the plants survive, take the total to five. C. franchetii Total 12. As far as we know, the plant is still present in seven of the eight sites previously reported. Of the four new records, three were seen in 1997: a bing complex at Easter Hessockrigg, a wood at Auchenshuggle, Glasgow and a path side at Provan Hall. The fourth was noted on the viaduct at Dalmamock in 2001 C. frigidus Total 2. No change, the plants survive at both sites. C. helmqvistii Total 4. The additional to the previous records is the occurrence noted in 1988 at a pathside at Stepps. C. horizontalis Total 26. Since the last report we have made 1 1 new records. This is the second most widespread cotoneaster in Lanarkshire, with 10 sites outwith the Rectangle. C. hylmoei Total 2. The strong plant on a rough grassy slope at Laigh Mains, East Kilbride survives, but that at Cessnock has been exterminated by industrial development. C. integrifolius Total 8. Our only previous record was that of a seedling between granite setts at the Custom House Quay. The new occurrences include a plant adjacent to a dockland fence and another with numerous seedlings on semi-bare ground in an old industrial site. C. lacteus Total 1. No change, a plant is still present at the site of the old Botanic Gardens Station. C. rehderi Total 2. The plant previously seen at Eastfield, Rutherglen can no longer be found, although there is no evidence of disturbance to the area. In 1999 a plant was noted adjacent to an abandoned building at the back of the King George V Dock. This is the only record for which we have doubt regarding status— it could have been planted at some time in the past. C. salicifolius Total 18. We know that one of the 12 previously reported has been lost due to development, but have not made an effort to check on the others. The additional six are in habitats such as abandoned industrial sites and a track side. 40 C. sherriffii Total 1. We have been unable to refind this plant, but as it grew in thick undergrowth it could well have been missed. C. simonsii Total 54. With 34 records, this was the most common cotoneaster taxon in our previous report, and the addition of a further 20 keep it in that position. It has the most widespread distribution in the vice-county with 16 sites outwith the Rectangle. C. sternianus Total 2. In our last report we noted that the plants at Cessnock were thriving and increasing in number, but during 2001 they were obliterated by site development. Those in a shrubby wood near the South Rotunda are still present. C. X suecicus Total 8. Of the six previous records, only those at the edge of a wood in the grounds of the Southern General Hospital and on the abandoned viaduct at Dalmamock persist. Of the two new records one is from an abandoned industrial site at Shettleston. With regard to the other, this is the only cotoneaster which we have found in the extreme south of Lanarkshire. A plant was seen in 2000 growing up against a grating which had been placed as a filter across a small burn. The initial thought was that the plant had been washed down and caught by the grating where it took root, but perhaps it is more likely to have been sown by a bird perched on the metal work. In addition to the above, in 1995 the cv. Skegholm was noted adjacent to a roadside wall at Plains, Airdrie. C. villosulus Total 1 . No change. C . X watererii Total 16. Only one of the eight previous records has been lost. Habitats for the additional eight include the edge of a small wood and waste ground. All but one are within the Rectangle. DISCUSSION The vast majority of plants occur alongside railings, parapets or buildings or under trees, indicating spread by bird dropping. It was noted before that this was particularly evident on an abandoned railway viaduct at Dalmamock, where plants were growing at the sides of the parapets. At a re-survey in 2001 , an additional four species were seen in this location. Over a 20 year period we have recorded 33 taxa of cotoneaster in VC 77, a higher total than that given in any of the vice-county floras that we have consulted. This could be the result of a greater number of taxa being grown locally in gardens and/or as amenity planting, or because we have made a practice of sending most of the plants seen for specialist identification. ACKNOWLEDGEMENT We are exceedingly grateful to Jeanette Fryer for identifying the large batches of specimens sent over the years. REFERENCES Dickson, JH, Macpherson, P & Watson, K (2000). The Changing Flora of Glasgow. Edinburgh University Press. Macpherson, P & Lindsay,ELS (1992). Cotoneasters in the Glasgow Section of VC 77. Glasgow Naturalist 22, 111-114. Macpherson, P & Lindsay, ELS (1993). Cotoneaster Update. Glasgow Naturalist 22, 239-242. Macpherson, P & Lindsay, ELS (1996). Cotoneasters Continued. Glasgow Naturalist 23,11-13. 41 42 Glasgow Naturalist 2002. Volume 24. Part 1. Pages 43-46. BARBEL BARBUS BARBUS IN THE RIVER CLYDE: A NEW FISH SPECIES ESTABLISHED IN SCOTLAND Peter S. Maitland' & William W. Miller^ Fish Conservation Centre, Gladshot, Haddington, EH41 4NR ' 7 Brooklands Avenue, Glasgotv, G71 7AT ^ ABSTRACT The Barbel is a new fish species to the Scottish fauna. It is believed that, following earlier attempts at introduction, adult fish were introduced to the River Clyde from England during the early 1990s and are now establishing there, with juvenile fish being caught during 2000. Although adding to the diversity of coarse fish available to Clyde anglers, the presence of the Barbel is likely to be disadvantageous to native fish there, especially Atlantic Salmon which is still re-establishing after a century’s absence. INTRODUCTION Scotland has seen numerous introductions of alien species over the last 150 years and more than one third of the Scottish ichthyofauna is now made up of species introduced from abroad (Adams & Maitland, 2002). The most notable introduction in recent years was the Ruffe Gymnocephalus cernuus, which was introduced to Loch Lomond about 1980 (Maitland et at., 1983; Adams & Maitland, 1998) and is now well established there and in other waters in Scotland (e.g. Loch Ken and the Forth and Clyde Canal). The latest in this series of alien fish is the Barbel Barbus barbus, which appears now to have become established in the River Clyde. One of the earliest fish introductions to the River Clyde was the Grayling Thymallus thymallus, which was first released there in December, 1855, when three dozen fish were placed in the river near Abington. Some 150 years later, the Grayling is well established and distributed throughout most of the river (and much of Scotland from the River Tay southwards) and a species favoured by many anglers (Miller, 1987). There is every reason to expect the Barbel to do equally well in time, although its northern credentials are not as good as those of the Grayling and it may not be so favoured during cooler summers. Judging by its favoured habitats elsewhere, the River Clyde should prove suitable for the Barbel. Like the Rivers Avon and Severn, in which it occurs in England, the Clyde is a large river, with extensive stretches of deep fast flowing water for adults, many suitable areas of gravelly shallows for spawning, and plenty of sheltered slow-flowing backwaters and still pools which are favoured by the young fish. Much of the main River Clyde and many of its main tributaries should prove suitable for this species and, as in other rivers further south into which it has been successfully introduced, the Barbel population is likely soon to expand and occupy these favoured habitats. ORIGIN Full details of Barbel captured so far in the River Clyde are not available, for there is no systematic programme of recording fish catches there. Moreover, some anglers are coy about discussing the new arrival. Thus, as with so many other introductions of fish to Scotland, the details of the original introduction of this species are somewhat hazy. However, from hearsay information, it seems likely that the first Barbel were probably brought up from the River Swale or the River Ure by steel workers at Motherwell who released them in the River Clyde above Motherwell. Subsequently, local reports indicate that several batches, totalling about 50 adult Barbel altogether were transported from England and placed in the River Clyde by coarse anglers during the 1990s. The origin of this stock is believed to be the River Severn. Unlike the introduction of Ruffe Gymnocephalus cernuus to Loch Lomond, which is believed to have resulted from the casual dumping of live bait at the end of a fishing trip, there is little doubt that this transfer of Barbel was a deliberate attempt by anglers to establish the species in the River Clyde. The Barbel, though of no commercial importance in Great Britain is an important species to some anglers, and is respected for its cunning and fighting qualities. Maxwell (1904) noted that ‘Altogether this fish is well equipped with organs of propulsion, which enable him to make a grand fight when hooked.’ Regan (1911) agreed: ‘The Barbel is a strong and active, yet wary, fish, and affords fine sport to the angler.’ Barbel are now caught in the River Clyde on a regular basis, and in the year 2000 the following fish were reported to the authors; (a) A 1 lb (0.45 kg) Barbel caught and returned by someone fishing for Atlantic Salmon just downstream of Blantyre. This fish is much smaller than any of those known to have been stocked and is assumed to be one of their progeny, (b) A ‘large’ Barbel, caught and returned just downstream of Uddingston. (c) A 13.5 lb (6.18 kg) female Barbel, ’full of spawn’, taken by a salmon angler ‘a few miles upstream of Motherwell’, (d) Three large Barbel, 8.25 lb (3.71 kg), 8 lb (3.6 kg) and 5 lb (2.25 kg) caught by coarse fishermen near Motherwell (see Figure 1). These fish were of sufficient interest to feature in the ‘Daily Record’ (1 December, 2000) as ‘Fish of the Week’ . THE BARBEL The Barbel is found in Europe from western France across central Europe to the Black Sea (Maitland, 2000). In the British Isles, it was formerly confined to the east and south-east of England but has been 43 Figure 1 . Three fine Barbel caught on the River Clyde near Motherwell by specialist coarse anglers 44 re-distributed by angling interests to several southern river systems such as the Medway, the Severn and the Bristol Avon. It is absent from Ireland and was from Scotland until the last decade of the 20th century. Adult Barbel are usually some 40-60 cm in length and 1-2 kg in weight, but the species can grow up to 100 cm in length and reach a weight of 8 kg in very favourable waters. The present British rod-caught record is for a fish of 6.237 kg caught in the River Avon in Hampshire in 1962. No Scottish record has yet been ratified. The Barbel is characterised by the possession of two pairs of long fleshy barbels on the equally fleshy upper lip, one pair (the smaller), just in front of the snout, the other at the rear angles of the mouth. The mouth is placed ventrally back from the snout. The body, evolved for active swimming, is long and rounded with very little lateral flattening, although it is rather flat along the belly. The body is covered in medium-sized scales of which there are 55-65 along the lateral line. The Barbel is usually a brownish-green colour on the head and back which grades to a golden-brown on the sides and then to creamy white on the belly. The fins are a dull yellowish-green, sometimes with an orange tinge. The Barbel is a bottom-living fish which occurs usually in the lowland reaches of large clean rivers, where there are stretches of clean gravel and weed beds. Spawning takes place in May and June or even into July when adult fish may move upstream and congregate in large numbers near the spawning grounds, which are over clean gravel and among open weed beds in flowing water. The eggs are yellow, 2-2.5 mm in diameter, sticky, and adhere to weed and to the gravel. Development varies with water temperature, but normally takes 10-15 days. The fry start to feed on small crustaceans and insect larvae and as they grow they move to larger invertebrates including worms, crustaceans, molluscs (both snails and mussels), and mayfly and midge larvae. Large Barbel will take small fish when they can. In good habitats the young fish may reach 10 cm after one year and 15-20 cm after two years. They mature normally at about 4-5 years of age, the males usually about a year earlier than the females and correspondingly often smaller. DISCUSSION Anglers who catch Barbel and are unfamiliar with the species may be in for a surprise, not to mention some potential danger! Firstly, Gordon (1920) mentions of that ‘It is very quick of hearing, and often makes a noise when caught, and growls under the water.’ Secondly, and more seriously, though the flesh is highly regarded in some parts of Europe, the roe (and possibly even the flesh during the spawning period) is poisonous, causing severe stomach disorders. Juliana Berners (1486) gave one of the earliest warnings about eating Barbel: ‘The barbyll is a swete fysshe, but it is a quasy mete, and peryllous for mannys bodye.’ Evidence is given by John Hawkins who, in notes to his edition of The Compleat Angler (Walton, 1760), recorded ’that one of his servants, who had eaten part of a Barbel, but not the roe, was seized with such a violent purging and vomiting as had like to have cost him his life.’ Regan (1911) continued with further warnings ‘... opinions differ as to its value as food, the flesh being white and firm, but rather coarse; the eggs are more or less poisonous, sometimes inducing violent purging and vomiting, and also weakening the heart so much that fainting may result; the poisonous secretion is sometimes absorbed by the flesh of the lower part of the fish, which may thus produce similar effects, and to be safe it is best to eat Barbel only in the late summer and autumn, and to remove the roe as soon as possible after the fish is caught.’ The safest option is offered by Maxwell (1904): ‘Well, and what are you to do with your Barbel when you have got him? That is just the least satisfactory part of the performance. Were Barbel a culinary prize, like Salmon, the sport would be a noble one; but most people account the fish fit for nothing better than to feed pigs withal.’ The realisation that Barbel is established in the main stem of the River Clyde follows closely on the confirmation that the North American Signal Crayfish Pacifastacus leniusculus is also well established in another part of the Clyde system - though this fact only recognised there relatively recently (Maitland et al., 2001). This aggressive predator was first identified by the authors in the River Clyde near Elvanfoot and is now known to be common along several kilometres of the river there. Some 10,000 Signal Crayfish were taken from the river during 2000 and attempts are continuing to contain or eradicate the species. It is unlikely that these will be successful in the long-term, and more likely that it, and the Barbel, will eventually spread to many parts of the River Clyde. Wheeler & Jordan (1990) noted that, in past translocations of Barbel in Great Britain (which have almost always used adult fish), it was normal for the acanthocephalan parasite Pomphorhynchus /aevA to be transferred with it. Thus, though it has not yet been possible for the authors to examine any of the Scottish specimens in detail, it seems likely that this parasite has been brought into Scotland with the Barbel. Such transfers have ignored the advice that any translocations of Barbel should use parasite-free young fish from hatcheries and not the parasitised adult wild stock. As with freshwater invertebrates (Maitland et al., 2001; Maitland & Adams, 2002), this new fish species is likely to continue the scenario identified by Maitland (1987), of changing and unstable fish communities in many parts of the River Clyde. With improving water quality in the lower reaches, native species such as Atlantic Salmon Salmo solar and River Lamprey Lampetra fluviatilis are trying to make a comeback to this river after a century of absence. It is particularly unfortunate, therefore, that alien species such as Barbel and Signal Crayfish (and possibly others, as yet unidentified), likely to 45 act as both competitors and predators, are being introduced at this time. Will the new population of Barbel do any significant damage to native aquatic communities in the River Clyde? This question cannot be answered with certainty. However, there is every likelihood that Barbel will eompete for food and space with native fish speeies such as Brown Trout Salmo trutta. It is also likely to be a predator on the eggs and young of both Atlantie Salmon and Brown Trout, as well as other fish there. Studies on the introdueed Ruffe in Loch Lomond (Adams, 1991; Adams & Maitland, 1998) have clearly shown that this one small species has caused massive ecologieal changes in Loch Lomond and now at least four other alien species are established there. A simpler question is easier to answer. Will the Barbel be a positive or a negative influenee on the native aquatic communities of the River Clyde? Unfortunately, the answer must be negative. Hopefully, the Barbel will be listed among those fish species to be included in forthcoming legislation on controls on the keeping or release of non-native fish in Scotland, at present being prepared by the Scottish Executive. This will parallel existing legislation in England and Wales (The Prohibition of Keeping or Release of Live Fish (Specified Speeies) Order 1998) and will make introductions of non-native fish to Scottish waters illegal, unless carried out under licence. ACKNOWLEDGEMENTS We are very grateful to Thomas McGregor for information regarding Barbel in the River Clyde and to William Weir for photographs and other details of catches. Azra and Peter Meadows made very useful comments on an earlier version of the text. REFERENCES Adams, C.E. (1991). Shift in pike, Esox Indus L., predation pressure following the introduction of mffe, Gymnocephalus cernuus (L.) to Loch Lomond. Journal of Fish Biology 38, 663-667. Adams, C.E. & Maitland, P.S. (1998). The Ruffe population of Loch Lomond, Scotland: Its introduction, population expansion, and interaction with native species. International Association for Great Lakes Research 24, 249-262. Adams, C.E. & Maitland, P.S. (2002). Invasion and establishment of freshwater fish populations in Scotland - the experience of the past and lessons for the future. Glasgow Naturalist 23, 35-43. Berners, J. (1486). A Treatyse of Fysshynge wyth an Angle. London. Gordon, W.J. (1920). Our country’s fishes and how to know them. Simpkin, Marshall, Hamilton & Kent, London. Maitland, P.S. (1987). Fish in the Clyde and Leven systems - a changing scenario. Proceedings of the Institute of Fisheries Management Conference 1987, 13- 20. Maitland, P. S. (1987). Fish introductions and translocations - their impact in the British Isles. Institute of Terrestrial Ecology Symposium 19, 57-65. Maitland, P.S. (2000). Guide to freshwater fish of Britain and Europe. Hamlyn, London. Maitland, P.S. & Adams, C.E. (2002). Alien freshwater invertebrates in Scotland: increased biodiversity or a threat to native species? Glasgow Naturalist 23, 26-34. Maitland. P.S., East, K. & Morris, K.H. (1983). Ruffe Gymnocephalus cernua (L.), new to Scotland, in Loch Lomond. Scottish Naturalist. 1983, 7-9. Maitland, P.S., Sinclair, C. & Doughty, C.R. (2001). The status of freshwater crayfish in Scotland in the year 2000. Glasgow Naturalist 23, 26-32. Maxwell, H.C. (1904). British freshwater fishes. Hutchinson, London. Miller, W. (1987). Grayling with a Scottish accent. Trout and Salmon October 1987, 40-41. Regan, C.T. (1911). The freshwater fishes of the British Isles. Methuen, London. Walton, 1. (1760). The compleat angler. London. Wheeler, A. & Jordan, D.R. (1990). The status of the Barbel, Barbus barbus (L.) (Teleostei, Cyprinidae), in the United Kingdom. Journal of Fish Biology 37, 393-399. 46 Glasgow Naturalist 2002. Volume 24. Part 1 . Pages 47-50 NEST-SITE COMPETITION WITH BLUE TITS AND GREAT TITS AS A POSSIBLE CAUSE OF DECLINES IN WILLOW TIT NUMBERS: OBSERVATIONS IN THE CLYDE AREA James Maxwell 7 Lilac Hill, Hamilton, MLS 7HG INTRODUCTION Willow tits Parus montanus are resident and sedentary, defending their territory throughout the year, and showing a habitat preference for areas of wet woodland (Perrins, 1979). Although the numbers and range of this species across continental Europe are thought to be stable (Hagemeijer & Blair, 1997), the ‘new atlas of breeding birds’ showed the willow tit to have contracted in breeding range in Britain between 1970 and 1990 (Gibbons et al., 1993), with a particularly large reduction in Scotland. By 1990, willow tits nested only in small pockets of the borders, Galloway, Ayrshire and the Clyde, with a single record north of the central lowlands. This contraction in Scotland follows disappearance from Inverness, Ross, Angus, Perth, Fife and the Lothians in earlier decades, which has been tentatively ascribed to factors such as severe winters during the 1940s-50s and loss of habitat (Thom, 1986). Since 1990, declines in numbers and breeding range of the willow tit have continued, and perhaps accelerated, throughout its range in Britain. Willow tit numbers on Common Bird Census plots of the British Trust for Ornithology fell by 56% during 1988-1998, and showed a 29% fall from 1998 to 1999 (Sanderson et al., 2000). Lanarkshire is now the northern limit for the species in Britain, and the neighbouring small isolated populations in Ayrshire and Galloway are in serious decline to very few pairs (Murray, 2000). Winter conditions and habitat loss seem inadequate as explanations for this drastic population contraction. Willow tits are maintaining healthy populations in much colder climates, such as northern Norway, Finland and Russia (Hagemeijer & Blair, 1997), while many areas of apparently suitable breeding habitat in Scotland remain, but now lack willow tits. This study was therefore undertaken to try to find reasons for the decline of the willow tit, and in particular to test whether providing artificial nest sites within suitable breeding areas could help to maintain the remaining pairs in the now tiny and isolated population in Lanarkshire. METHODS Willow tit territories in Lanarkshire were located by listening for the characteristic song and contact calls of this species. Locating territories was enhanced by amplified playback of tape recordings of willow tit vocalisations, which was extremely effective in eliciting responses from willow tits within a distance of as much as 1 km. Once found, pairs of willow tits were visited throughout the year to record their presence and behaviour, especially in the early breeding season. Since there are no marsh tits Parus palustris in the Lanarkshire area, species recognition was simple even when based only on visual observation. Unlike blue tits P. caeruleus and great tits P. major, willow tits only nest in cavities that they excavate for themselves, normally in the rotten stump of a tree. Particular attention was given to the nest construction activities of willow tits in order to glean indications of a suitable design for a willow tit nestbox, and to monitor their breeding activity to see whether the population decline may be related to some cause of breeding failure. The “ nest box “ is basically a bark-covered plastic tube with an internal nest chamber, filled to the entrance hole with fine wood shavings (design details are given in Appendix 1); it was first placed in willow tit territories and used by the birds in 1995 .At present, there are 136 boxes distributed through 22 areas of Lanarkshire ( Figure 1 ) where willow tits have been found holding territory or where they have previously bred. Of these, 7 areas currently hold breeding birds; the species has always easilyjocated these purpose-designed boxes and excavated them readily^ In addition to monitoring breeding success of willow tits in natural sites and in nestboxes, since 1996 the nestlings have been marked with nest- specific colour rings in order to provide information on their post- fledging survival and natal dispersal. A number of adults have also been caught on their territory during winter by mist net and marked with individual colour ring combinations, allowing their survival and pairings to be recorded. RESULTS Habitat, nest sites, nest construction, incubation and fledging. Willow tit territories were located in a range of habitats, from mature woodland with minimal ground cover through regenerating woodland with moderate understorey of brambles etc., to thick birch carr with heather ground cover. Nest sites were found on flat or sloping ground, even on the sides of steep valleys. The preferred decaying timber is birch, with alder another favourite. The average hole-height above ground level in natural sites is 1.5 m, but cavities can be as high as 10.5 m from the ground. During March, willow tits were seen to make trial borings in several stumps, but by April the effort is usually concentrated on one site, with both birds actively engaged in excavation work. The entrance hole dug out is slightly oval (the long axis vertical) and about 3 cm across. Debris falls underneath the site at first, but as the excavation proceeds to form a chamber in a downward direction, the birds turn and fly out with shavings and carry them progressively further away, eventually to up to 20 m. At this stage, birds make up to 4 visits per minute, one bird waiting while the other excavates. This work is carried out in an open, busy fashion and can continue for 47 Figure 1. Locations of willow tit territories investigated on the R. Clyde system during the study from 1994-2001: l=Strathclyde CPark, Motherwell; 2=Calder Glen, East Kilbride; 3=Baron’s Haugh, Motherwell; 4=Chatelherault Cpark, Hamilton; 5=Candermoss, Stonehouse; 6=Garrion Gill foot, Overtown; 7=Mauldslie, Rosebank; 8=Ross Tip, Hamilton; 9=Harestonehill, Newmains; 10=Eastmuir Plan., Newmains; ll=Upper Clyde, New Lanark; 12=Millburn Glen, Larkhall; 13=Merryton, Larkhall; 14=Cleghorn Glen, Lanark; 15=Jock’s Gill, Carluke; 16=Greenhead Wood, Waterloo; 17=Drumpellier Cpark, Coatbridge; 18=Coltness Woods, Coltness; 19=Auchter Water, Newmains; 20=Garrion Gill head. Overtown; 21=Upper Millbum, Ashgill; 22=Canderwood, Stonehouse. several days. When the cavity is complete, the female then builds a minimal nest - a felt-like pad of fine hairs, grasses and fibres. She incubates, and is fed on the nest by the male, but occasionally leaves the eggs unattended, concealing them with ruffled nest material. Both parents share chick- feeding duties and when the young fledge, they are dependent for about a fortnight, then gradually disperse. Use of nest boxes, and breeding data The “nestbox “ (Appendix 1) has proved very successful, and is regularly adopted by willow tits. In 1996, 5 young fledged from a nest box at Strathclyde Country Park and a pair held territory at Baron’s Haugh. In 1997, two pairs bred in the Wishaw and Lanark areas, both in nestboxes, and produced 14 young. Five pairs were located in 1998, producing 28 young. In 1999, two pairs were successful in natural stumps, producing 14 young. In 2000 there were six pairs producing 47 young (two pairs in nestboxes, four in natural sites), and in 2001, three pairs in natural stumps had 17 young. A fourth pair at Coltness used a box to the nesting stage but then deserted (see further note). Sightings of unringed birds hinted at the existence of other breeding pairs in the area but there are probably no more than 8-20 pairs of willow tits now remaining in Lanarkshire . Site fidelity and dispersal A colour ringed male willow tit has bred at the same site at Eastmuir Plantation, Newmains, each year for four successive seasons, but each season 48 with a different female. Young birds given site- specific colour rings were found establishing a territory, on average, only about 1-2 km from their natal site. In Lanarkshire, the main stronghold of willow tit breeding is in the area centering on Newmains, Wishaw and Coltness, with other sites near Larkhall. One fledgling travelled 6 km from Coltness to a site at Baron’s Haugh. From one brood of 9 chicks at Larkhall (in 2000), one moved to Baron’s Haugh (5 km), one to Wishaw (4 km), one to Garrion Gill (3 km) and one moved 2 km, all in different directions. The Garrion Gill female has bred (in 2001) with a previously dispersed male bird from Larkhall (1998) - i.e. this pair shared the same parent and did the same journey, but two years apart. In spite of this kind of consistency in dispersal, many young, including 3 whole broods out of 16, have never been traced, so either dispersed to unknown locations or did not survive. The two Willow Tits that dispersed all the way to Baron’s Haugh were males and apparently did not find females in 2001 and have left that area. Interactions with blue 0 tits At nut feeders during winter, willow tits are subordinate to great tits and blue tits. Similarly, at nest sites, willow tits can be displaced by great tits or blue tits. Observations of willow tits in the Lanarkshire study area showed that these birds frequently lose their newly excavated nest site to a pair of blue or great tits. During five years of study till 2000, of only some 30 willow tit pair-years, I observed 1 8 nest take-overs by blue tits and two by great tits. Of these, 16 were take-overs of newly excavated nestboxes, and four were take-overs of newly excavated natural sites. This probably underestimates the frequency of take-over of natural sites since it was easier to make observations at nestboxes. One example of this interaction was recorded at the territory of a pair of willow tits at Strathclyde Country Park in 1995. This pair excavated five successive nestboxes, losing each in turn to pairs of blue tits. In mid-May, they at last got to the egg stage in a nest-box, only to fail, due to heavy snow. Eventually in late May, their last box was excavated, but, due possibly to energy depletion, only to a depth just under the nest-hole. In the later stages, this inevitably left the young openly vulnerable to predators and the brood was lost ( Nilsson 1984 ). Meanwhile the Blue Tits had raised families successfully in all their acquired sites. In the 2001 breeding season alone, there were 7 Blue Tit and 6 Great Tit take-overs from the nest- boxes. In the afore-mentioned Coltness failure at the nest stage, both Blue Tit and Great Tit pairs were seen showing great interest in the nest, although they did not use the box. It may well be that even at the nest stage, aggression by the dominant tits can be sufficient to make a pair desert, as was witnessed at Jock’s Gill in 1999, when the Willow Tits were witnessed being chased around the nest area before deserting. DISCUSSION In colder countries such as Russia and Finland, young birds usually join adults in adjoining areas and capitalize, in the winter, on food that the older birds have stored (Cramp & Perrins, 1993). In Lanarkshire, storing food seems not to be so vital, and youngsters are often found holding territories on their own in the post-natal year. Willow tits in Lanarkshire seem rarely to join winter mixed flocks of tits, but remain in their territory throughout the year. The dispersal of juveniles from their natal territory to establishing a territory of only a few hundred meters to a few kilometres away is typical of willow tits in Britain; Cramp & Perrins (1993) state that 78% of willow tit ring recoveries came from within 5 km of the ringing location. Only one recorded movement in Britain exceeded 50 km. This would suggest that the Lanarkshire population of willow tits is effectively a closed, isolated population, since the nearest (Ayrshire) population is almost extinct, and the (declining) Galloway and Borders populations are about 100 km away, so are unlikely to provide immigrants to the Lanarkshire area. The specially designed nestboxes were readily accepted by willow tits and good numbers of young have been reared in these boxes. Wired to a tree at knee to shoulder height they look like bark-covered stumps with a tempting bit of decay, which attracts investigative pecking. They satisfy several criteria; being light to carry, cheap to make, waterproof, fairly long-lasting and unobtrusive, so that they are not noticed when in vandal-prone areas. They have also proved invaluable in monitoring the presence of Willow Tits in new areas, as they are always excavated if the species is there. Although it is unlikely that willow tits are limited by a lack of natural sites for nesting, the nestboxes are particularly easy to excavate, and so the energetic cost of cavity construction will be much reduced. This may be important given the problems willow tits face from blue tits and great tits. My observations strongly suggest that an important factor in the decline of the willow tit has been the tendency for blue tits and great tits, having witnessed all the afore-mentioned excavation activity , to take over newly excavated willow tit nest chambers, forcing the willow tits to start a new excavation elsewhere. A series of such thwarted attempts to breed may impair the body condition of adult willow tits, and it is a general feature of birds that late breeding tends to be less successful and fledglings reared late in the season tend to have low survival prospects (Perrins & Birkhead, 1983), (Bromssen and Jansson, 1980). It is likely that blue tit and great tit populations have increased over recent decades as a result of increased provision of winter food in suburban gardens. It is also possible that these species, facing gradually reducing nesting facilities due to the great development of improved housing etc. since the fifties, have expanded into typical willow tit habitat to breed; if so, their liking for new, ready-made sites may be the main cause of 49 the demise of willow tits. This hypothesis invites further research. At the suggestion of Chris Mead (BTO), placing willow tit nest boxes in pairs may help to mitigate this interaction; blue tits take over the first excavated nest box, but permit the willow tits to nest in the adjacent box. Preliminary evidence (in 2000) suggested that this was an effective strategy, but although many nest-box placings were converted to this system in 2001, there was still no significant improvement. Further experimenting along these lines will take place. ACKNOWLEDGEMENTS I am most grateful to the Hamilton branch of the RSPB and Neil Darroch for help in willow tit surveying fieldwork. Matt Mitchell for nest-box work, and Iain Livingstone for ringing the birds. I thank North Lanarkshire Council for their interest and support, the BTO, SNH and B&Q for funding this study, and Bob Furness for generous advice in the preparation of this paper. REFERENCES Bromssen, A. Von & Jansson, C. (1980) Ornis scandinavica 11: 173-178 Cramp, S. & Perrins, C.M. (eds.) (1993). The Birds of the Western Palearctic Vol. VII. Oxford University Press. Gibbons, D.W., Reid, J.B. & Chapman, R.A. (1993). The New Atlas of Breeding Birds in Britain and Ireland: 7988-7997. T& AD Poyser. Hagemeijer, W.J.M. & Blair, M.J. (1997). The EBCC Atlas of European Breeding Birds. T & AD Poyser. Murray, R. (2000). 1998. Scottish Bird Report No. 31. Scottish Ornithologists Club Nilsson, S. G. (1984) Ornis scandinavica 15: 167-175. Perrins, C. M. (1979). British Tits. Collins. Perrins, C.M. & Birkhead, T. R. (1983). Avian Ecology. Blackie. Sanderson, F., Marchant, J. & Glue, D. (2000). Changes in breeding bird populations, 1998-99. BTO News 228, 10-13. Thom,V.M. (1986). Birds in Scotland. T&ADPoyser. APPENDIX 1. Details of the willow tit “ nest box “ design used in this project. The Willow tit nest box is constructed from a plastic tube of 1 1 cm diameter, covered with Cypress bark held in place by chicken mesh. Two tin lids are laid on screws to limit the size of the nest chamber to 22 cm high, with crumpled newspaper used to fill the space above the upper lid. The tube between the two lids is filled with wood shavings. An entrance hole of 3 cm diameter is cut in the tube, 10 cm from the top. The top of the tube is sealed with a plastic cover. The nest box is attached to a tree trunk at about 1-2 m off the ground. Placing nest boxes in pairs, about 3 m apart, seems to be the best way to ensure that willow tits are able to nest in one of the pair of boxes if blue tits or great tits are active in the area. 8cm 30cm 50 Glasgow Naturalist 2002. Volume 24. Part 1. Pages 51-58. THE SCOTTISH LYNX: IS REINTRODUCTION A POSSIBILITY? Katie McDonald Division of Environmental and Evolutionary Biology, Institute of Biomedical and Life Sciences, Graham Kerr Building, University of Glasgow. G12 8QQ. INTRODUCTION The reintroduction of a species must fulfil two criteria to be viable. First it must be feasible from the point of view of all involved - the subject (the species to be returned), the host (the ecosystems into which the subject is to be introduced) and the activators (the conservation body undertaking the reintroduction). Second it must be of benefit to the subject species and/or the host ecosystem. The aim of this review is to discuss the extent to which the reintroduction of the Eurasian lynx to Scotland fulfils these criteria, and thus to consider whether, despite the length of its absence, the lynx has a potential role as the large carnivore of Britain. Before doing this it is necessary to summarise the natural history of lynx and the circumstances under which the species lived in, and was lost from, the UK. THE BACKGROUND Under Article 11 (2) of the European Union’s Convention of European Wildlife and Natural Habitats, contracted parties, of which the British Government is one, should “encourage reintroduction of native species... when this would contribute to the conservation of an endangered species”, conditional on a preliminary study to establish the likelihood of the reintroduction being “effective and acceptable.” (Anon, 1979). Parties are next instructed to “strictly control the introduction of non native species.” In neither the document nor its appendices is a definition of a native species given. What kind of criteria are being used to decide? It is clear from the quote above that contemporary residence is not a prerequisite, but is it a sufficient determinant? Scottish Wildlife Trust (SWT), a major independent conservation organisation in Scotland, advocates complying with these recommendations. SWT does provide definitions of introduction and reintroduction. It is opposed to introduction, which it defines as “the intentional or accidental dispersal by human agency of a living organism outside its historically known native range”. It is however cautiously in favour of reintroduction, which it specifies as - “The intentional movement of an organism into part of its native range from which it has disappeared or become extirpated in historic times as the result of human activities or natural catastrophes,” (SWT, 2000). These statements imply that residence in a country does not make a species native: a country’s native animals are those which colonised independently. What is also apparent is that the return of presently absent native animals is desirable but non native arrivals are not welcome. The modern flora and fauna of the Northern Hemisphere could only begin to establish themselves when temperatures climbed out of the lows of the most recent Ice Age, that is, no earlier than 12,000 - 10,000 bp. The final stadial (glaciation) of this last cold period came to an end about 10,000bp in Europe (Bell & Walker, 1992). The severing of the land bridge with Europe c.9500bp effected Britain’s final isolation from the mainland (Yalden, 1999). To all intents and purposes, colonisation by terrestrial animals ceased, therefore the indigenous land animals of the island must have arrived before this event. THE POTENTIAL INTRODUCTION OF THE LYNX AND OTHER MAMMALS SWT, along with Scottish National Heritage and the Mammal Society, has approved plans for the reintroduction of the European beaver {Castor fiber) to Scotland in 2003 (Kitchener, 2002). Beavers are believed to have last existed in Britain in the fifteenth century (Kitchener & Bonsall, 1997). The beaver will be the only mammal ever to be actively reintroduced to the UK. It is also the only native rodent species to have become extinct in Britain since the modem fauna became established (Kitchener, 2002). All three of our large carnivores, the wolf (Canis lupus), the bear (Ursus arctos) and the less widely known Eurasian lynx {Lynx lynx) were lost in historic times. The last Scottish wolf is recorded as being killed in the 1740s, three centuries after the beaver’s demise but a plan to return wolves to Britain has never come near to the ratification stage. No one can deny that if the beaver is a native of the UK the wolf should be regarded as native also. The situation demonstrates the added cultural and practical complications of reintroduction when the subject is a carnivore. “Nativeness” is by no means the only criterion the species in question can be asked to satisfy. Wolves, and to some extent bears, have received a great deal of negative publicity through the ages. A lynx reintroduction proposal is less likely to encounter opposition from people whose views are prejudiced by folklore. Compared with the much maligned ‘big bad wolf, the lynx is little known and, therefore, a lynx reintroduction proposal is perhaps less likely to encounter opposition before it is even drafted. The Eurasian Lynx, Lynx lynx, belongs to the cat family, Felidae. Felidae is one of the seven 51 families, which make up the order Carnivora. Also included in this order are the family Ursidae, which accepted answer. The major wildlife agencies do not attempt to answer it. Their primary concern when considering a reintroduction proposal is Table 1. The Eurasian Lynx, Lynx lynx. Morphological and Natural History Variables Body length Height to shoulder Weight Dental formula Coat Distinctive morphological features Life span Daily food requirement Gestation Litter no. Breeding frequency 70-130 cm 60-75 cm 12-35kg, average weights: 18.1kg (female), 21. 6kg (male) 13/3, Cl/1, P2/2, Ml/1 Yellow to brown showing seasonal and latitudinal change Short tail with black tip, tufted ears, cheek ruffs, wide spreading paws 17 years in wild 1-2.5 kg 63-73days 1-4 kittens, usually 2 or 3 1 litter per year includes the brown bear, and the Canidae to which the wolf belongs. There is not complete consensus over the lower level systematics of the lynx. The system used here is in agreement with Council of Europe Action Plan (2000). This is to regard the lynx as a genus within the family Eelidae with four species: the Eurasian lynx - Lynx lynx, the Bobcat - L. rufus, the Canadian Lynx - L. canadensis and the Iberian/Spanish Lynx - L. pardinus. L. lynx is often quoted as having several subspecies - the type occupying Northern Europe being L. 1. lynx the boreal lynx. Some authorities include the Spanish lynx as one of these subspecies (L /. pardinus) (Kitchener, 1991). Other variations of the classification scheme include regarding L. pardinus and L. lynx as two of four species in the subgenus Lynx of the genus Felis (Nowark 1999; Corbet, 1991). Although the lynx is not listed as endangered by Convention on International Trade in Endangered Species (CITES), the Council of Europe (CoE) maintains it is in need of conservation as a result of population fragmentation. The once pan-European species now exists in several relatively small isolated units. Small populations are generally felt to be less robust as there is a greater likelihood of any stress causing numbers to fall below the viability threshold. The loss of even one local population is unfortunate, but the more drastic consequence of such population structuring is that the species as a whole is at no less risk than the strongest of its isolated constituents. The CoE regarded the lynx situation as serious enough to merit including the species in “the Big Five” (brown bear, grey wolf, Eurasian lynx, Iberian lynx and the wolverine); predatory mammals who form the subject of their initiative aimed at maintaining and restoring the large carnivores of Europe. It is true that Eurasian lynx become extinct from Scotland over a thousand years before the beaver, but how much importance should be put on this? At what point does the past become recent enough to be relevant? This is a question with no generally whether the species in question is more likely to enhance or to disrupt the ecosystem into which it is placed. They follow the International Union of the Conservation of Nature 1995 Guidelines for Re- introduction (Anon, 1995). These call for assurance that: a) there is a suitable source population which will provide animals of appropriate genetic make up without this source itself being compromised, b) an area of adequate size of suitable habitat and food resources is available, c) there are enough qualified staff to undertake the preliminary work, the actual introduction and subsequent monitoring, and d) the costs of all stages have been assessed. Where a species has been lost from an area in the distant past it must be possible to demonstrate using experience gathered within Europe, “that this species has an integral role in the relevant habitat”. THE BIOLOGY AND ECOLOGY OF THE LYNX The Eurasian Lynx, Lynx lynx, is the biggest of the cats in the European region. Its large paws, thick coat and ear tufts are prominent characteristics of the genus Lynx. Table 1 shows other morphological data. At the full extent of its historical range there were populations in western and eastern European countries, much of the former USSR and as far as Iran and China (Alderton, 1993). This remarkably extensive range, one of the widest known for a cat species, spans many types of terrain. Lynx by preference frequent dense cover, but populations live successfully in a variety of habitats. In Europe they are found mainly in deciduous woodland or pine forest. In central Asia, however, they inhabit open, thinly wooded areas. The most northerly extent of their range includes landscapes of tundra and rocky slopes (Jackson, 1996). Despite obvious morphological adaptation to cold and snowy climates the species is not restricted to high altitudes. In the Carpathians the highest density of the species is found between 700 and 1100 meters, but individuals occur as low as 150 meters. Reports of lynx living at heights of 2000m, 52 demonstrate the animal’s ability to tolerate extreme environments (Bjarvall & Ullstrom, 1986). An animal able to live successfully in a range of habitats must be capable of behavioural adaptability, as well as being physically robust. The wide geographical range of the lynx is more understandable when one appreciates that a description of the lynx ecology contains more preferences than absolutes. This facilitates the species’ wide geographical range Tables 2 and 3 list estimated home ranges in different regions of the overall species’ range. Lynx are classed as nocturnal hunters, although they prefer to rest in the darkest hours and are most active in the hour before nightfall (Jackson, 1996; Burton, 1991). Like other felids, however, they rely on sight and sound when hunting and this practice is ineffective under conditions of poor visibility. Thus, in bad weather the lynx will resort to daytime activity (Kitchener, 1991). Felids, to which the lynx belongs, are arguably the mammal group most highly adapted to meat eating (Gittleman, 1989). For them, as for all dietary specialists, lifestyle - how and where they live - is dominated by the availability of the appropriate food source in sufficient quantities. The other species in the genus lynx are specialist lagomorph hunters and it is often assumed the Eurasian lynx also relies on hare populations. Unlike the Canadian lynx {Lynx canadensis) , which preys almost exclusively on the snowshoe hare throughout the year, there is seasonal and regional variation in the L. lynx diet (Kloor, 1999). The Table 2. Ranges of lynx from various types of terrain. (area in km_) Source Home Range Territory (core area) Male Female Male Female Council 180-27880 98-759 of Europe Action Plan 2000 Council 71-450 45-197 of Europe Action Plan 2001 Wildcats 200-400 100-150 of the World Mammals Few dozen km, of Britain central Europe to and 1000km Scandinavia Europe Cat 264 168 Specialist (n=23) ( n=64) Group lUCN (given as averages) Eurasian lynx is the largest of the species and appears to be most successful when feeding on small ungulates (Jackson, 1996). In the Pan-Alpine conservation strategy for the lynx (Adamic et al, 2001), one section states “many items can be found in the lynx diet” and in another the lynx is described as having a “specialist prey requirement”. This seems contradictory but not when data on lynx populations over the species’ range are brought together. Lynx specialise to the extent and in a manner that their local habitat allows. Populations take as their staple food the most abundant of a range of prey species. The relative abundance of the local ‘favourite’ item and the other potential prey species influences the extent to which the favoured item dominates the diet. In Scandinavia red deer {Cervus elaphus) form the bulk of the diet. In Switzerland chamois {Rupicapra rupicapra) and roe {Capreolus capreolus) constitute 85% of all catches. Roe are also the main prey item in the Carpathians. Although in areas where ungulates are scarce lynx are small and in less than peak condition, there is evidence that populations do exist, in north Finland and Siberia for example, where they have to live on a diet almost completely made up of lagomorphs (Jackson, 1996). A specialized meat eater cannot supplement animal food with non-meat items, like berries and roots, a choice available to omnivorous carnivores such as the bears. A single lynx requires on average 1kg to 2.5kg of meat per day. Survival depends on getting access to a specific resource that requires substantial energy output to obtain. Lynx do this by defending a territory in which they have exclusive hunting rights. The result is a solitary existence. The only association that takes place among adult lynx occurs between males and females during mating periods. Two females never hold the same territory, although the territories of males can overlap. Male territories are larger than those of females and can encompass parts of several females’ territories. The absolute range of both sexes depends greatly on the productivity of the land on which they are residing. Where ungulates are at a high density, ranges are small. As ungulate density decreases ranges have to increase in size for a lynx to have access to sufficient resources. (Breitenmoser, 1998) (Table 2). Two points should be noted about the terminology used in Table 2. Firstly the home area is the area usually around the domicile over which an animal travels in search of food. Secondly, the territory applies to the area within the home range occupied more or less exclusively by an animal or group of animals of the same species and held through overt defence, display or advertisement (Burton, 1979; Bjarvall & Ullstrom, 1986; Kitchener, 1991; Alderton, 1993; Breitenmoser et al., 2000). 53 HISTORY OF THE LYNX IN SCOTLAND There is a dearth of information about former British lynx populations. As with all prehistorically extinct species, period of residence can only be estimated. The oldest fossil find tells us only that the animal existed at this date. The most recent find indicates no more than that the animal was still present at this time. The palaeontological record for Scotland in the Quaternary is not good. Ice covered the country during much of the Pleistocene and there is a shortage of suitable cave sites. The high acidity of the soil is another factor which mediates against fossilisation (Kitchener & Bonsall, 1997). Only one of the meagre number of lynx fossils found in the UK comes from a Scottish site (Kitchener, 1983; Yalden, 1999). Many British specimens come from sites of 18th and early 19th century excavations so were processed before expertise in recording fossil sequences and handling finds were well developed (Jenkinson, 1983; Kitchener & Bonsall, 1997). Most literature has dated lynx fossil finds by associated species with more complete records, and by the presence of artefacts associated with particular prehistoric cultures (Yalden, 1999). It has been suggested that lynx were in the UK during the last cold stage (Guggesberg, 1975, quoted in Yalden, 1999). There is little fossil evidence for this. Of the fifteen British finds only four come from sites with pre-Flandrian (<10,000 year ago) sequences. At only one of these did Jenkinson (1983) in his review, regard the occurrence of lynx remains as a genuine indication of the species living contemporaneously with the other creatures present, rather than an artefact of geomorphological mixing or bad handling of recovery operations. Yalden (1999) suggested a postglacial arrival. Roe deer have a reasonable fossil record and some remains can be dated (pollen and carbon 14 dating techniques). They are believed to have entered Britain in the Mesolithic (c.l0000-5000bp). It is likely that the lynx would have appeared around this time, obviously before the opening of the English Channel (Yalden, 1999). Sizable lynx populations prior to colonisation by small ruminants are known to have existed only in three areas, all in the northern part of the felid’s range (Hedmark, Finland and Ural Mountains) (Breitenmoser et al. 2000). The latest reliable record, according to Jenkinson (1983), comes from Steely Cave, Yorkshire, where a Mesolithic artefact was found and the pollen sequence matched that of zone Vila (c. 7000- SOOObp). Subsequently, radiocarbon dating has been performed on a partial skeleton from Reindeer Cave, Inchnadamff, Sutherland, (formerly believed to be no more recent than pollen zone HI - c. 1 1,000-10,000 bp.). It was dated at cl880bp, indication of the species’ survival in Scotland at least 3200 years after pollen zone Vila. The above section on lynx ecology emphasised the animal’s ability to adapt its feeding strategy, range and diet depending on its circumstances. The precise nature of the previous British population must be assumed to depend on when it was resident, because the environment was changing over time. The late Devensian saw a rise in temperature above present day levels throughout Europe, but at the very end of the Pleistocene temperature fell once more (1 1,000-10, OOObp). Ever since rapid climatic amelioration terminated the final stacial, (c. 1 0, OOObp), European temperatures have oscillated within a range rarely greater than 1-2° above or below present levels. Mid-July temperatures in late glacial were 7-9°. Mesolithic average July temperature was 15° (Bell and Walker, 1992). The warming considerably altered the British landscape through its long-term effect on the vegetation. Pollen records show a trend of increasing forestation through most of the Mesolithic (cl0,000-5000bp) (Price, 1983). The woodland matured and its constituents altered partly a result of adapting to fluctuating precipitation levels, which remained dynamic despite temperature stability. Table 3. Densities of lynx in different habitats. Area Poland Switzerland, Jura mountains Switzerland, North Alps Switzerland, central Alps South Norway Sweden Bialowieza Russia Density (individuals/lOOkm) 1. 9-3.2 0.94 1.2 0.34-0.74 10-19 The modern habitat most similar to the British lowlands in the first half of the Flandrean is thought to be that of the Bialowieza forest, Poland (Yalden, 1999). Figures show Bialowieza to have the highest densities of lynx so far recorded (Cat Specialist Group web site). Table 3 shows how density varies from area to area. This is likely to be an indication of near optimum conditions. The 580km^ forest provides a habitat of wide rivers, reed beds and lagoons. The rich landscape is able to support about 3700 red deer and 2700 roe, ample prey for the 19 lynx and around equal numbers of wolves believed to reside in Bialowieza (Yalden, 1999). Yalden (1999) calculated that at this density the whole of the UK (230,367km^) could have 54 supported 7543 lynx. If he is correct Mesolithic British lynx were certainly not dwindling. A British lynx extinction during the Mesolithic in Pollen zone Vila does not have an obvious cause. Climate change alone does not seem a likely explanation (Yalden, 1999). The range of the Eurasian lynx spans an area over which the geographical temperature range is greater than the temporal variation Britain is thought to have experienced during the Mesolithic. It is likely the species would have had the capacity to cope with the altering temperature and weather conditions (Kitchener & Bonsall, 1997; Yalden, 1999). The much more recent history of lynx decline on the continent gives clues as to the factors which are likely to undermine the species. During the eighteenth and nineteenth centuries deforestation to make way for an expanding agricultural industry drastically reduced the amount of natural woodland on the continent of Europe. The increasing human population overexploited what remained. This resulted in severe degradation of the lynx’s preferred habitat. Since the lynx uses an ambush hunting technique, the percentage of attacks that are successful is highest in areas of dense forest (Breitenmoser, 1998). Deforestation has also directly influenced the contemporary fall in wild ungulates numbers. (Breitenmoser, 2 0 0 0; Breitenmoser, 1998) Hungry predatory animals resorted to supplementing their diet with ungulates from the increasing domestic herds, thus farmers were added to gamekeepers and fur trappers as direct persecutors of the lynx. Drastic reduction of woodland took place in Britain centuries before it became a major issue on the European mainland. Jenkinson’s review reports of “woodland clearance by human groups” beginning at the end of the Mesolithic (Jenkinson, 1983). This substantiates his belief of a Mesolithic extinction of the lynx. However, Bell & Walker, (1992) while agreeing that clearance of woodland did take place in the late stages of the Mesolithic stated that the effect of hunter gathering societies on the landscape is limited to a local level. At the very end of the Mesolithic the lifestyle of humans in Britain changed fairly abruptly from hunter to farmer. They then began to have a significant impact on the landscape (Bell & Walker, 1992). It is plasticity of lifestyle which allows the lynx to survive in areas less than ideally suited to its ecology. If the productivity of the habitat declines, the territory size of each lynx must increase if the animals are to survive. This will cause the population density to fall and eventually the population will become extinct. In prehistoric Britain the lynx would have been put under additional pressure as those animals on British soil turned into island populations, limiting the area available for expansion. Yalden (1999) suggests that the strain, when combined with hunting pressure, could have led to the lynx’s disappearance from Britain. Thus it appears the survival of British lynx into the Neolithic is more understandable than its extinction before the end of the Mesolithic. The latest data on the Inchmadamph skeleton suggest the lynx’s time of residence could have extended into ‘historic’ times. This discovery has important implications for the debate on the wisdom of lynx reintroduction to Scotland. The time between the Neolithic and the Roman occupation was when the characteristic fauna of modern Britain developed. By the time of the Roman invasion, human clearance had transformed the island into what was essentially an agricultural landscape (Yalden, 1999). DISCUSSION In the introduction to this article I have stated my view of the requirements for a successful reintroduction. I then reviewed the biology and ecology of the lynx, and described the history of the lynx in Scotland. I now discuss the potential introduction of lynx to Scotland in more detail, and evaluate the extent to which founding a lynx population in Scotland fulfils the requirements and the Council of Europe’s Action plan. I have divided the discussion into two parts - feasibility of reintroduction, and benefits of reintroduction. Feasibility Had climate change played a major role in the downfall of the British lynx population, reintroduction would be inappropriate and, most likely, impossible. The latest evidence, however, suggests lynx survived in Scotland into Roman times. This increases the legitimacy of arguments for the species’ reintroduction to Northern Scotland because it implies that maintaining a lynx population in modem Britain could be ecologically viable and there could be a niche in our ecosystem for these cats. Something other than unsuitable natural habitat must therefore have been responsible for the loss of the species from Britain. One recommendation of Council of Europe’s 2000 action plan is that “The historical decline of the lynx should be analysed, the threats to the population identified and measures to remove limiting factors taken.” (Breitenmoser, 2000). In the case of the British lynx this would be a more complex task, since the period under investigation would be the ancient rather than the recent past. The century, and therefore the environmental conditions, in which the lynx perished are rather uncertain. Investigation of the history of the lynx in Britain is a crucial part of piecing together the circumstances of its demise, but one cannot simply extrapolate from this to the steps required to allow its return. Examination of the way in which modern day lynx are faring in various types of habitats and the success or otherwise of management and conservation can provide some information. The European Lynx lynx population reached an all time low on the continent in the 1950s and ‘60s. At this time it was extinct from south, west and northern countries, remaining only in Fennoscandia (Norway, Sweden, Finland), the Eastern 55 Carpathians and White Russia (present day Belarus), (Bjarvall & Ullstrom, 1986). Lynx reintroduction programmes began in the 1970s but conditions had begun to recover before this (Breitenmoser, 1998). By the end of the nineteenth century authorities had started to appreciate the environmental importance of forest regeneration (Breitenmoser, 1998). As conditions improved ungulate numbers recovered. The lynx was able to recolonise parts of its former range. Recovery was aided in many countries by the introduction of legal protection (Breitenmoser et al. 2000). Alderton (1993) noted that only three of nine reintroductions appeared to have been viable in the long term, two Swiss projects (Dinaric population, Alpine population) and the Slovenian (Carpathians) programme Successful introductions have been in areas of dense forest and high ungulate density. Of the ten recognised European populations only two do not cross national boundaries. Pan-European collaboration on the regeneration of Europe’s carnivores means lynx can move freely over vast tracts of land. The forests of Britain were decimated long before their importance was appreciated and earlier than those in most mainland European countries. Serious attempts at reversal began later. All the evidence suggests that successful reintroduction requires forestation or other suitable cover over a sufficient area and, certainly in the early stages, legal protection and population monitoring. Even if Britain were to sign up to the Action Plan of the Council of Europe a UK population would remain insular. The importance of this is made apparent by the knowledge that some small countries in mainland Europe do not have sufficient area to support an independent population. In addition, where reintroduction has been tried in areas where natural recolonisation is not a possibility, the amount of work required is even greater and success is even less certain (Breitenmoser et al., 2000). Rural Britain is does not lack areas where the concentration of deer should be adequate to support a reasonable number of lynx (Yalden, 1999). Suitably sized sites of dense, upland forest are more limited. Scotland is better endowed with appropriate expanses than England. More effort would have to be put into reforestation, specifically expanding existing woodland sites, if a population of lynx in Scotland was to have a chance of becoming ecologically feasible. The next issue is how well such a population could coexist with the resident Scottish biota. Of particular importance is whether the introduction of a large predator would lead to competition with the present Scottish carnivores, and how our deer populations would respond to a serious wild hunter. The member of our fauna most closely related to the lynx is the Scottish wild cat (Felis silvestris grampia). It might be expected therefore that wild cats would be the indigenous carnivores put at greatest risk by a lynx return. Wildcats are protected under a schedule of the Wildlife and Countryside Act 1981. Any venture that could threaten them is likely to meet with strong opposition from many conservationists. Although wildcats inhabit most of the same general parts of the continent as lynx, they are by and large at lower altitudes (<500m) and catch only smaller prey items (Corbet and Harris, 1991). None of the literature cited here on lynx reintroduction mentions significant competition between the two species. It seems therefore, the Scottish wildcat and the lynx could coexist as part of a carnivore guild, provided that deer did not become scarce. There is no consensus on either the extent or the nature of the lynx’s effect on wild ungulates. In a review of the lynx in Switzerland, Breitenmoser (1998) stated that on fairly barren territory lynx home ranges are large and not a major factor in determining the number of small ungulates: where prey is plentiful lynx hunting in a concentrated area can significantly shrink populations of deer at a local level. Other reports, cited in the Council of Europe’s Action Plan for the Lynx, (2000), suggested that it is at the margins of ungulate ranges where population is least dense that lynx predation most greatly disrupts ungulates. It appears that the results of predation are heavily dependent on the structure of the ungulate population, but the relationship is not straightforward and other factors are also involved (Breitenmoser et al., 2000). The human species is itself a member of an area’s biota. The manner in which the people of Britain would receive the lynx is hard to determine. Ultimately, it could tip the balance between the success or failure of the venture. On the continent the anthropogenic difficulties faced by large predators are most acute where they are reintroduced into areas from which they have long been absent. The largest difficulties are where sheep are left to graze unattended (Breitenmoser, 1998). This is the practice in Britain. Presenting humans as ‘just’ another member of the biota is misleading. Apart from the magnitude of our influence, the most important feature is that our perception of the effect another creature has on us is as influential as the effect itself Definite evidence that lynx in Scotland would threaten the welfare or economic interests of the Scottish people is scarce: the threat which human hostility could pose to the successful reintroduction of the species to Scotland is very real. The first stage of any reintroduction must be a feasibility study (Breitenmoser et al., 2000). Certain conditions are made apparent by the above. A lynx population would have to be confined to a designated area which must contain sufficient resources to sustain the population without the area being compromised in any way. The site would have to be located so that conflict between people and the lynx was minimised and road casualty, a major cause of death of reintroduced lynx. 56 (Jackson, 1996; Council of Europe, 2000) avoided as far as possible. If this information can be simplified into a set of fundamental requirements computer modelling techniques can be used to provide information which helps in the planning the venture. Geographic Information System (GIS) is an analysis technique which can determine the number of sites, in designated size categories, available in the study area (Scotland) and how many animals each could support. If supplied with the necessary biological variables PVA (population viability analysis) programmes allow an estimation of the likely number of release animals required if there is to be reasonable certainty of the population surviving on average for a specified length of time A 95% chance of survival for at least 50 years was the level used by Leaper et al. (1999). The lynx is not an endangered species but, as has been mentioned, many populations in Europe are felt to be vulnerable. The source animals for the reintroduction of a British population would need to be taken from an area where density was reasonable and not in decline. With reintroduction of a long absent species there is not, as there is with population supplementation programmes, the danger of contaminating an indigenous subspecies with another subspecies. However, it would be judicious to use source animals from an area with similar ecology to the terrain being repopulated. Much depends on the value organisations such as SWT would place on returning the lynx to Britain and how it would rate in importance relative to their other commitments. Benefits A primary purpose of the CoE in reintroducing lynx to places they formerly inhabited is the establishment of viable populations or sub-populations of viable metapopulations. Returning the lynx in Scotland would have very little impact on the success or failure of the redevelopment of a pan-European population. So such an action would be of negligible benefit to the species as a whole If one’s only concern were the welfare of the species, returning it to Scotland would be a waste of human and financial resources. Given the potential disruption to the source population, it could be seen as counterproductive. As a component of an action plan with the aim of improving the host ecosystem, however, it could be extremely useful. The pattern in Britain, common to most areas of significant human colonisation, has been to increase ungulates numbers, which are a source of food and a focus of sport. The large carnivores, potential competitors for game, are persecuted. The natural equilibrium is destroyed and not allowed to redevelop. The red deer population now numbers c. 300 ,000. Their browsing causes economic and environmental damage, and the food requirement of the increasing population is beyond the capacity of the available habitat (SNH web site). Insufficient nutrition can compromise the health of all age groups in the herds. Culling measures have so far been insufficient to halt the escalation. The presence of lynx is not suggested as a complete solution, but reintroduction of a natural predator is a measure so far untried, which could assist deer regulation. Lynx have had much less effect on sheep populations than either bears or wolves in Europe and they never attack cattle (Breitenmoser, 1998). They are the most likely of the three to remain in remote areas and thus least likely to inhabit areas close to human settlement. On the continent they are not even perceived as ‘man eaters’, which both the bear and the wolf often are (Yalden, 1999). A proposal has been put forward for setting up a lynx population on the Island of Rum (Nevard & Penfold, 1978). Manufacturing an artificial community on an island without requiring attitudes to alter would not greatly broaden our understanding of the changes that would be required to make the countryside of the Scottish mainland suitable for a functional lynx population. Large predators are “keystone species’. They are the top of the food chain so their survival depends on the lower level of the chain functioning well. Investigating the viability of establishing a modern day British lynx population, even if the conclusion was not in the affirmative, would be a worthwhile exercise. The presence of a healthy lynx population is indicative of a healthy ecosystem. Aiming to create ‘a lynx friendly environment’, however slight the chances of complete success, would benefit a substantial proportion of the resident Scottish biota. CONCLUSION The lynx would not enter the minds of most British people should they ever be asked to name the creatures of the country’s wilder past. Yet the Eurasian lynx is indeed native to Britain, in the sense that the species colonised this island independently during the Mesolithic, before Britain became separated from Europe (c9500bp). Recent work done on the Inchmadamph skeleton dated it at cl880bp and suggests the species survived in Scotland into historic times - post the Roman invasion of Scotland (c.43 bp). With this knowledge it seems more likely that the reintroduction of the lynx to Scotland would conform to lUCN criteria designed to ensure projects do not undermine donor populations or host ecosystems. Its ability to hunt in different ways, combined with a tough physique, allowed the lynx to become one of the most widespread of the Felidae. These same characteristics make it quite possible that the species could find food and survive in the Scottish climate. There is no resident native British carnivore that would be at risk of intense competition for resources with the lynx. Even wild cats and foxes, the largest of our present carnivores, are much smaller and foxes are considerably more inclined to be omnivorous. 57 Interested parties are not in agreement over whether lynx are likely to prey intensively on livestock. Evidence does suggest that of the three possible candidates for reintroduction of a large carnivore to Britain, wolf, bear and lynx, the lynx is the least liable to cause a serious problems to farmers. The conclusion that lynx reintroduction is theoretically feasible is not in itself justification for undertaking a scheme. A major objective of CoE lynx initiatives is the establishment of populations that contribute to the stability of metapopulations. Returning the lynx to Scotland would not significantly further this aim. It would create yet another discrete population, and in this case, there would not be the possibility of eventual amalgamation with populations in neighbouring countries. Unlike the projects on the continent, the primary benefit of restoring a Scottish lynx would be for the existing fauna of the host country rather than for the lynx itself. We live in a country where forests and wild areas were removed to make way for agriculture. The wild animals have all been affected to a greater or lesser extent by habitat destruction. In addition game species have been encouraged, resulting in great rises in numbers of individuals. Meanwhile predators have suffered persecution and the outcome is the disappearance of all large carnivores. An investigation into the adjustments in land management necessary to enable a reintroduced population of lynx would highlight problems that directly and indirectly compromise many of our indigenous species. The first is whether sufficient suitable habitat exists, or could be created, in Scotland to sustain a viable population. The second issue is anthropogenic difficulties, such as the widespread perception that the introduction of a large carnivore will inevitably cause major disruption. With the present climate being relatively favourable to the general idea of reintroduction it would seem an appropriate time to investigate seriously the return of the lynx to Scotland. Bearing in mind the potential problems mentioned above, however, it is crucial that a feasibility study puts emphasis on determining how many lynx could be expected to live in the country, and whether it would be possible for British people to coexist with a major predator population without the much feared side effects. REFERENCES Adamic, M., Breitenmoser-Wursten, C., Breitenmoser, U., Easel, M., Huber, T., Kaczensky, P., Koren, I., Molinari, P., Molinari-Jobin, A., Rotelli, L., Stahl, P., Stanisa, M., Vandel, J. and Wolfl, M. 2001. Pan-Alpine Conservation Strategy for the Lynx. Convention on the Conservation of European Wildlife and Natural Habitats. Council of Europe, Strasbourg. Anon. 1979. Convention on the conservation of European Wildlife and natural habitats. 19.IX.1979, European Treaty Series/104, Bern. Alderton, D. 1993. Wild Cats of the World. Blandford, London. Bell, M. and Walker, M.J.C. 1992. Late Quaternary Environmental Change: Physical and Human Perspectives. Longman, Harlow. Bjarvall, A and Ullstrom, S. 1986. The Mammals of Britain and Europe. Croom Helm, London. Breitenmoser, U. 1998. Large predators in the Alps: the fall and rise of man’s competitors. Biological Conservation 83, 279-289. Breitenmoser, U., Breitenmoser-Wursten, Ch., Okarma, H., Kaphegyi, T., Kaphegyi-Wallmann, U. and Muller, U.M. 2000. Action plan for the conservation of the Eurasian Lynx in Europe. Nature and Environment Series 1 12: 1-69, Strasbourg, Council of Europe. Burton, R. 1979. Carnivores of Europe. Cox & Wyman, London. Corbet, G.B. 1984. The Mammals of the Palaearctic region: A Taxonomic Review. British Museum, Natural History, London. Corbet, G.B. and Harris, S. 1991. The Handbook of British Mammals third edition. Blackwell Scientific Publications, London. Gittleman, J.L. 1989. Carnivore Behaviour, Ecology and Evolution. Chapman & Hall, London. Jackson, P. and Nowell, K., 1996. Wild Cats: a status survey and conservation action plan. lUCN, Gland, Switzerland. Jenkinson, R.D.S. 1983. The recent history of northern lynx, (Lynx lynx, Linne) in the British Isles. Quaternary Newsletter A\, 1-7. Kitchener, A. 1991. The Natural History of the Wild Cats. Christopher Helm, London. Kitchener, A.C. 2002. Alien mammals: wreaking havoc or missing the boat? Glasgow Naturalist, 23, Supplement. Kitchener, A.C. and Bonsall, C. 1997. AMS radiocarbon dates for some extinct Scottish mammals. Quaternary Newsletter ^2), 1-11. Kloor, K. 1999. Lynx and biologists try to recover after a disastrous start. Science 285, 320-321 . Leaper, R., Massei, G., Gorman, M.L. and Aspinall, R. 1999. The feasibility of reintroducing Wild Boar {Sus scrofa) to Scotland. Mammal Review 29, 239-259. Nevard, T.D. and Penfold, J.B.. 1978. Wildlife conservation in Britain: an unsatisfactory demand. Biological Conservation 14, 25-44. Nowark, R.M. 1999. Mammals of the World 6"' Edition (Vol I). Baltimore and John Hopkins University Press, London. Price, J.R., 1983. Scotland’s Environment during the Last 30,000 years. Scottish Academic Press, Edinburgh. Reintroduction Specialist Group. 1995. lUCN/SSC Guidelines for Re-Introductions. lUCN Species Survival Commission, Gland, Switzerland.. Scottish Wildlife Trust, 2000. Policy on Introductions, Re-introductions and Translocation of Species. Scottish Wildlife Trust, Edinburgh. Yalden, D.W. 1999. The History of British Mammals. T & A.D. Poyser, London. Web Site SNH web site: http://www .snh .org .uk/ 58 Glasgow Naturalist 2002. Volume 24. Part 1. Pages 59-63. A FORMER WATER-MEADOW IN THE UPPER CARRON VALLEY, STIRLINGSHIRE John Mitchell 22 Muirpark Way, Drymen, Glasgow G63 ODX "The meadows are filled with every kind of field herbs, and watered with perennial streams". Sir John Clerk of Penicuik (1679-1755) INTRODUCTION The paper describes an investigation into a former water-meadow in the Upper Carron Valley, Stirlingshire. Until it was briefly uncovered by an exceptional fall in water level during the late summer of 1984, the site (NS 6885) had lain submerged by the Carron Reservoir (Fig. 1) since 1939. At over 250 ha, the Carron Meadow is reputed to have been the largest of its kind in Scotland. BACKGROUND TO WATER-MEADOWS IN SCOTLAND Two hundred years ago water-meadows were attracting considerable attention from agriculturists in Scotland, but today their creation and use is a virtually forgotten episode in the country's agrarian history. A water-meadow is defined as a grazing or hay field over which flowing water - often-silt- enriched - can be fed by a network of carefully graded channels. Equally important, a water- meadow can be laid dry at will. Following irrigation in spring such a meadow undergoes rapid growth, providing a much needed 'early bite’ for overwintered stock. The meadow would then be closed to animals, any trampling damage to the channels repaired and the vegetation watered again before being left to grow on as hay. After the hay crop was cut and gathered in, the meadow was watered once more, offering fresh grazing in late summer at a time when the drier grasslands on the farm were past their nutritional best. To judge from papers (e.g. Singers, 1807) published in the early Transactions of the Highland Society of Scotland and contemporary county agricultural reports, the peak period in the laying-out of water-meadows in Scotland was from the end of the 18th century into the early part of the 19th century. This was a time of enhanced prices for farm products, a direct result of the drawn out war with France. Although poorly documented, there does appear to have been a slightly earlier phase of water-meadow use in Scotland, of which the Carron Meadow may have been part. MAPS AND DESCRIPTIONS OF THE CARRON BOG OR MEADOW Along with the rest of the county, the Carron Valley was surveyed by the military sometime between 1749-1751. On the resultant 1:36,000 unpublished Roy map of Scotland, the Carron Bog or Meadow appears as the upper flood plain of the Water of Carron. Thirty years or so later, the same area is shown with the caption ‘a fine meadow' on Charles Ross's 1:63,360 Map of Stirlingshire issued in 1780. Not until the end of the 18th century and the publication of the first Statistical Account of Scotland is there a description of the Carron Bog or Meadow. In his contribution for the Parish of Fintry, the Reverend Gavin Gibb (1793) wrote "The [River] Carron rises in the west end of the parish, and runs east in a straight direction, watering the Carron Bog in its passage". In giving the dimensions of the Carron hay-meadow, which the Reverend Gibb considered the largest in Scotland, he also included parts which extended into two neighbouring parishes "Beginning in [the parish of] Fintry, it runs east between the parishes of Kilsyth and St. Ninians to the extent of 4 miles (6.44 km); is in some places 2 miles (3.22 km) in breadth, and in no place less than 1 mile (1.61 km); containing about 500 [Scottish] acres (630 English acres / 255 hectares) in one continuous plain. It affords sustenance during the winter to cattle of the surrounding farms. This remarkable meadow, besides its utility, adds great liveliness to the general face of the country. The scene it exhibits during the months of July and August, of 20 or 30 different parties of people employed in hay-making, is certainly very cheerful. And during winter, the greater part of it being overflowed by the Carron, which runs through the middle of it, ""which is then industriously led over the whole extent" (present writer's emphasis). This latter statement would appear to confirm that a system of water-meadow management was in operation in the Upper Carron Valley. John Grassom's 1:42,240 map of the County of Stirling (1817) is of interest from the aspect of river engineering. It shows that, compared with the Roy and Ross maps of the previous century, the meandering stretch of the River Carron running through the western part of the bog or meadow had been straightened out, presumably to improve water flow. The New Statistical Account for the Parish of Fintry (Smith, 1841) includes the following footnote on the Carron Meadow "The most abundant source of meadow hay is the Carron Bog, an immense tract of meadow land on the banks of the River Carron, commencing in [the parish of] Fintry, and thence passing into Kilsyth and St. Ninians. The meadow extends in length fully 3 miles (4.38 km) and averages in breadth about 400 yards (366 m), though in some places its breadth is little short of 1 mile (1.61 km). It contains 358 acres (145 ha)". The reason for the apparent shrinkage of the Carron Bog or Meadow is made clear with the publication 20 years later of the Ordnance Survey 1:2.500 map and accompanying Book of Reference covering the area. The original marshy ground was sufficiently agriculturally improved since conversion to water- meadow, that it had been found practicable for the best drained land to be put to the plough. Rotational cropping adopted by that time included 59 60 Fig. 3. Looking south-east: internal water channels of the fonuer Carron Water-meadow. Fig. 4. Looking north-east: internal water channels of the former Carron Water-meadow. 61 cultivated varieties of grass suitable for hay. Also shown on the OS map is the water-meadow's main water carrier diverting from the River Carron at Finnich Haugh (the farm steading later renamed The Binns), which then contoured the Carron- Endrick watershed in order that water could be gravity fed into the meadow from the opposite side of the valley (Fig. 2). In common with all water- meadows, a gentle declivity to the water carrier was essential to prevent the outlet sluices onto the meadow becoming continually clogged with moving gravel. Significantly, the name Row Burn given to the feature implies a slow-running water- course. As part of a nationwide project, an agricultural assessment of Stirlingshire was undertaken in the 1930s, fortuitously just before most of the low- lying farmland in the Upper Carron Valley was flooded following the construction of a reservoir. The field by field survey showed that rotation grass for hay was still one of the principal crops of the district (Stamp, 1946). THE CARRON RESERVOIR The feasibility of siting a reservoir in the Upper Carron Valley was first considered at the turn of the 19th century by the Carron Iron Company. Founded in 1759 alongside the lower reaches of the River Carron, the iron works was beset with water shortages during the drier summer months almost from the start. The problem was exacerbated in 1790 with the opening of the Forth and Clyde Canal, which in its Bonnybridge stretch depended on one of the main tributaries of the River Carron for its water (Campbell, 1961). Several sites for additional water storage were investigated by the Carron Company, including the Upper Carron Valley. According to a communication from the Reverend John Graham - the Minister for Fintry 1805-1822 - the local residents successfully raised an objection to the Carron Valley scheme on the grounds that their hill farms would be rendered unviable without the hay they harvested from the low-lying meadowland (Stirling, 1817). Faced with the increasing need for additional water supplies for domestic and industrial use within their district during the early 1930s, the Stirlingshire and Falkirk Water Board promoted a plan for a major new reservoir in the Upper Carron Valley. It was to be an unusual undertaking in that the proposed site straddled the central watershed between East and West Scotland, the reservoir requiring a dam at both ends. Parliamentary approval for the scheme was given in June 1935. Four years in its construction, the Carron Reservoir was formally opened on 14 July 1939 (Anon, 1939). At 5 km long, an average of 0.78 km wide and a surface area of 388 ha, the reservoir covered almost the entire floor of the Upper Carron Valley. Some of the dispossessed farm owners did however, continue to work the meadows for hay until the land was actually submerged; and at least one was still utilising the network of channels for watering his cultivated grass crop right up to the end (Mitchell, 1997), seemingly the last known instance of water- meadow management in Central Scotland. Further demand for water in the newly created Central Region administrative area led to an enlargement of the Carron Reservoir in the mid 1980s. The top water level at the east dam's overflow sill was raised by about 0.5 m, the upgrading operational by the winter of 1987/88. THE SITE OF THE CARRON WATER- MEADOW EXPOSED A visit to the Upper Carron Valley was made by the author in the late summer of 1984 on being informed that the level of the reservoir was abnormally low. The surface level had initially been dropped to facilitate structural alterations to the two dams (Dr.A.B. Bailey pers comm), but this was then followed by an unusually prolonged period of low rainfall (Harrison, 1987). The overall result was a lowering of the reservoir's water level by an unprecedented 4.98 m, a situation which more or less continued to the end of September. Examination of the western end of the receding reservoir confirmed two features already noted on 19th century maps. First was the channelised section of the River Carron, which on inspection was found to have been faced with timbers to strengthen the re-aligned banks. Second was the line of the main water carrier, which led off the River Carron and circled around the Carron- Endrick watershed to eventually rejoin the river further downstream. Not shown on any published map, however, was any evidence of a series of internal channels which, in a water-meadow, would have been necessary to distribute the water evenly over the ground. Despite a subsequent history of arable ploughing, the lay-out of the internal water channels was still visible on the drying-out bed of the reservoir (Figs. 3 & 4). Following the onset of the autumn rains, the water level gradually rose and the site of the old water- meadow was once more lost to view. With the capacity of the reservoir substantially increased since the study was carried out in 1984, it would seem that only in very exceptional conditions will this intriguing piece of Stirlingshire's agricultural history be so fully revealed again. THE ORIGINAL VEGETATION OF THE CARRON BOG OR WATER-MEADOW To the naturalist-historian, a question raised by this investigation is: what was the Carron Bog or Meadow's natural vegetation before the land was agriculturally improved in the 19th century?. One indication is to be found along the River Carron's ungrazed banks just downstream of the east dam. Seen to advantage beside the Forestry Enterprise riverside car park and picnic area, the plant assemblage is typical of northern tail-herb wet meadows formerly cropped for marsh hay, with globe flower Trollius europeus, ragged robin Lychnis flos-cuculi, large bitter cress Cardamine amara, meadowsweet Filipendula ulmaria, water avens Geum rivale, wood cranesbill Geranium sylvaticum, •mgelicz. Angelica sylvestris and water 62 sedge Car ex aquatilis well represented. Except where a vigorous growth of reed canary-grass Phalaris arundinacea has overtaken the community, there are no dominant grass species present. Cooper & MacKintosh (1996) have recommended that this type of riparian tail-herb assemblage be treated as a variant of Filipendula ulmaria - Angelica sylvestris mire M27 (see Rodwell, 1991) in the National Vegetation Classification. ACKNOWLEDGEMENTS In carrying out the field work for the above project, I am grateful to Central Region Council Water & Drainage Department and the Forestry Commission for granting me access to the Carron Reservoir and surrounding area during the summer of 1984. My thanks also to Frank Stone of Scottish Water for providing additional information on the enlargement of the reservoir in the mid 1980s. Mary Bruce of the Drymen and District Local History Society kindly commented on the first draft of the paper, and Norman Tait prepared the accompanying map and figures for publication. REFERENCES Anon (1939). Carron Valley Reservoir [commemorative booklet]. Stirlingshire & Falkirk Water Board. Campbell, R.H. (1961). Carron Company. Oliver & Boyd, Edinburgh. Cooper, E. & MacKintosh, J. (1996). NVC Review of Scottish Grassland Surveys. Scottish Natural Heritage, Battleby. Gibb, Rev.G. (1793). 'Parish of Eintry' in The Statistical Account of Scotland Vol.l 1, pp.371-382. William Creech, Edinburgh. Harrison, S.J. (1987). Annual Climatological Bulletin No. 6, 1984. Forth Naturalist and Historian 9, 3-24. Mitchell, J. (1997). Wet Meadows in Lowland West Central Scotland - an almost forgotten Botanical Habitat. Botanical Journal of Scotland 49, 341-345. Rodwell, J.S. (1991). British Plant Communities Vol.2, Mires and Heaths. Cambridge University Press. Singers. Rev.W, 1807. General Observations on the Practice and Principles of Irrigation. Prize Essays and Transactions of the Highland Society of Scotland 3,224-256. Smith, Rev.W .G. (1841). 'Parish of Eintry' in The New Statistical Account of Scotland No.32, pp.38- 47. William Blackwood, Edinburgh. Stamp, L.D. (1946). 'Stirlingshire' in The Land of Britain Pt.23, pp .355-378. Geographical Publications, London. Stirling, Rev.W.McG. (1817). Nimmo's History of Stirlingshire (2nd ed.). Andrew Bean, Stirling. 63 64 Glasgow Naturalist 2002. Volume 24. Part 1. Pages 65-68. LOCH LOMONDSIDE DEPICTED AND DESCRIBED 5. EARLY NATURAL HISTORIANS John Mitchell 22 Muirpark Way, Drymen, Glasgow G63 ODX “I have taken the Loch Lomond district as my special charge, for I feel convinced if we are ever to have a perfect knowledge of the bird-life of this country, it is only by having many workers, and by each worker having a special district. In this way scraps of information, which would otherwise be passed over, are secured, and these, when published, in due time become of value.” James Lumsden (1882). INTRODUCTION To ascribe a starting date to natural history recording on Loch Lomondside is no easy task. The 12th century chronicler Geoffry of Monmouth’s colourful tale of sixty so-called ‘eagles’ nesting on rocks in and around Loch Lomond (Giles, 1842) is but an oft- repeated legend that cannot be taken at face value. On the other hand, the 16th century map-maker Timothy Font’s comments on the yews Tcaus baccata of Inchlonaig, fallow deer Dama dama on Inchmurrin and adders Vipera berus on several other islands in the loch (Mitchell, 1907) are apparently based on personal observations in which one can have more trust. From the mid-18th century onwards both travelers and local residents with a background of scientific investigation or the collecting of trophy specimens began to take an increasing interest in the Loch Lomond area, and equally importantly committed their findings to print. Which of these early naturalists should be chosen to head a short paper such as this inevitably comes down to personal choice, but most biographers would agree that the first person to visit Loch Lomondside for the express purpose of biological recording was the Reverend John Lightfoot; and it is with his botanical exploration of Scotland that this account begins. John Lightfoot (1735-1788) Intent on collecting material for the first-ever flora of Scotland, the Reverend John Lightfoot - Chaplain to the Dowager Duchess of Portland - accompanied the antiquary Thomas Pennant and his illustrator on a summer tour of the country in 1772 (Bowden, 1989). Reaching Drymen manse on their approach to the Highlands early on 13th June, it is generally assumed that it was at this point in the journey where they were joined by the Reverend John Stuart, a newly qualified Church of Scotland minister who was to act as their gaelic interpreter and guide. While Pennant was taking in the view of Loch Lomond, the two divines spent a productive day botanising Ben Lomond, recording not only flowering plants, but cryptogams as well. When Flora Scotica (Lightfoot, 1777) first appeared it was initially badly received. As Thomas Pennant observed ... “No sooner did it come out, than Envy emptied her whole quiver” ... and the two-volume work was quietly withdrawn and not re-issued until 1789. However, the Flora was not as widely condemned as the criticism would lead one to believe. The University of Glasgow, for example, offered John Lightfoot a doctorate in recognition of his outstanding achievement, an honour he modestly declined. John Stuart (1743-1821) The Reverend John Stuart first came to prominence in scientific circles as the traveling companion to Thomas Pennant and the Reverend John Lightfoot during their tour of the Highlands in 1772 (see above). As a botanist, John Stuart’s knowledge of the Scottish mountain flora was almost without equal, with a string of first British records to his credit (Mitchell, 1986 & 1992). Later, in the resident minister’s description of the Parish of Luss (Stuart, 1796) for the first Statistical Account of Scotland, he provides us with a tantalising glimpse of his other natural history interests. Over 100 different species of birds, 21 mammals, 13 fishes plus 6 reptiles and amphibians are tabulated, a commendable total considering the indifferent quality of the identification books of the time. David Ure (1749-1798) The Reverend David Ure was yet another cleric who was drawn to the natural world. For much of his ministerial career, Ure was only able to secure assistant posts, but this did present him with the opportunity of accepting commissions from Sir John Sinclair to work on the first Statistical Account of Scotland and compile agricultural reports for several Scottish counties (Burns, 1993). As the stand-in contributor of the Parish of Killeam description (Ure, 1795) for the Statistical Account, he was not in the same position as the Reverend John Stuart in being able to produce long species lists built up over a period of time, but his experience did enable him to pick up on several worthwhile records for the area which might otherwise have been overlooked. One of these was Loch Lomondside’s last pair of lowland golden eagles Aquila chrysaetos nesting on the north face of the Campsie Fells. David Ure also noted that Finnich Glen was a haunt of the wild cat Felis catus, the animal’s former presence at this lowland locality still recalled in the place name Craighat (Creag a’ Chait). There are a few additional natural history snippets to be gleaned from his General View of the 65 Agriculture in the County of Dumbarton (lire, 1794). Included amongst them are references to the fallow deer herds on Inehlonaig and Inchmurrin, together with local uses made of certain wild plants. John Colquhoun (1805-1885) John Colquhoun’s father was Sir James Colquhoun of Luss, who had inherited the family estate on Loch Lomondside in the same year as the birth of his son. Colquhoun’s passion for the outdoors began early, but it was not until 1833, when he gave up a military career, that he was able to devote a great deal of his time to hunting, shooting and fishing, together with assembling of a comprehensive collection of stuffed mammals and birds (Maitland, 1908). As a writer, Colquhoun published the first of several semi- autobiographical works in 1840. These he brought together as a 4th edition of The Moor and the Loch (Colquhoun, 1878), which is still considered a minor classic. The content includes vivid descriptions of field sports of the period, from wildfowling on Loch Lomond to ptarmigan Lagopus nuitus shooting on the surrounding hills. Particularly revealing is Colquhoun’s account of his participation as a youth in the destruction of all birds and beasts of prey considered to be a threat in the rearing of game. In older and wiser years, however, he not only championed the greater importance of good habitat, but the necessity of predators to ensure healthy stocks of red grouse L . lagopus. In such views John Colquhoun was away ahead of his contemporaries. John Hutton Balfour (1808-1884) In 1841, John Hutton Baltour was appointed to the Chair of Botany at the University of Glasgow, but remained only four years before returning to his native Edinburgh as Professor of Botany at the University and Keeper of the Royal Botanic Garden (Bettany, 1908). Although his term of office at Glasgow was brief. Balfour extended the practice started by his predecessor Professor William Hooker of summer field trips for the botany students. Even before the advent of railways, most of Loch Lomondside was accessible to excursionists because of the regular steamer services on the River Clyde and the loch itself. John Balfour’s detailed field note books for 1846-1878 - extracts of which were published posthumously (Balfour, 1902) - show that even after moving to Edinburgh he continued to undertake botanical recording visits to the Loch Lomond area. With Ben Vorlich and Glen Ealloch conveniently to band, the Inverarnan Hotel at the loch’s steamer terminus was a popular base for Professor Balfour’s student parties. Robert Gray (1825-1887) Robert Gray’s long connection with west central Scotland began around 1845, when as a young man he took up an appointment with a Glasgow bank. As part of a group of like-minded associates, in 1851 Gray became a founder member of the Natural History Soeiety of Glasgow, holding the positions of Treasurer and Secretary in turn (Gray, 1908). During the thirty or so years he resided in Glasgow, Gray made frequent visits to Loch Lomond, the completion in 1 850 of an interconnected steamer and railway link between the City and Balloch having facilitated travel to the area still further. His first important contribution towards documenting the region’s fauna was a text with annotated species lists of mammals, birds and fishes which was appended to a guide book to Loch Lomond and the Trossachs compiled by fellow Natural History Society member William Keddie (Keddie, 1864). Robert Gray’s reputation as an ornithologist was further enhanced with the publication of his Birds of the West of Scotland (Gray, 1871) (Fig.l), Loch Lomondside being well represented in its carefully researched pages. George Hector Leith Buchanan (1833-1903) Sir George Hector Leith (his mother’s name of Buchanan was not adopted until 1877) inherited his title before the age of nine. His interest in shooting and forming a collection of stuffed birds also began at an early age, soon attracting the attention of the zoological pundits of the day by obtaining Scotland’s first example of Bonaparte’s gull Larus philadelpia on Loch Lomondside in April 1850. However, like John Colquhoun before him, it was not until retiring from the army in 1 859 and taking up residence at his mother’s family seat at Ross Priory on the south shore of the loch that he was able to take up field sports in earnest. As a corresponding member of the Natural History Society of Glasgow, Sir George Leith Buchanan more than anyone was responsible for drawing attention to the ornithological richness of the wetlands in the lower flood plain of the River Endrick (Mitchell, 1985). John Stirling (1832-1900) & Robert Kidston (1852-1924) Two 19th century floras prepared by Glasgow botany professors which made mention of Loch Lomond and surrounds - Flora Scotica (1821) by William Hooker and the Clydesdale Flora (five editions between 1 865 and 1891) by Roger Hennedy - were essentially compilations of casual records. A more systematic plant recording scheme eovering Stirlingshire, which took in the eastern portion of Loch Lomondside, was initiated by Colonel John Stirling and Robert Kidston, both members of the Stirling Natural History and Archaeological Society (Anon, 1900; Edwards, 1984). With the aim of evenness in cover, the county was divided into recording areas based on the main geological formations, such as the red sandstone and metamorphic rock zones of the region. As the field work progressed, their flora was published in two main parts with eight supplements over a period of nine years in the Transactions of the Stirling Society (Stirling & Kidston, 1891 et seq). 66 BIRDS OF THE WEST OF SCOTLAND THE NATURAL HISTORY OF ^ LOCH LOMOND i 67 Fig. 1 . Gray's Birds of the West of Scotland (1871). Fig. 2. Lumsden & Browns' Natural History of Loch Lomond and Neighbourhood {\S95) Francis Buchanan White (1842-1894) A former student of Professor Balfour at Edinburgh University, Dr Francis Buchanan White was an outstanding all-round naturalist, playing a prominent role in establishing the Perthshire Society of Natural Science in 1867 and becoming the first Editor of the Scottish Naturalist in 1871 (Taylor, 1986). His major published work The Flora of Perthshire (Buchanan White, 1898) - which divided the county into districts by the principal watersheds - was unfortunately delayed until four years after the author’s premature death. Dr. Buchanan White’s ‘Lomond District’ in the Flora took in all of the River Falloch drainage area north of Inveraman. James Lumsden (1851-1911) & Alfred Brown (1840/1-1902) Natural history recording on Loch Lomondside came of age with the publication of A Guide to the Natural History of Loch Lomond a n d Neighbourhood (Lumsden & Brown, 1895) (Fig.2), although, despite the all-embracing title, coverage was confined to the vertebrates of the area. The joint authors both enjoyed the advantage of living in the district and had previously written papers on Loch Lomondside’s vertebrate fauna for scientific journals (Anon, 1902; Mitchell, 1998). In collaborating for the book. James Lumsden - a member of the Natural History Society of Glasgow - prepared the chapters on mammals and birds, while Alfred Brown - Secretary of the Loch Lomond Angling Association - took responsibility for reptiles, amphibians and fishes. Although much has been written on Loch Lomondside’s wildlife since the late 19th century, Lumsden & Browns’ Guide still holds a place amongst the standard works. ACKNOWLEDGEMENTS My special thanks to Allan Stirling and Dr. Jack Gibson for their valued comments on the draft paper, and to Norman Tait who kindly prepared the two figures for publication. REFERENCES Anon. (1900), Colonel John Stirling. Obituary: Stirling Journal & Advertiser, 25 May 1900. Anon. (1902). Alfred Brown. Obituary: Lennox Herald, ?>\ May 1902. Balfour, I.B., (1902). Notes o/Botanical Excursions made by Professor John Hutton Balfour. The Royal Botanic Garden, Edinburgh 7,21-497. Bettany, G.T. (1908). ‘John Hutton Balfour (1808-1884)' in Dictionary of National Biography (2nd ed.), Vol.l , p.976. Smith & Elder, London. Bowden, J.K. (1989). John Lightfoot: His Work and Travels. Royal Botanic Gardens, Kew. Buchanan White, F. (1898). The Flora of Perthshire. Perthshire Society of Natural Science, Edinburgh. Burns, J.H. (1993). David Ure (1749-1798): Breadth of Mind and Accuracy of Observation. Glasgow Naturalist 22, 259-275. Colquhoun, J. (1878). The Moor and the Loch 2 vols. William Blackwood, Edinburgh. Edwards, D. (1984). Robert Kidston: The Most Professional Palaeobotanist. Forth Naturalist and Historian 8,64-93. Giles, Rev.J.A. (1842). The British History of Geoffrey of Monmouth. James Bohn, London. Gray , J .M . ( 1 908) . ‘ Robert Gray ( 1 825- 1 887) ’ In Dictionary of National Biography (2nd ed.), Vol.VIlI, pp .462-463. Smith & Elder, London. Gray, R. (1871). The Birds of the West of Scotland. Thomas Murray, Glasgow. Keddie, W. (1864). Highland Route No. VII: Edinburgh and Glasgow to the Lake of Menteith, the Trossachs, Loch Katrine and Loch Lomond. Maclure & Macdonald, Glasgow. Lightfoot, Rev.J. {Mil). Flora Scotica 2 vols. B. White, London. Lumsden, J. & Brown, A. (1895). A Guide to the Natural History of Loch Lomond and Neighbourhood. David Bryce, Glasgow. Maitland, J.A.F. (1908). ‘John Colquhoun (1805-1885)’ in Dictionary of National Biography (2nd. ed.), Vol.IV, p.858. Smith & Elder, London. Mitchell, Sir A. (1907). MacFarlane’s Geographical Collections relating to Scotland Vol.II. Scottish History Society, Edinburgh. Mitchell, J. (1985). ‘Sir George Leith Buchanan’ in Early Scottish Ornithologists on the Loch Lomondside Scene. Loch Lomond Bird Report 1984 13. 12-13. Mitchell, J. (1986 & 1992). The Reverend John Stuart D.D. (1743-1821) and his contribution to the discovery of Britain’s Mountain Flowers. Glasgow Naturalist 2 \ , 119- 125 & 22, 103-105. Mitchell, J. (1998). James Lumsden: a late 19th century Bird Recorder. Clyde Birds 10, 144-145. Stirling, J. & Kidston, R. (1891 et seq). Notes on the Flora of the North-Western Portion of Stirlingshire. Transactions of the Stirling Natural History and Archaeological Society 13,88-102. Stuart, Rev.J. (1796). ‘Parish of Luss’ in The Statistical Account of Scotland Vol.l7, 238-271 . William Creech, Edinburgh. Taylor, M.A. (1986). Francis Buchanan White (1842-1894) and Scottish Botany. Scottish Naturalist 1986, pp. 157-173. Ure, Rev.D. (1794). General View of the Agriculture in the County of Dumbarton. W. Smith, London. Ure, Rev.D. (1795). ‘Parish of Killeam’ in The Statistical Account of Scotland Vol.16, 100-129. William Creech, Edinburgh. Glasgow Naturalist 2002. Volume 24. Part 1. Pages 69-82. ENVIRONMENTAL IMPACT ON THE SEA BED CAUSED BY TRAWLING FOR THE NORWAY LOBSTER, NEPHROPS NORVEGICUS IN THE CLYDE SEA AREA. A GEOTECHNICAL ASSESSMENT. Azra Meadows, Peter S. Meadows and John M. H. Murray Biosedimentology Unit, Division of Environmental and Evolutionary Biology Institute of Biomedical and Life Sciences, University of Glasgow, G12 8QQ, UK Email: gbza21@udcf.gla.ac.uk. Web Site: www.gla.ac.uk/Acad/IBLS/Biosed ABSTRACT The Norway Lobster, Nephrops norwegicus, is a common benthic crustacean living on the sea bed in the Clyde Sea area, Irish Sea, North Sea, and Mediterranean. It lives between 50 metre and 400 metres water depth in fine mud and constructs large burrow systems. It is a high-value catch for local fishermen, and the most usual method of fishing is with an otter trawl. Otter trawls cause considerable disturbance to the sea bed. There have therefore been a number of investigations of the fishery, and on its environmental impact on the seabed and biological communities that live there. Sediment cores were taken from high, medium and low trawl impact sites in the Clyde Estuary using a new sea-bed coring device developed at the University Marine Biological Station Millport (UMBS) by Dr P.R.O. Barnett. The new coring device provided sediment cores of c. 14 to 18 cm depth, that were undisturbed and were covered with in-situ overlying water. Two stations were selected at each of the three impact sites. The stations represented a range of sedimentary environments from sandy mud to fine mud. Sampling was conducted at four times during the 2000 - 2001 year: Spring 2000, Summer 2000, Autumn 2000, and Winter 2000/Spring 2001 . Immediately after collection, the sediment cores from the pairs of stations at the high, medium and low impact sites were submitted to six quantitive geotechnical tests in the laboratory. The geotechnical tests used were shear strength, load resistance, water content, dry bulk density and particle size mean and sorting. Measurements were taken at 0 - 2 cm, 4-6 cm, and 8 - 10 cm in the sediment cores. Shear strength, load resistance and bulk density increased, and water content decreased, with increasing depth in the sedimentary column. There were no obvious changes in mean particle size or sorting with increasing depth in the sedimentary column. No marked seasonal changes were noted in any of the geotechnical parameters. The six geotechnical parameters used all detected differences in the sediment structure and characteristics between the low, medium and high impact sites. There was a decrease in sediment shear strength, load resistance, dry bulk density, mean particle size, and particle size sorting, as trawling impact increased. There was an increase in sediment water content as trawling impact increased. The degree to which the geotechnical parameters detected changes in sediment structure between the low, medium and high impact sites differed markedly. Shear strength and load resistance showed the largest differences and are therefore the most robust of the tests used. Shear strength decreased by 52% at the medium impact site and 87% at the high impact site, when compared with the low impact site. The equivalent changes for load resistance were a decrease of 47% and 86% respectively. The changes for bulk density were a decrease of 18% and 50%, and the changes for water content were an increase of 9% and 61%, at the medium and high impact sites respectively. The changes for phi mean particle size were an increase of 1 1% and 19%, and for phi sorting an increase of 3% and 5%. In terms of the speed with which the geotechnical tests can be conducted, shear strength and load resistance are the fastest tests (4 hours), followed by water content and bulk density (7 days), and particle size mean and particle size sorting (14 days). A Speed Robustness Index (SRI) has been developed for the six geotechnical parameters. This quantifies the effectiveness of the six parameters in terms of their percentage change as trawling intensity increases, and of the time taken to conduct the tests. Shear strength and load resistance have the highest SRI's of 17.4 and 16.6. Water content and bulk density have SRI's of 0.63 and 0.61, and mean particle size and sorting have SRI's of 0.18 and 0.05. The results from this survey therefore show that shear strength and load resistance are the most effective of the six geotechnical parameters used to assess the environmental impact of Otter trawling on the structure of the sea bed in the Clyde Estuary. They are both rapid tests and both show large changes as trawling impact increases. For the future, we suggest that the six geotechnical parameters used in this report are assessed in other high medium and low impact trawling areas of the sea bed in the Clyde Sea area and then further afield, to confirm their relative suitability and robustness in a wider range of environmental conditions. This should initially concentrate on areas where the Norway Lobster, Nephrops norwegicus, is abundant. It should then be extended to other commercially important species and to ecosystems where man’s impact on the sea bed is thought to be deleterious. 69 INTRODUCTION The Norway Lobster, Nephrops norwegicus, is a common benthic crustacean living on the sea bed in the Mediterranean, North Sea, and Irish Sea (Maynou et al. 1998; Relini, 1998; Sarda, 1998a, b; Abello et al. 2002). It is also found in considerable numbers off the west coast of Scotland, being well known from the Clyde Sea area and Firth of Clyde (Field et al. 1998; Tuck et al. 2000; Briggs & McAliskey, 2002; Parslow-Williams et al. 2002). The species lives subtidally, between about 50 metre and 400 metres water depth on fine muddy substrates where it constructs large burrow systems. Nephrops norvegicus is a high-value catch for local fishermen, and the most usual method of fishing is with an otter trawl (Abella et al., 1999; Madsen & Nansen, 2001; Campos et al. 2002; Poulard & Leaute, 2002). These trawls and similar fishing gear are known to cause considerable disturbance to the sea bed during their operation (De Groot, 1984; Jones, 1992; Jennings & Kaiser 1998; Schwinghamer et al. 1998;Thrush et al. 1998), and there is some concern about the effects of this on the biodiversity of benthic communities (Hall et al. 1990; Kaiser et al. 1996; MacDonald et al. 1996; Hall & Harding, 1997; Engel & Kvitek 1998; Poiner et al. 1998; Ball, Fox, & Munday, 2000; Ball, Munday, & Tuck, 2000;Collie et al., 2000). There are also environmental effects caused by the discarded dead organisms from Nephrops trawlers (Stratoudakis et al. 2001; Bergman et al. 2002a, b). As a result, there have been a number of investigations of the fishery, and on its environmental impact on the seabed and biological communities that live there. The objectives of our research was to find out whether simple physical measurements on sediments could identify the impact of otter trawl fishing for Nephrops norwegicus on the seabed in the Firth of Clyde which would then have application to other areas in which the species was abundant. This is particularly important in relation to benthic invertebrate communities living in the same environment as Nephrops. Many of the animals in these communities, including many molluscs, annelids, and echinoderms, are very sensitive to changes in the physical and chemical properties of the sediments on or in which they live (Meadows & Meadows, 1991). If these properties change, animals often cannot feed and burrow properly. They are therefore likely to die or move elsewhere. This reasoning has broader implications. The same sort of effects will be true of other marine environments in which anthropogenic disturbance takes place, such as the intertidal zone (Meadows & Tufail, 1986; Tufail, Meadows & McLaughlin, 1989; Meadows, Meadows & McLaughlin, 1998). This zone is often the site of digging for clams or edible cockles, and in some areas significant disturbance takes place by bathers and by research sampling. The problem has received little attention. Specifically, we aimed to answer the question 'Can geotechnical parameters detect changes in the structure of the sea bed associated with an increase in Otter trawl activity?'. We therefore gave detailed consideration to the suite of tests that should be used. The tests should include standard as well as novel experimental protocols and equipment. They should provide a range of different geotechnical signatures for any change in sediment structure. The speed with which the results can be conducted should also receive detailed consideration. The techniques that we used are part of a range of geotechnical measurements that are routinely used by civil engineers and environmentalists to quantify the properties of soils on land and sediments in freshwater and marine ecosystems (Meadows & Meadows, 1994). These are shear strength, load resistance, dry bulk density, water content, mean particle size and particle size sorting (standard deviation). These geotechnical techniques are all used routinely by the present authors and their co- workers (Meadows & Tufail, 1986; Meadows & Tait, 1985, 1989; Meadows & Meadows, 1994; Meadows et al. 2000, Murray et al. 2000), and have been tested over a number of years by us in a wide range of intertidal, continental shelf, continental slope and abyssal plain environments. Shear strength is known to quantify sediment structure in an engineering context, and load resistance is a novel microscale approach. Dry bulk density and water content are standard geotechnical parameters, and are likely to change with sediment disturbance. Mean particle size and sorting are routinely used by sedimentologists to distinguish between different sedimentary ecosystems, particularly in terms of different degrees of disturbance associated with high and low energy sea bed environments MATERIALS AND METHODS Stations, sampling dates and number of tests High, medium and low trawling intensity sites, were identified in the Clyde Estuary by Dr R.J. A. Atkinson, and colleagues at the University Marine Biology Station Millport. Two stations were selected that were representative of each of the three sites, and were sampled during spring 2000, summer 2000, autumn 2000 and winter 2000/01 - a total of 24 stations in all. The pairs of stations at the high, medium and low trawling intensity sites, with their dates of sampling, latitude and longitude, and water depth, are given in table 1 . Six geotechnical tests were conducted on cores from each station, hence in all 24 x 6 = 144 geotechnical tests were undertaken during the research. Geotechnical Techniques All the geotechnical techniques were conducted on cores of sediment that had been sampled from the sea bed within 24 hours. A new sea bed corer was used developed by Dr P.R.O. Barnett of UMBS. The sediment cores were contained in sealed 70 Table 1. Detailed log of the High, Medium and Low impact sites and their stations. Trawling Intensity sites, High (stations: WCUM Wee Cumbrae, BROD Brodick), Medium (stations ; INCH Inchmarnock, LOLL Lower Loch Long) and Low (stations : HUHU Hush Hush, MLL Mid Loch Long) in the Clyde Sea area. Trawling Intensity High Medium Low Spring 2000 WCUM BROD INCH LOLL HUHU MLL Date 13.04.00 13.04.00 13.04.00 12.04.00 14.04.00 12.04.00 Lat. W Long. N 55° 41' 29” 04° 58' 33" 55° 35' 21" 05° 03' 24" 55° 44' 40" 05° 09' 32" 56° or 30” 04° 52' 59" 55° 47' 25" 04 56' 19" 56° 03' 27" 04° 53' 19" Water Depth (m) 81 97 81 61 37 86 Summer 2000 Date 28.07.00 03.08.00 28.07.00 27.07.00 03.08.00 27.07.00 Lat. W 55° 41' 51" 55° 35' 22" 55° 45' 24" 56° or 55" 55° 47' 32" 56° 03' 56" Long. N 04° 58' 35" 05° 03' 18" 05° 09' 18" 04° 53' 11" 04° 56' 18" 04° 52' 54" Water Depth 86 95 59 64 39 87 (m) Autumn 2000 Date 31.10.00 24.10.00 24.10.00 25.10.00 31.10.00 23.10.00 Lat. W 55° 41' 30" 55° 35' 46" 55° 45' 36" 56° or 18" 55° 47' 31" 56° 03' 36" Long. N 04° 58' 37" 05° 03' 27" 05° 09' 40" 04° 53' 04" 04° 56' 19” 04° 53' 19" Water Depth 80 94 60 64 38 80 (m) Winter 2000/1 Date 22.02.01 26.02.01 26.02.01 23.02.01 22.02.01 23.02.01 Lat. W 55° 4r.22" 55° 35' 44" 55° 45' 19" 56° or 39" 55° 47' 36" 56° 03' 50" Long. N 04° 58' 50" 05° 03' 20" 05° 09' 20" 04° 53' 16” 04° 56' 20" 04° 53' 11” Water Depth 83 95 60 65 41 90 (m) 71 transparent core liners, internal diameter 85 cm height 30 cm. The sediment cores were 14 to 18 cm in height and were covered by in-situ overlying water. In the laboratory, the overlying water was siphoned off the sediment core. Shear strength was then measured on the surface of each sediment level, the sediment was then extruded and sliced, and sediment samples were taken from the sliced sections for water content, dry bulk density and particle size. Load resistance was measured on a separate core taken close to the first core on the sea bed. The overlying water on this second core was not siphoned off before load resistance was measured. Data for all the geotechnical parameters measured are presented for the following depths of the sedimentary column: 0-2 cm, 4-6 cm, and 8 - 10 cm. These three levels were chosen to represent the top, middle and bottom of the sediment core. Shear Strength was measured using a Geonor Fall Cone apparatus (Meadows & Tait, 1985, 1989; Meadows & Meadows 1991; Meadows et al. 1994). It is expressed as kN/m^ in the tables of results. Load resistance in kN was measured using a miniature load resistance penetrometer developed by Muir Wood et al. (1993), Meadows, P.S. et al. (1998) and Mun-ay et al. (2000). The experimental protocols for measuring load resistance described in these three papers were followed with minor modifications. Water Content was measured by weighing wet sediment samples, and then drying at 80° C for 24 hours. This was followed by cooling in a dissector for 3 hours, after which the samples were weighed. Water content was then calculated as ((wet weight - dry weight)/dry weight)xl00. It is presented as % water content in the tables of results (BS1377, 1975). Dry bulk density was measured on the same samples and was calculated as (volume of sediment)/ (dry weight of sediment). It is presented as g/ml in the tables of results. Particle size was measured as follows. Sediment from the three levels of the cores (0-2 cm, 4 - 6 cm, and 8-10 cm) was supplied to the Marine Laboratory Aberdeen for granulometric analysis. The Marine Laboratory Aberdeen provided granulometric data as percentages at half phi intervals from 1 to 11. This data was then used by the authors to calculate the phi mean and phi sorting of each sample (Buchanan, 1984). The data are presented as phi mean and phi sorting in the tables of results, where phi = -(log 2 particle diameter in mm). Hence - 1 phi = 2 mm, 0 phi = 1 mm, +1 phi = 500 micron, +2 phi = 250 micron, -i-3 phi = 125 micron and so on. RESULTS The results of each of the geotechnical parameters measured are presented in the same tabular format. There is one table for each parameter: each presents the data for spring 2000, summer 2000, autumn 2000 and winter 2000/spring 2001. In each table data are presented for 0 - 2 cm, 4-6 cm, and 8-10 cm sediment depths. These are presented for the two high impact stations (Wee Cumbrae and Brodick), the two medium impact stations (Inchmamock and Lower Loch Long) and the two low impact stations (Hush Hush and Mid Loch Long) . Sediment Shear Strength and Load Resistance Sediment shear strength and load resistance data are presented in tables 2 and 3. An inspection of these eight tables show a number of common features. Shear strength and penetration resistance are closely related. Low shear strength occurs in a sediment having a low load resistance, and high shear strength occurs in sediment having a high load resistance. This is to be expected from previous work by the authors (Meadows, P.S. et al. 1998; Murray et al. 2000) and indicates that the two techniques are measuring similar properties of the sedimentary column. Both shear strength and load resistance increase slightly with increasing sediment depth. This is a common feature of such data (Meadows & Tait, 1985; Meadows & Tufail, 1986; Meadows & Meadows, 1991; Meadows, P.S. et al. 1998; Meadows, A. et al., 2000; Murray et al., 2000). There is very little difference in the two parameters between the four seasons. In other words there is no marked seasonal signature in the results. There is some suggestion in the data that there is a decrease in shear strength and a decrease in load resistance as trawling impact increases, however there is considerable variation. This effect is considered in more detail below (section 5) Water Content and Dry Bulk Density The data for water content and dry bulk density are presented in table 4 and 5. It is clear from the data in these two tables that water content and dry bulk density are inversely related. When water content is high, dry bulk density is low, and when water content is low dry bulk density is high. This is as expected. When a sediment contains less water the individual particles are more tightly packed together which in turn produces a higher bulk density. There is a decrease in water content and increase in dry bulk density with increasing sediment depth at all the stations. This effect is well known. There is very little difference in the two parameters between the four seasons. There is hence no seasonal signature shown by either parameter. There is some indication from the data that water content increases and bulk density decreases as trawling impact increases, however there is considerable variation. This effect is considered in more detail below (section 5). Particle Size. Mean and Sorting The data for mean particle size and sorting (standard deviation) are presented in table 6 and 7. The units used are phi units, following standard sedimentological procedure (cf. Buchanan, 1984). In these units a higher value of mean phi represents a smaller particle size, and a lower value for mean phi represents a larger particle size. This is not so of the phi sorting (phi standard deviation). Here a 72 higher phi standard deviation (defined as a lower sorting) indicates a smaller scatter of data - just as sorting) indicates a wider scatter of data, and a in normal standard deviations, lower phi standard deviation (defined as a higher Table 2. Sediment shear strength (kN. m'^) mean ± s.d. values (n = 6) at top (0-2 cm), mid (4-6 cm) and bottom (8-10 cm) depths down a sediment core (core internal diameter = 10 cm). Three Trawling Intensity sites, High (stations :WCUM Wee Cumbrae, BROD Brodick), Medium (stations : INCH Inchmarnock, LOLL Lower Loch Long) and Low (stations : HUHU Hush Hush, MLL Mid Loch Long) in the Clyde Sea area. First set of data: Spring 2000; second set: Summer 2002; third set: Autumn 2000; fourth set winter 2001. Trawling Intensity High WCUM BROD Low HUHU MLL Sediment Depth INCH LOLL 0-2 cm 0.137 0.102 0.149 0.502 1.396 0.150 ± 0.062 ±0.043 ±0.072 ±0.064 ±0.412 ±0.028 4-6 cm 0.423 0.373 1.140 0.573 2.704 0.354 ±0.094 ±0.150 ±0.625 ±0.066 ±0.964 ±0.072 8-10 cm 0.345 0.511 2.439 0.505 10.103 0.422 ±0.132 ±0.205 ±0.604 ±0.094 ±4.736 ±0.127 0-2 cm 0.107 0.118 0.255 0.122 0.666 0.118 ± 0.027 ±0.049 ±0.137 ±0.045 ±0.457 ±0.028 4-6 cm 0.616 0.432 1.619 0.507 8.240 0.492 ±0.264 ±0.153 ±0.334 ±0.100 ±5.780 ±0.113 8-10 cm 2.370 0.552 5.299 0.816 14.157 0.816 ±1.706 ±0.097 ±2.594 ±0.298 ±11.329 ±0.171 0-2 cm 0.129 0.096 0.250 0.150 0.788 0.139 ±0.053 ±0.026 ±0.161 ±0.085 ±0.553 ±0.039 4-6 cm 0.384 0.522 4.974 0.412 5.837 0.352 ±0.127 ±0.170 ±2.849 ±0.114 ±1.426 ±0.007 8-10 cm 0.889 0.531 1.345 0.781 7.237 0.799 ±0.224 ±0.072 ±1.428 ±0.297 ±2.642 ±0.158 0-2 cm 0.151 0.090 0.487 0.114 0.968 0.135 ±0.070 ±0.030 ±0.242 ±0.040 ±0.918 ±0.071 4-6 cm 0.589 0.463 3.134 0.549 6.680 0.401 ±0.140 ±0.075 ±1.761 ±0.164 ±5.220 ±0.084 8-10 cm 0.866 0.730 15.210 0.847 22.84 1.142 ±0.223 ±0.132 ±7.47 ±0.154 ±12.81 ±0.293 73 Table 3. Load resistance (kN) mean ± s.d. values (n = 26) at top (0-2 cm), mid (4-6 cm) and bottom (8-10 cm) depths down a sediment core (core internal diameter = 10 cm). A 30 mm diameter penetration probe was used. Three Trawling Intensity sites. High (stations :WCUM Wee Cumbrae, BROD Brodick), Medium (stations : INCH Inchmarnock, LOLL Lower Loch Long) and Low (stations : HUHU Hush Hush, MLL Mid Loch Long) in the Clyde Sea area. OS: load resistance over scale, value of greater than 100 N. First set of data; Spring 2000; second set: Summer 2002; third set; Autumn 2000; fourth set winter 2001. Sediment Depth Trawling Intensity High WCUM BROD INCH Medium LOLL HUHU Low MLL 0-2 cm 0.072 0.370 0.959 0.784 6.532 0.861 ±0.079 ±0.218 ±0.615 ±0.459 ±5.467 ±0.637 4-6 cm 1.564 2.464 6.208 4.960 55.91 4.198 ±0.305 ±0.590 ±1.114 ±0.630 ±12.50 ±0.409 8-10 cm 4.254 10.04 13.02 14.77 66.71 6.735 ±0.505 ±1.547 ±0.443 ±2.144 ±1.261 ±0.356 0-2 cm 0.323 0.456 3.194 0.854 7.498 0.324 ±0.275 ±0.299 ±2.386 ±0.688 ±6.006 ±0.22 8 4-6 cm 3.432 3.050 38.57 6.329 85.19 3.234 ±0.574 ±0.310 ±6.663 ±0.834 ±11.97 ±0.658 8-10 cm 5.784 5.998 85.62 10.38 7.652 ±0.230 ±0.530 ±7.825 ±0.531 OS ±0.499 0-2 cm 0.695 0.646 1.064 1.076 4.100 0.563 ±0.507 ±0.424 ±0.611 ±0.642 ±2.528 ±0.385 4-6 cm 6.000 4.429 8.010 5.910 44.54 3.398 ±0.698 ±0.748 ±1.931 ±0.740 ±11.89 ±0.401 8-10 cm 10.95 9.694 24.02 8.842 6.582 ±0.975 ±0.744 ±2.332 ±0.239 OS ±0.630 0-2 cm 0.858 0.306 4.040 0.652 4.455 0.890 ±0.498 ±0.232 ±3.266 ±0.392 ±3.913 ±0.514 4-6 cm 4.810 2.382 31.90 3.039 47.54 4.116 ±0.561 ±0.427 ±4.940 ±0.433 ±10.88 ±0.470 8-10 cm 8.032 4.752 72.96 6.629 8.672 ±0.473 ±0.253 ±6.932 ±0.740 OS ±1.039 74 Table 4. Water content (%) mean ± s.d. values (n = 4) at top (0-2 cm), mid (4-6 cm) and bottom (8-10 cm) depths down a sediment core (core internal diameter = 10 cm). The three Trawling Intensity sites, High (stations :WCUM Wee Cumbrae, BROD Brodick), Medium (stations : INCH Inchmarnock, LOLL Lower Loch Long) and Low (stations ; HUHU Hush Hush, MLL Mid Loch Long) in the Clyde Sea area. First set of data: Spring 2000; second set: Summer 2002; third set: Autumn 2000; fourth set: winter 2001. Sediment Depth Trawling Intensity High Medium Low WCUM BROD INCH LOLL HUHU MLL 208.45 252.75 116.70 205.53 40.87 232.05 0-2 cm ±16.022 ±25.189 ±21.765 ±11.022 ±4.279 ±4.208 167.10 196.10 78.15 209.73 36.095 196.53 4-6 cm ±1.977 ±8.163 ±2.765 ±90.854 ±2.551 ±5.525 163.35 185.55 73.33 158.425 29.325 177.85 8-10 cm ±9.840 ±5.444 ±2.210 ±8.019 ±0.807 ±11.388 0-2 cm 240.83 224.65 79.73 229.60 43.70 255.68 ±45.723 ±9.110 ±16.596 ±11.257 ±4.975 ±26.109 4-6 cm 149.70 194.33 47.70 170.33 31.60 183.13 ±4.491 ±3.732 ±0.970 ±4.371 ±1.052 ±6.562 121.83 178.88 53.05 142.65 27.23 167.08 8-10 cm ±5.090 ±4.001 ±2.330 ±2.483 ±0.967 ±7.475 0-2 cm 217.70 254.48 110.05 233.45 40.25 246.73 ±22.241 ±12.891 ±15.765 ±25.501 ±4.234 ±6.620 4-6 cm 172.65 187.93 67.33 178.80 30.95 201.65 ±14.214 ±5.902 ±3.784 ±6.351 ±0.790 ±4.966 8-10 cm 142.95 177.68 68.50 152.95 30.05 174.03 ±0.889 ±3.745 ±1.874 ±4.088 ±0.778 ±1.752 0-2 cm 214.06 253.50 54.01 243.20 36.93 255.50 ±12.57 ±16.82 ±6.93 ±23.20 ±3.360 ±38.00 4-6 cm 154.23 197.34 39.75 179.57 29.11 191.35 ±6.61 ±8.07 ±1.55 ±4.81 ±4.62 ±4.59 140.82 167.54 40.69 157.25 27.10 158.02 8-10 cm ±3.11 ±4.41 ±1.80 ±3.38 ±0.319 ±2.02 75 Table 5. Dry bulk density (g.ml ') mean ± s.d. values (n = 4) at top (0-2 cm), mid (4-6 cm) and bottom (8-10 cm) depths down a sediment core (core internal diameter = 10 cm). Three Trawling Intensity sites. High (stations :WCUM Wee Cumbrae, BROD Brodick), Medium (stations : INCH Inchmarnock, LOLL Lower Loch Long) and Low (stations : HUHU Hush Hush, MLL Mid Loch Long) in the Clyde Sea area.. First set of data: Spring 2000; second set: Summer 2002; third set: Autumn 2000; fourth set winter 2001. Trawling Intensity WCUM High BROD Medium INCH LOLL Low HUHU MLL Sediment Depth 0-2 cm 0.713 0.600 1.146 0.714 2.281 0.668 ±0.050 ±0.052 ±0.164 ±0.021 ±0.105 ±0.008 4-6 cm 0.865 0.769 2.097 0.773 2.388 0.773 ±0.025 ±0.017 ±0.040 ±0.167 ±0.092 ±0.013 8-10 cm 0.874 0.799 2.162 1.197 2.620 0.821 ±0.041 ±0.022 ±0.038 ±0.333 ±0.049 ±0.053 0-2 cm 0.654 0.676 1.518 0.654 2.183 0.591 ±0.090 ±0.021 ±0.207 ±0.032 ±0.138 ±0.041 4-6 cm 0.956 0.763 2.091 0.847 2.545 0.801 ±0.026 ±0.012 ±0.022 ±0.011 ±0.026 ±0.030 8-10 cm 1.124 0.817 1.998 0.979 2.703 0.862 ±0.031 ±0.020 ±0.061 ±0.011 ±0.064 ±0.038 0-2 cm 0.689 0.581 1.240 0.656 2.060 0.616 ±0.062 ±0.021 ±0.182 ±0.069 ±0.457 ±0.018 4-6 cm 0.840 0.794 1.745 0.828 2.624 0.744 ±0.067 ±0.021 ±0.073 ±0.022 ±0.038 ±0.015 8-10 cm 0.980 0.841 1.680 0.925 2.679 0.848 ±0.011 ±0.015 ±0.028 ±0.021 ±0.023 ±0.014 0-2 cm 0.398 0.342 1.100 0.355 1.351 0.338 ±0.018 ±0.022 ±0.0926 ±0.0307 ±0.069 ±0.051 4-6 cm 0.536 0.428 1.323 0.467 1.532 0.448 ±0.016 ±0.017 ±0.0340 ±0.012 ±0.064 ±0.014 8-10 cm 0.578 0.497 1.308 0.527 1.610 0.519 ±0.009 ±0.008 ±0.027 ±0.007 ±0.020 ±0.008 76 Table 6. Particle Size. Phi mean ± s.d. values (n = 4) at top (0-2 cm), mid (4-6 cm) and bottom (8-10 cm) depths down a sediment core (core internal diameter = 10 cm). Three Trawling Intensity sites, High (stations :WCUM Wee Cumbrae, BROD Brodick), Medium (stations ; INCH Inchmarnock, LOLL Lower Loch Long) and Low (stations : HUHU Hush Hush, MLL Mid Loch Long) in the Clyde Sea area.. First set of data: Spring 2000; second set; Summer 2002; third set: Autumn 2000; fourth set winter 2001. Trawling Intensity High WCUM BROD INCH Medium LOLL Low HUHU MLL Sediment Depth 0-2 cm 5.810 6.595 5.271 5.496 4.280 6.009 ±1.861 ±1.356 ±1.882 ±1.848 ±2.024 ±1.656 4-6 cm 4.771 6.244 5.412 5.712 4.272 6.100 ±2.784 ±2.195 ±2.011 ±1.978 ±2.039 ±1.471 8-10 cm 3.120 5.314 5.544 5.773 4.287 6.159 ±2.879 ±2.900 ±1.836 ±2.055 ±1.971 ±1.500 0-2 cm 5.849 6.266 4.828 6.157 3.920 6.073 ±1.754 ±1.545 ±2.018 ±1.412 ±1.975 ±1.476 4-6 cm 5.995 6.214 4.961 6.029 4.127 6.214 ±1.681 ±1.959 ±2.000 ±1.540 ±1.954 ±1.523 8-10 cm 5.924 6.238 5.407 6.011 3.988 6.096 ±1.680 ±1.820 ±2.256 ±1.619 ±2.014 ±1.496 0-2 cm 5.946 5.942 5.461 5.758 3.845 5.907 ±1.770 ±1.985 ±1.687 ±1.656 ±1.928 ±1.580 4-6 cm 6.052 6.330 5.724 5.924 3.978 5.725 ±1.687 ±1.554 ±1.658 ±1.558 ±1.966 ±1.726 8-10 cm 6.096 6.302 5.894 6.056 4.304 5.912 ±1.691 ±1.700 ±1.672 ±1.457 ±1.909 ±1.756 0-2 cm 6.335 6.523 4.619 6.177 3.938 6.224 ±1.601 ±1.425 ±2.073 ±1.523 ±2.018 ±1.353 4-6 cm 6.390 6.606 4.608 6.080 3.740 6.220 ±1.412 ±1.424 ±2.142 ±1.545 ±1.466 ±1.407 8-10 cm 6.448 6.632 5.010 6.337 3.837 6.254 ±1.519 ±1.522 ±2.202 ±1.403 ±1.943 ±1.592 In contrast to the previously described parameters, there is no consistent increase or decrease in the particle size mean or sorting as sediment depth increases. However in common with the previously described parameters there is very little difference between the four seasons. There is some indication from the data that as trawling impact increases, the mean particle size decreases (increase in phi mean particle size), and the particle sorting decreases (increase in phi particle size standard deviation). This effect is considered in more detail below (section 5). 77 CONCLUSIONS Increased Trawling Impact and Changes in the Measured Geotechnical Parameters Careful consideration has been given to the way in which the effects of increasing trawling for Nephrops norvegicus and its impact on the measured geotechnical parameters could be quantified. It was eventually decided to calculate the means and standard deviations for each parameter in turn for the high impact stations, for the medium impact stations, and for the low impact stations. This was done using the data for all three sediment depths, all four seasons, and the two stations at each impact level. This resulted in a series of means and standard deviations based on individual means from three sediment depths, two stations, four seasons. Each mean and standard deviation is therefore base on 3 x 2 x 4 = 24 readings. The results are given in table 7. Statistically, care needs to be exercised in calculating means of means, and in basing deductions on them. However the data in table 7 suggest that increasing trawl impact does have effects on all the geotechnical parameters measured. These effects are as follows. Sediment shear strength and sediment load resistance decrease with increasing trawl impact. Sediment water content increases with increasing trawl impact. Dry bulk density decreases with increasing trawl impact. Sediment mean particle size decreases (increasing phi mean) with increasing trawl impact. Sediment sorting decreases (increasing phi standard deviation) with increasing trawl impact. The results in table 7 therefore indicate that any one of the six sedimentary geotechnical parameters measured can detect differences caused by trawl impact on the sediment fabric within the top 10 cm of the sedimentary column. An Interpretation of the Observed Effects If one assumes that Otter trawling for Nephrops norvegicus will disturb the sediment fabric to a depth well in excess of the 10 cm sediment cores investigated in the present study, a mechanism for the observed changes in the sedimentary geotechnical parameters can be suggested. Trawling over and through the surficial layers of a sediment will both mix the sediment and disperse it vertically and horizontally. This will immediately lead to a lower shear strength, load resistance and bulk density, and to a higher water content. It will also lead to a reduction in sorting (increased standard deviation of particle size). It is not so obvious however, why trawling should lead to a reduction in mean particle size - as observed in the present study. Perhaps finer material is released from deeper parts of the sedimentary column by the effect of the trawling. Alternatively, the finer material is captured in some way from the overlying water column by persistent trawling, and then incorporated into the surficial layers of the sedimentary column. Physico-chemical flocculation processes and biological activity of one form or another may play a role in both of these hypothesised processes. Speed and Robustness of the Six Sedimentary Parameters as Indicators of Trawling Intensity and Sediment Disturbence on Nephrops grounds Our aim was to assess and develop a rapid methodology for quantifying environmental impacts of Otter trawls on the sea bed. Consideration has therefore been given to assessing the speed of the geotechnical techniques used above and to their robustness. A quantitative assessment has been made of the time required to conduct each of the six techniques, and to collate and calculate the results from each. This has led to the conclusion that shear strength and load resistance are the quickest, followed by water content, bulk density, particle size mean and particle size sorting in that order - particle size mean and sorting being the slowest. Our assessments indicate that final results for shear strength and load resistance can be available in table form on disc, within about four hours of receiving the sediment cores in the laboratory. This contrasts with a period of about seven days for water content and bulk density, and about fourteen days for particle size mean and sorting. These times are based on one person conducting each of the tests, using the methodology and experimental protocols adopted in the present work. Robustness is here defined as the ability of a given geotechnical test to detect differences in sediment structure between high, medium and low trawling impact sites. In order to assess this, the data in table 7 were expressed as percentage changes in relation to the low impact site data for each of the geotechnical parameters. These percentages are given in table 8. They show clearly that shear strength and load resistance are the most robust of the tests used, and particle size mean and sorting are the least robust, in terms of percentage change from the low impact site value. For example, shear strength decreased by 52% at the medium impact site and 87% at the high impact site, when compared with the mean value at the low impact site. The equivalent changes for load resistance were a decrease of 47% and 86% respectively. The changes for bulk density were a decrease of 18% and 50%, and the changes for water content were an increase of 9% and 61%, at the medium and high impact sites respectively. The changes for phi mean particle size were an increase of 1 1% and 19%, and for phi sorting an increase of 3% and 5%. The ranked order based on the percentage changes at the medium and high impact sites when compared with the low impact site (table 8) is hence shear strength and load resistance (most robust), water content and dry bulk density, mean particle size, and particle size sorting (least robust). 78 Table 7. Summary table of trawling impact is assessed by changes in sedimentary parameters. Dry bulk density (g/ml), particle size (phi) mean and sorting, shear strength (kN/m2), load resistance (kN), water- content (%). Each mean and standard deviation is based on 24 values (two sites, four seasons : spring, summer, autumn, winter, and three sediment depths: top, middle and bottom). Trawling Impact DRY BULK DENSITY PARTICLE SIZE MEAN SORTING SHEAR STRENGTH LOAD RESISTANCE WATER CONTENT High 0.7204 5.998 1.821 0.4713 3.797 190.2 ±0.1948 ±0.7416 ±0.4457 ±0.4793 ±3.360 ±37.34 Medium 1.179 5.594 1.793 1.755 14.74 128.8 ±0.5461 ±0.5006 ±0.2634 ±3.198 ±22.24 ±67.69 Low 1.438 5.059 1.739 3.624 27.91 118.4 ±0.8611 ±1.053 ±0.2405 ±5.604 ±36.60 ±90.19 Table 8. Summary table of trawling impact as assessed by changes in sedimentary parameters. Dry bulk density, particle size mean and sorting, shear strength, load resistance, water content. Percentages calculated as follows: means in table 22 are expressed as percentage increases or decreases from the low trawling impact means (100%). SRI = Speed Robustness Index. Trawling Impact DRY BULK DENSITY PARTICLE SIZE MEAN SORTING SHEAR STRENGTH LOAD RESISTANCE WATER CONTENT High -50% -1-19% +5% -87% -86% +61% Medium -18% + 11% -1-3% -52% -47% +9% Low 100% 100% 100% 100% 100% 100% SRI 0.61 0.18 0.05 17.4 16.6 0.63 79 An Index of Speed and Robustness of the Geotechnical Parameters We have combined the assessments of speed and robustness of each of the techniques in a Speed Robustness Index (SRI) for each technique. The SRI is defined as follows: SRI = 1 a + b I /2c where: a = % increase or decrease in the observed parameter at the high impact site compared with the low impact site, b = % increase or decrease in the observed parameter at the medium impact site compared with the low impact site, c = hours taken to conduct the test, and to collate and present the data in hard copy and on disc, based on an eight hour working day. Application of the SRI to the percentages in table 8 and using the times taken to conduct the tests provides a quantitative assessment of the overall efficiency of the six tests used in terms of their speed and robustness. The SRI values are given at the bottom of table 8. Using the SRI it is clear that shear strength and load resistance are the best tests, and particle size and sorting are the worst tests, for the assessment of the impact of trawling for Neprhops norvegicus on sea bad structure in the Clyde Estuary. Shear strength and load resistance have the highest SRI's of 17.4 and 16.6. Water content and bulk density have SRI's of 0.63 and 0.61, and mean particle size and sorting have SRI's of 0.1 8 and 0.05. FUTURE WORK The geotechnical parameters used in the current investigation have demonstrated that differences in seabed structure attributable to Otter trawl impact can be detected by relatively simple and rapid technology. However the local characteristics of the sea bed in the area sampled may not be fully representative of the environments in which Otter trawling is used on continental shelves elsewhere. For the future, therefore, we suggest that the six geotechnical parameters used in this report are assessed in other high, medium and low impact trawling areas of the sea bed, to confirm their relative suitability and robustness in a wider range of environmental conditions. We recommend that the new corer designed by Dr Barnett should be developed to take four cores of a longer length. We strongly recommend that the load resistance equipment used in the current investigation is modified to quantify additional geotechnical parameters, and then developed for laboratory operation and for in situ operation on the sea bed with automatic data logging facilities in real time. In a broader context, it would now be interesting to use the same group of geotechnieal tests to assess sea bed disturbance in other areas, which could be linked to changes in benthic community structure and function caused by anthropogenic and natural sea bed disturbance. In a sense, this has already begun, as we have conducted similar work over a number of years in the Pacific and Atlantic, and also in the Arabian Sea These studies show clearly how geotechnical and geochemical parameters of the sea bed are related to the number and species of benthic organisms, benthic community structure and function, and more generally to the biodiversity of intertidal, near shore, and deep sea environments. ACKNOWLEDGEMENTS We thank Jim Atkinson and Geof Moore, University Marine Biology Station Millport, for inviting us to take part in the EC project. We thank Rupert Ormond, Director of UMBS, who has been very supportive and helped us in a number of ways. 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Meso- and micro-scale heterogeneity in benthic community structure and the sedimentary environment on an intertidal muddy-sand beach . Proceedings of the 22’^^ European Marine Biology Symposium, Barcelona. Scientia Marina 53: 319-327. 82 Glasgow Naturalist 2002. Volume 24. Part 1. Pages 83-91. A NEW SPECIES OF PARASITIC COPEPOD, SPHAERONELLA KEPPELENSIS N.SP. (SIPHONOSTOMATOIDA: NICOTHOIDAE), FROM THE AMPHIPOD ORCHOMENE NANUS (KROYER, 1846) IN THE FIRTH OF CLYDE, SCOTLAND. Myles O’Reilly Scottish Environment Protection Agency South West Area, 5 Redwood Crescent, Peel Park, East Kilbride G74 5PP e-mail: mvles.oreillv@seDa.org.uk INTRODUCTION The copepod genus Sphaeronella Salensky, 1868 contains over 70 species which live as parasites mainly within the marsupium of marine peracarid crustaceans. Over 40 species are recorded from amphipods (Costello & Myers, 1989) with around a dozen of these from amphipods in British Waters (Gotto, 1993, O’Reilly & Geddes, 2000, O’Reilly et a/., 2001). In 1993-94 during a study of the amphipod Orchomene nanus (Fam. Lysianassidae) undertaken at the University Marine Biological Station, Millport, a small number of the amphipods were found to be infested with copepods which were tentatively identified as S.callisomae Scott, 1904 - a species inadequately described almost 100 years ago by Thomas Scott. Of 990 amphipods examined only 8 were found to be parasitised (Moore & Wong, 1996). As the copepod has never been recorded since its original description, the opportunity was taken to undertake a full description of the female, the first description of the male, and also the first description of the copepodite stage. On completion of the descriptive work some new S.callisomae material was recovered from the amphipod Scopelocheiros hopei (Fam. Lysianassidae) collected in Loch Riddon, in the Firth of Clyde. This amphipod was the original host species of S.callisomae and examination of the new material, a mature female and 3 copepodites, revealed an excellent match to Scott’s original description. However, it also indicated that the copepod from Orchomene nanus actually exhibited significant differences from S.callisomae, and indeed from all other known Sphaeronella and thus it was considered necessary to establish a new species. The new species is described here. Its congener, S.callisomae, will be re-described in a subsequent work. METHODS The hosts and parasites were fixed in 1 0% formalin before transfer to methylated spirit for storage. The copepods were examined as temporary whole mounts in lactic acid or permanent mounts in polyvinyl lactophenol. Drawings were executed at magnifications up to x 1,000 with the aid of phase contrast and a camera lucida drawing tube. Host nomenclature follows Costello & Myers (1989). Sphaeronella keppelensis n.sp. Material examined All collected at Keppel Bight, Millport, Isle of Cumbrae, Firth of Clyde (55° 45.75’N, 04°54.48’W), depth 5-6m. (Each sample represents a single host amphipod) Material deposited in the National Museum of Scotland, Edinburgh: Sample T33 C9 - 19/4/94 - Holotype - 1 female, mature + 1 ovisac (NMSZ 2002. 111.1) Sample T24 B8 - 9/12/93 - Allotype - 1 male, mature, mounted on slide (NMSZ 2002. 1 1 1 .2) Sample T7 A3 - 20/5/93 - Paratype - 1 female mature, cephalon dissected and mounted on slide (NMSZ 2002.111.3) Sample T19 B1 - 22/9/93 - Paratype - 1 female immature (NMSZ 2002.111.4), 9 copepodites mounted on slide (NMSZ 2002.11 1.5), 2 unmounted copepodites (NMSZ 2002.111.6-7), and host amphipod ( Orchomene nanus) Sample T26 B3 - 22/2/94 - Paratype - 1 female mature + 4 ovisacs (NMSZ 2002. 111.8). Additional material retained in author’s collection: Sample T21 B1 - 26/10/93 - 4 ovisacs (copepod missing!). Sample T22 B3 - 9/11/93 - 1 female, mature (mounted on slide, poor condition), 1 copepodite dissected and mounted on slide. Sample T24 B8 - 9/12/93 - 1 female mature (poor condition) -f- 3 ovisacs, 1 female, immature, mounted on slide. Etymology: Named after the type locality - Keppel Bight. DESCRIPTION Female (Fig. 1, a, b): Body subspherical comprising a large spherical trunk and a small protruding head. Maximum length 0.85 mm, maximum width 0.72 mm. Ovisacs, irregularly ovoid, detached from female, diameters 0.4-0. 5 mm. Cephalon (Fig. 2. a) with naked frontal margin, lateral border margins with fine hairs. First antennae, A.l (Fig. 2.a,c) relatively short, 3- segmented; proximal segment with 3 setae distally; second segment much shorter without setae, third segment with 9 setae and an aesthetasc (chemosensory appendage). Second antennae, A.2 (Fig. 2.a,b) reduced, comprising a single segment surmounted by a long seta. Oral disc (Fig. 2. a) encircled with fine hairs, with central aperture 83 through which the tips of the mandibles, Md, are visible. First maxillae, Mx.l (Fig. 2. a) with 2 long filamentous processes directed anteriorly and a shorter filament on the inner edge. Second maxillae, Mx.2 (Fig. 2.a,d) 2-segmented; proximal segment (syncoxa) robust with stout median process on posterior surface and a row of spinules adjacent to articulation with distal segment; distal segment (basis) forms a curved claw, surmounted with fine curved spine-like process. Maxillipeds, Mxpd (Fig. 2. a) 3-segmented, first segment stout, elongate, with transverse row of fine setules proximally and another row medially on anterior face, second segment considerably smaller with short spine distally on inner edge, third segment curved with minutely tridentate tip. Ratio of maxilliped segments approximately 5:2: 1.5. Maxillipeds in paratype (Fig. 2.e) apparently with shorter first segment and 2 spines distally on second segment. The sub-median skeleton (Fig. 2. a) is just visible (dotted) extending back from the base of the first maxillae. Two distinct parallel chitinous ridges are visible between the second maxillae and extend between the base of the maxillipeds to form acute triangular processes. A weak transverse chitinous bar occurs posterior to the maxillipeds. Trunk (Fig. l.b) with small spinules scattered throughout. Legs minute (Fig. 3.a,b), consisting of a single short segment with 1 -2 setae apically. In the first legs, L. 1 , the setae are approximately equal in length but the inner seta is much stouter than the outer. Second leg, L.2, a little smaller, the setae are of similar thickness but the outer one is about twice the length of the inner, although 1 seta is sometimes missing. First legs occasionally with an extra seta (Fig. 3.c). The genital area (Fig. 3.d), is mostly covered with spinules through which a pair of oval seminal receptacles and semi-circular genital apertures are visible. The spinules extend posteriorly between the genital apertures towards the caudal rami. The caudal rami (Fig. 3.d,e,f) are short with pronounced conical process on the distal inner edge, and each ramus is ornamented with 2 naked setae, which are about 2-3 times the length of the ramus. Male: (Fig. l.c, 2,g) Body small, rhomboidal, maximum length 0.28 mm, maximum width 0.23 mm, head and trunk region each comprising around half of the body. Anteriorly the head is produced medially into a bilobed pseudorostrum. It has pronounced lateral lobes between the pseudorostrum and the first antennae. Lateral lobes, naked ventrally but covered in fine spinules dorsally. The spinule patch is observable even when viewed ventrally through the lateral lobes giving the lobes a dotted appearance. A tuft of these spinules extends around ventrally on either side at the base of the pseudorostrum. The cephalic shield forms a cape, which partly conceals the base of the second maxillae. The shield margins are ornamented with fine spinules. The exterior border (“shoulder”) area is covered with numerous conspicuous nodules. First and second antennae, oral disc, and first and second maxillae, apparently similar to female, although some details difficult to observe. Maxillipeds similar to female but first segment stouter and with row of fine setules distally at articulation with second segment, and third segment claw is strongly bidentate. The chitinous processes between the base of the maxillipeds are produced posteriorly into two strong spines. The trunk region is densely covered with long spinules, except for its anterior part. The legs are reduced and very difficult to see. The first legs are mostly concealed by the base of the maxillipeds. The interpretation shown (Fig. 3.h) was gained by focusing through the maxilliped and may not be entirely accurate. Apparently comprising an inner an outer branch with a short rounded lobe between them. The branches have 2 and 3 short apical setae respectively with a longer seta at the base of the inner branch and also on the medial lobe. The second legs (Fig. 3.i) are party obscured by the dense abdominal spinulation. They comprise 2 short simple branches, the inner a little shorter, and each branch with a stout apical seta. The large oval paired spermathecae are visible through the body wall between the legs. The caudal rami are placed close together at the posterior end of the body. Their structure is similar to the female with the distinctive conical process, though the seta are a little longer. Copepodite: (Fig. 4. a) Length 0.19 mm (excluding caudal setae), width 0.15 mm, cephalothorax oval, dorsally shield like, covering almost the entire body, only the last 2 segments of the urosome and the caudal rami protruding. The dorsal surface is finely granular with a transverse suture around the midpoint. The dorsal margin has 4 pairs of long setules positioned symmetrically, on anterior margin, and laterally level with the oral cone, mid-dorsal suture, and legs. Frontal border with naiTow rostral brim, and a pair of shorter setules ventrally. Lateral margins slightly in-rolled ventrally. First antennae(Fig. 4.b) similar to adult but setae and aesthetasc on third segment much longer. Second antennae (Fig. 4.c) 4-segmented with 2 terminal setae on distal segment. Oral disc and first maxillae apparently similar to adult. Second maxillae (Fig. 4.d) 3-segmented. First segment stout with row of fine setules, second segment narrow elongate, third segment forming a curved claw. Maxillipeds (Fig. 4.e) 4-segmented, first segment naked, greatly enlarged, second and third segment considerably smaller, third segment with a long seta distally which lies parallel to (and is around 75% the length of) the fourth segment which forms a long slender claw. Ratio of maxilliped segments approximately 9:1:1 :6. 84 a Figure 1: Sphaeronella keppelensis n.sp. - a) Paratype - Female gravid (T.26 B3) dorsal, with ovisacs, b) Holotype - Female gravid (T.33 C9), habitus ventral c) Allotype - Male (T.24 B8), habitus, ventral, to scale, d) Copepodite (T.19 Bl) - habitus, ventral, to scale. 85 Figure 2: Sphaeronella keppelensis n.sp. - a) Holotype - Female, mature (T.33 C9) - cephalon, ventral, Md.- mandible, A. 2 - second antenna, A. 1 - first antenna, Mx.l - first maxilla, Mx.2 second maxilla, Mxpd.- maxilleped, b) Paratype - Female (T.7 A3) - right & left second antennae (A. 2), c) Female (T.24 B8) - first antenna (A.l), left, d) Paratype - Female (T.7 A3) - second maxilla (Mx.2), left, anterior view, e) Paratype - Female (T.7 A3) - maxilliped (Mxpd.). 86 Figure 3: Sphaeronella keppelensis n.sp. - a) Holotype - Female (T.33 C9) -F' and 2"'* leg pairs, b) Paratype - Female (T.26 B3) -1'‘ and 2"'' leg pairs, c) Female (T.24 B8) -F' leg right, d) Holotype - Female (T.33 C9) - genital area and caudal rami, e) Paratype - Female (T.26 B3) - caudal rami, f) Female (T.24 B8) - caudal ramus, right, g ) Allotype - Male (T.24 B8), habitus, ventral, right A.2 and Mx.l omitted, h) Male (T.24 B8), first leg, left, interpretation, i) Male (T.24 B8), second leg, left, interpretation. 87 Figure 4: Sphaeronella keppelensis n.sp - Copepodite (T.19 Bl) a) habitus, ventral, (A1 & leg setae truncated), b) first antenna (A.l), right, c) second antenna (A. 2), right, d) second maxilla (Mx.2), left, e) Maxilliped (Mxpd.), left, f) left leg, rami & setae folded inwards, g) left leg, rami & setae folded outwards. 88 A medial prosomal swelling forms an approximate hexagonal shape between the maxillipeds and the legs. It has a smooth surface and is without furrows exhibited by copepodites in many other species of Sphaeronella. There are 2 pairs of biramous legs (Fig. 4.f,g) which are similar in structure, each pair joined ventrally by a narrow intercoxal plate. They have a long basal segment with a single long seta distally on the outer edge. The inner ramus (endopod) is very small with 3 long plumose setae. The outer ramus (exopodite) is a little larger with 4 long plumose setae and 2 shorter naked setae on outer edge. The first legs are incorporated into the cephalosome. The somite of the second legs expands laterally and posteriorly where it form the rear edge of the cephalothorarax. The urosome comprises 3 segments of decreasing size, and a pair of caudal rami. The first segment has 2 pairs of setae distally on the outer edges. The second and third urosomal segments are un- omamented. The caudal rami each have 3 relatively short lateral setae and an extremely long inner seta, which is around 75% the length of the cephalothorax. DISCUSSION The S.keppelensis material was initially attributed to S.callisomae on the basis of the unusual combination of a 3-segmented maxilliped with 3- segmented first antennae and 1 -segmented second antennae. Its utilisation of a lysianassid host, and its occurrence in the same Sea area added weight to this assertion. However, Scott’s description of his single female of S.callisomae does show some significant differences from S.keppelensis. The most notable of these is the presence of a “conspicuous tubercle” projecting from the frontal border of the cephalon of S.callisomae. In addition its first antennae has only a single setae on the first segment, and a longer third segment with 7 setae (and no aesthetasc). The size and shape of the first and second maxillae also appear to be slightly different and the maxilliped has a shorter basal segment with some spinulation at the articulation with the second segment. Preliminary examination of the new S.callisomae female from Loch Riddon substantiates these differences. Bradford (1975) tabulated the then known species of Sphaeronella described from amphipods (41 spp.), isopods (7 spp.), cumaceans (7 spp.), and ostracods (17 spp.) and arranged them into groups based on key features of their morphology. Among these she recognised one group of eight amphipod-infesting species, where the females have a 3-segmented maxilliped (and 3-segmented first antennae), as shown in Table 1. This group which can informally be called the “S.giardii Group” is also characterised by males in which the legs are relatively small with short setae (although males are only known for S.giardii, S.bonnieri, S.aeginae, and S.longipes). The inclusion of S.valida in the group is based on the re-description by Green (1958) who figures a 3- segmented maxilliped, although the original figure of Scott (1905) is clearly 4-segmented. Among five species considered by Bradford as “not well described” and not placed in any group were S.callisomae and S.cluthae both described from the Firth of Clyde by Scott (1904). They are depicted by Scott with 3-segmented maxillipeds suggesting they ought to belong to the '‘S.giardii Group”. The description and figure of the maxilliped of a third species, S.pilosa Blake, 1929, is unclear although it shares the same host species (and geographical location) as S.photidas. Table 1. Sphaeronella species of the “5. giardii Group” S. giardii Hansen, 1897 Host Protomedeia fasciata Locality Denmark S. bonnieri Hansen, 1897 Host Protomedeia fasciata Locality West Greenland S. longipes Hansen, 1897 Host Ampelisca tenuicornis Locality Denmark, British Isles S. amphilochi Hansen, 1897 Host Paramphilochoides odontonyx Locality Denmark, British Isles S. dulichiae Hansen, 1897 Host Dyopedos monacanthus Locality Denmark S. valida Scott, 1905 Host Megamphopus cornutus Locality British Isles S. aeginae Hansen, 1923 Host Aeginina longicornis Locality Iceland / Faroes S. photidis Blake, 1929 Host Photis reinhardi Locality New England, USA It is clear that S.keppelensis should also be included in the "S.giardii Group” based on the female maxillipeds and reduced male legs. In addition to the 3-segmented maxilliped S.keppelensis possesses two features which are unusual within the group; the 1 -segmented second antennae and the conical process on the caudal rami. These characters are exhibited by both sexes. The 1 -segmented second antennae appears to be shared only with S.callisomae whilst the conical process appears to be unique within the group. It is impossible to compare adequately the new S.keppelensis material with S.cluthae, known by a single female from the amphipod Harpinia pectinata (Fam. Phoxocephalidae). Only the habitus (lateral and dorsal) and maxilliped of S.cluthae were figured, although the latter is not inconsistent with S.keppelensis there is insufficient information overall to draw any inference regarding the affinity of S.cluthae. 89 The male S.keppelensis has additional features which also appear to be unique within the ''S.giardii Group”, notably the pronounced bilobed pseudorostrum and lateral lobes of the cephalon and also the conspicuous nodules on the exterior border area. The male has 3-segmented maxillipeds in common with the female but this does not appear to be the case in all males of the "'S.giardii Group” as S.giardii and S.aeginae males are figured with 4- segmented maxillipeds. The legs of the male S.keppelensis appear to be generally similar to S.bonnieri. The larval or copepodite stage has only been described for around a third of Sphaeronella species; 10 from amphipods, 3 from isopods, 5 from cumaceans, and 9 from ostracods (including 2 among the latter for which only the copepodite stage is known). In almost all Sphaeronella copepodites the entire urosome protrudes beyond the cephalothorax, the only exception being S.bradfordae Boxshall & Lincoln, 1983 (an isopod parasite) where only the setae of the caudal rami protrude. S.keppelensis falls between these two extremes with just the tip of the urosome visible in dorsal view. The legs of the S.keppelensis copepodite are also unique with their reduced rami bearing only plumose or shorter simple setae. All other described Sphaeronella copepodites have legs with elongate rami (as long as or longer than the basal segment) and possess both plumose setae on the inner edge and spines or simple setae on the outer edge. Similar reduced rami are present in the copepodite of Sphaeronelliodes vargulae Bradford, 1975, sole representative of an allied genus from an Antarctic ostracod, but the basal segments of its legs are much shorter and its urosome is only 2-segmented with the caudal rami fused to the last segment. These, along with other unusual features of the mature female Sphaeronelliodes, warranted its separation as a distinct genus. It is apparent then that S.keppelensis is clearly different from any other described Sphaeronella species with both the female and male, as well as the copepodite stage, exhibiting distinctive morphologies. Most Sphaeronella appear to be relatively host- specific occurring in only one or two allied host species. Orchomene nanus and Scopelocheirus hopei occupy a similar niche as scavengers and were recently found to be the dominant amphipods in carrion-baited traps in the Firth of Clyde (Bergmann et al., 2002). In view of their close association, both phylogenetically and ecologically, it is not surprising that their copepod parasites may also be closely related. Only two other Sphaeronella species have been described from lysianassoid amphipods: S.norvegica Hansen, 1905 from Tmetonyx similis collected in Norway and S. australis Boxshall & Harrison, 1988 from the genus Amaryllis collected in Tasmania. However, both of these have 4-segmented maxillipeds excluding them from the ^"S.giardii Group”. ACKNOWLEDGMENTS I am greatly indebted to Professor Geoff Moore and Mr Tym Wong of the University Marine Biological Station, Millport, who kindly supplied the parasitised material of Orchomene nanus. REFERENCES Bergmann, M., Wieczorak, S.K., Moore, P.G., & Atkinson, R.J.A. (2002). Utilisation of invertebrates discarded from the Nephrops fishery by variously selective benthic scavengers in the west of Scotland. Marine Ecology Progress Series, 233, 185-198. Blake, C.H. (1929). New Crustacea from the Mount Desert Region. In: Proctor(1933)(ed.), Biological survey of the Mount Desert Region, The Wistar Institute Press, Philadelphia, Part 3, 1-34. Bradford, J.M. (1975). New parasitic Choniostomatidae (Copepoda) mainly from Antarctic and Subantarctic Ostracoda. New Zealand Oceanographic Institute, Memoir No.67, 1-36. Boxshall, G.A. & Lincoln, R.J. (1983). Some new parasitic copepods (Siphonostomatoida: Nicothoidae) from deep-sea asellote isopods. Journal of Natural History 17, 891-900. Boxshall, G.A. & Harrison, K. (1988). New nicothoid copepods (Copepoda: Siphonostomatoida) from an amphipod and from deep-sea isopods. Bulletin of the British Museum, Natural History (Zoology) 54(6), 285- 299. Costello, M.J. & Myers, A. A. (1989). Observations on the parasitism of Aora gracilis (Bate) (Amphipoda) by Sphaeronella leuckartii Salensky (Copepoda), with a review of amphipod- Sphaeronella associations. Journal of Natural History 23, 8 1 -9 1 . Gotto, V. (1993). Commensal and Parasitic Copepods associated with Marine Invertebrates (and Whales). Synopses of the British Fauna (New Series) No. 46. Published for the Linnean Society and the Estuarine and Coastal Sciences Association by Universal Book Services/Dr. W.Backhuys, 264 pp. Green, J. (1958). Copepoda parasitic on British Amphipoda (Crustacea), with descriptions of a new species of Sphaeronella. Proceedings of the Zoological Society of London 131,301-313. Hansen, H.J. (1897). The Choniostomatidae. A family of Copepoda, parasites on Crustacea Malacostraca. Pp.l- 206, pi. 1-13. A.F.Host & Son, Copenhagen Hansen, H.J. (1905). Two new forms of Choniostomatidae: Copepoda , parasitic on Crustacea Malacostraca and Ostracoda. Quarterly Journal of Microscopical Science 48, 347-358, pl.22. Hansen, H.J. (1923). Crustacea, Copepoda. II. Copepoda parasita and hemiparasita. Danish Ingolf-Expedition 3,(7), 1-92, 5pl. Moore, P.G. & Wong , Y.M. (1996). A note concerning associates of the scavenging amphipod Orchomene nanus (Lysianassoidea). Journal of the Marine Biological Association of the U.K. 76, 259-261. O’Reilly, M.G. & Geddes, D. (2000). Copepoda. pp.217- 281, in Vol. 1 of: Foster-Smith, J. (ed.)(2000). The Marine Fauna and Flora of the Cullercoats District: Marine Species Records for the North East Coast of England. Vol.l (546pp.), Vol. 2 (561pp.) A Dove 90 Marine Laboratory Publication, Penshaw Press, Sunderland. O’Reilly, M., Hamilton, E. & Heaney, L. (2001). New records of amphipods and leptostracans from the Forth Sea area, with notes on their copepod parasites (Siphonostomatoida; Nicothoidae). Glasgow Naturalist 23(6), 35-42. Scott, T. (1904). Notes on some rare and interesting marine Crustacea. 22'"' Annual Report of the Fishery Board for Scotland, 1903, Part 3, pp. 242-261, plates 13-15. Scott, T. (1905). On some new and rare Crustacea from the Scottish Seas. 22"“' Annual Report of the Fishery Board for Scotland, 1904, Part 3, pp. 141-153, plates 10-13. 91 ' ft Glasgow Naturalist. 2002. Volume 24. Part 1. Pages 93-103 SHORT NOTES Glasgow Naturalist. 2002. Vol 24. Part 1. 93 PILL MILLIPEDE GLOMERIS MARGINATA ON JURA Glyn M. Collis & V. Dawn Collis Brookside House, Fosse Way, Moreton Morrell, Warwick, CV35 9DF. The Pill Millipede Glomeris marginata (Villers) is common and widespread in England, Wales and Ireland, extending to Southern Scotland. It is well known for its habit of rolling into a ball in a manner similar to the Pill Woodlouse Armadillidium vulgare (Latreille). Blower (1985) reports that it is not recorded north of the firths of Clyde and Forth except for one record from Wester Ross (VC 105). This record is presumably unverified since an updated distribution map recently published by the British Myriapod Group (Newsletter number 32, Spring 2000), indicates no records north of the Clyde-Forth line. Mr Blower’s records are included in the database from which this map is derived. The only record shown from a Scottish Island is a single 10km square on Arran (Hancock, 1991). In July 2000 we found G. marginata at two sites on the Isle of Jura. One record was from a garden in Craighouse, NR526671, the principal settlement on Jura. The garden was not one that has received much attention other than a regularly mown lawn and some basic tidying. The range of garden plants was small although there were some well- established Fuchsias plus other shrubs. G. marginata was found among old garden rubbish under the shrubs. The other site was low moorland, about 100m from the sea at McDougall’s Bay NR443680, not close to human habitation but beside a loop of road that was abandoned when the “main” road was straightened. A short stretch of mortared wall bordered the old road where it crossed a bum, and G. marginata was found among mbble at the base of this wall. References Blower, J. G. (1985). Millipedes. Synopses of the British Fauna (new series) No. 35. The Linnean Society and the Estuarine and Brackish-Water Sciences Association, London. Hancock, E.G. (1991). Pill Millipede on Arran. Glasgow Naturalist 22, 84-85. Glasgow Naturalist. 2002. Vol 24. Part 1. 93 PLA TYARTHRUS HOFFMANSEGGI IN KIRKCUDBRIGHT (VC 73) V. Dawn Collis & Glyn M. Collis Brookside House, Fosse Way, Moreton Morrell, Warwick, CV35 9DF. Platyarthrus hoffmanseggi Brandt is a small blind white woodlouse, well known as an inhabitant of ants’ nests in southern Britain. In Harding & Sutton (1985) the only Scottish site recorded for this species is Inverkeithing, Fife (VC 85), with recent records from the same locality where it was found at the turn of the last century by Evans ( 1 900). On 24th October 1999 P. hoffmanseggi was found by us in a nest of yellow ants under a stone on coastal grassland at Knockbrex NX578498. With a favourable climate, the Solway coast is known to support a number of species that generally have a southerly distribution in Britain, including woodlice of the genus Armadillidium (Harding, 1975). We found Armadillidium vulgare (Latreille) at Knockbrex just a few metres away from the P. hoffmanseggi site. References Evans, G. (1900). Platyarthrus hoffmanseggi Brandt in Fife. Annals of Scottish Natural History 35, 186. Harding, P.T. (1975). Armadillidium in South-west Scotland. Glasgow Naturalist 19 (3), 175-177. Harding, P.T. & Sutton, S.L. (1985). Woodlice in Britain and Ireland. Institute of Terrestial Ecology, Huntingdon. Glasgow Naturalist. 2002. Vol 24. Part 1. 93-94 LIME TREE (TILIA SPP.) REGENERATION 2001 R.K.S. Gray “Langdale”, 6 Prince Albert Road, Glasgow G12 9JX Previous papers in this Journal have drawn attention to the phenomenon of regeneration of lime trees by seed. Readers will therefore be aware that we are near the northern limit of such natural regeneration in these islands and that consequently its occurrence is worthy of record. They will also realise that naturally occurring lime trees in this country are either large leaved (T. platyphyios), small leaved (T. cordata) or common (T. X europaea) and that successful fertilisation of the small leaved lime is more temperature sensitive than that of the other two. Also it is important to realise that a suitable temperature at the time of fertilisation is not the only factor that determines successful fertilisation and subsequent germination of lime trees. The monthly mean temperature maxima for July and August of 1999 were 19.9‘’C and 19. iV respectively. The summer temperatures 1.5 years before germination are considered to be critical in determining fertilisation and subsequent germination of lime seed. The 2001 numbers for T platyphyios and T x europaea are comparable to those of 1999 when the corresponding 1997 temperature figures were 20.3 and 21.6°C. These increased amounts of lime regeneration at these latitudes are in keeping with the general trend of climatic warming. Regeneration has been observed in Glasgow’s West End at the time of writing in April 2002 and a report has been received of regeneration in Milngavie. Observations have also been made of a few survivors in addition to those collated below. Readers are requested to forward information about observations they make to R. Gray. 93 T. platyphyllos T. cordata T.x. europaea Glasgow West End 134 1 225 Outwith Glasgow 6 - 2 Survivors 13 1 5 The table is a summary of the numbers of seedlings and survivors (seedlings which have lived through at least one winter since germination) found by or reported to us in Scotland in 2001. This report therefore extends the records obtained each year since 1997. Acknowledgements N.R. Grist, M.H. Hansen and P. Macpherson of the Society and M. Matthews of the Met. Office. Reference Gray, R.K.S. & Grist. N.R. (2001). Natural Regeneration of Limes (Tiliaspp.) in Scotland. The Glasgow Naturalist 23 (6), 19-21. Glasgow Naturalist. 2002. Vol 24. Part 1 . 94 ORANGE LADYBIRD Halyzia 16-guttata IN HYNDLAND Norman R. Grist 5A Hyndland Court, 6A Sydenham Road, Glasgow G12 9NR The short note in the last Glasgow Naturalist (Putter and Putter 2001) drew my attention to the interest of my own sighting of a single orange ladybird on May 22, 1999. I noticed this unusual ladybird in our garden, on a rose leaf I think, one sunny afternoon, collected and photographed it and released it next morning. The photographs were shown to E.G. Hancock, more recently to Richard Weddle and Richard Sutcliffe, and the record with photographic evidence was duly entered as the first in Glasgow for this species in Glasgow Biological Records. The possible association of the species with sycamores was mentioned by Putter and Putter (2001), and there is one mature sycamore in the garden about 12 meters from where I found it. The orange ladybird is primarily a mildew feeder (Majerus 1994), but aphids and honeydew are listed as secondary foods. Most trees in our urban garden are common limes (Tilia x europaea) which usually generate much honeydew and black mould on vegetation below, and did so in 1999. Are these handsome ladybirds increasing in this northern latitude? References Putter, S. & Putter, K. (2001). Orange Ladybird Halyzia 16-guttata (L.) in Dumbarton. The Glasgow Naturalist 23 pt.6, 120. Majems, M. (1994). Ladybirds. Harper Collins, London. Glasgow Naturalist. 2002. Vol 24. Part 1. 94-95 SUMMERTIME SWIFTS Norman R. Grist 5A Hyndland Court, 6A Sydenham Road, Glasgow G12 9NR Por me summer in Glasgow begins when swifts (now ’Common. Swifts ’) Apus apus arrive. My first sightings this year (2001) as usual were from the roof garden of our Hyndland flat (NS560 675). I see them clearly silhouetted against the dusk sky over Gartnavel chimney tower, Bingham’s Pond and greened areas to its south. This location appears to act as a convenient muster point for them to gather each evening before flying up to perhaps 3000m in order to miero-sleep on the wing without collisions and heading into any wind to avoid drifting too far from their starting point (Bromhall 1980; Blackman 2001). We wonder how they judge the direction of wind when aloft in the dark without reference points during night when it is not always clear and unclouded. They numbered 3 on May 11th (3 days later than last year’s arrival, also of 3). Those last seen also numbered 3 on August 20th (2 on Aug. 18 2000) - their departure signalling the end of summer. Their numbers rose to 22 by the end of May with 1 6 the average maximum in July - August. These totals were somewhat fewer than last year when 24 to 30 were seen in July. Maximum counts in earlier years were 25 (‘94), 28 (‘95), 20 (‘96-97), 18 (‘98), 20+ (‘99). It is always a challenge to count these rapidly moving birds about half a kilometer away as they wheel individually and in groups. Most fly in loose pairs and show up black and clear against the bright dusk sky, becoming almost invisible when they turn head-on or tail-on to the observer, to reappear as they bank side-on. Patient scrutiny with the binoculars for several minutes can eventually achieve a reasonably steady count, avoiding double-counting as far as possible and confirmed by consistent counts over several consecutive evenings. The numbers in early weeks are modest, around 8 to 12, increasing to roughly double in July, then falling in August to about the starting numbers. Perhaps this represents the addition of the young birds of the year that in August fly back to Africa unaccompanied. This leaves their parents free of responsibility to enjoy some remaining weeks here on their own before they too return to Africa to escape our winter. That the youngsters are genetically programmed to migrate on time to the right place without parental guidance is one of those amazing things. Another is the speed with which parental mating and egg laying, and the hatching, fledging and achievement of flight by the youngsters are completed in the few weeks of our midsummer - a tribute to the quantity and quality of our ‘aerial plankton’, mainly invertebrates windbome over the city. Clare Darleston, Coordinator of ‘Concern for Swifts (Scotland)’, has drawn attention to the problems of urban swifts with fewer nesting places available in most of the city. My own observations do not throw much light on this problem, but I hope that my systematic observations and records may 94 help to document the population trends in this area near Anniesland where she identified some nests. References Blackman, S. (2001). Life’s a Breeze. BBC Wildlife 19 no. 7: 17, citing Backman & Alerstam. Proceedings of the Royal Society of London Series B 268: 1081-7. Bromhall, D. (1980). Devil Birds, The Life of the Swift. Hutchinson. Glasgow Naturalist. 2002. Vol 24. Part 1 . 95-96 URBAN FOXES IN HYNDLAND, GLASGOW Norman R. Grist 5A Hyndland Court, 6A Sydenham Road, Glasgow G12 9NR During the time we have lived here (since 1970) or in nearby Lorraine Rd. (1954-70) we saw no fox until the severe winter of 1996 when we saw a pair in the front garden at 7.30 am on January 3 0th. A neighbour reported seeing two “wolves" a few days before. Having prospected the territory, the foxes adopted it and established a nearby den. Since then they have been resident, breeding, scavenging and interacting with us humans, with cats, squirrels & magpies. Casual observations have made us familiar with the favoured routes and behaviour of the foxes, varying as generations succeeded one another. We have made use of their scavenging for eco-friendly disposal of kitchen scraps, left-over foods (bonanzas around Christmas!) sometimes supplemented by cat-food, dog-food or other items. Pasta & potatoes are ignored, but biscuits, peanuts & cake are readily accepted, and we understand that the foxes enjoyed packet dates put out by a nearby neighbour. This sophisticated town food seems to suit them well - their condition has remained splendid, contrasting with the scrawny foxes seen around Glasgow Airport by Riccardo Lazzarini. Their numbers remained fairly stable with surviving litters of one to three. Sightings recorded in Table 1 were not from specific, planned observations but from casual sightings from windows or from ground level (garden, car parking, garage), sometimes provoked by hearing the characteristic harsh, high-pitched barking. Peak sightings were in June > July > September. Some encounters were close-range with foxes who showed little fear but much curiosity about our actions and the chance that we might have food. One fox tried to snatch a plastic bag of cake crumbs I was putting out one evening for birds next day. The plastic slipped from his jaws. 1 tossed him a couple of fragments to show it would not interest him (I thought) but they were gobbled with pleasure - however, I have not taken to buying cake for foxes! On average we put out food scraps, often supplemented with a few dog or cat biscuits, which vanish each night - even occasional quite large bones, which disappear completely - not gnawed as playthings as a dog might do. Table 1: Times seen/Number seen of foxes, by quarter Year Jan- April- July- Oct.- Notes March June Sept. Dec. 1996 2/2 25/ 26/ 5/ Mobbed by 30 32 3+ mapies, June. Dish cleared nightly, Nov. 1997 4/5 5/6 44/ 13/ [“(+)” = daily (+) (+) 52 15 sightings of young by neighbours next door] 1998 10/11 9/9 12/ 9/ Grazing 15 10 peanut/biscuit; Cat spitting standoff in May 1999 11 / 12 14/ 14/ 11/ 17 April 16 17 11 03.00h, squabble, chased off by white cat 2000 6/6 20/ 29/ 15/ 22 35 16 2001 5/5 26/ 19/ 9/9 3 June chased 35 25 etc. Interactions with other animals include squirrels, one of which clambered up our brick building when disturbed by a fox investigating peanuts dropped for the squirrel. Generally the squirrels avoid them without problems - just as they elude cats, often with contempt, confident in their wide field of vision, speed, agility - and claws. A young fox recently amused us by perching on the roof comer of a garage building to watch a squirrel, a few metres below, enjoying peanuts from a feeder on a tree trunk - so tantalising! Cats rarely chase a fox and a fox rarely chases a cat. After a noisy face-to- face encounter one night both withdrew without actual aggression. A neighbour's garden provides a sheltered spot for resting, sunbathing, for juniors to play with or without attendant vixen. A wall about 1.5m high provided a sheltered space when heavily overgrown with ivy, and one summer day Mrs Grist while gardening below was surprised to find a young fox staring down at her through the cover One successful vixen, "Stumpy" with half a tail, first seen in 1997, has reared several families, and showed prowess by chasing off the large white cat which is currently dominant among the felines. Stumpy in turn was chased one June night in 2001 at full racing speed along Linfern Road into our garden and vanished through the shrubby hedge. The large pursuing dog appeared but gave up and left with his owner. Stumpy reappeared and giving a series of loud, sharp calls - danger warning? summoning help? She went back through the hedge, but shortly her big mate Tippy" with the 95 unusually well-marked white tail tip came through, stared down the track, walked back towards the road & onto the comr of the garden wall to survey the scene for a few minutes. Satisfied all was well, he trotted off to forage on his own account. Another episode that year was when Stumpy led her two well-grown offspring up steps onto the garage roof and sat gazing out, apparently to show how this vantage point gave a wide view including the length of Linfem Road. The youngsters gambolled with one another, over and round Stumpy, and eventually all departed. Next morning from the stairs I noticed the two alone on the car park, gazing in opposite directions - "what to do next"? As 1 descended one strolled off behind our building. The other strolled a short distance in the other direction and watched me with relaxed interest as I emerged and opened our garage. It hesitated about approaching in case I had goodies, but decided not and strolled away. Next night again fox barking was heard in 3 directions - suggesting family contact as the youngsters dispersed. As in many other urban areas in Britain, foxes have thus become an established part of our local fauna, as a stable population presumably in equilibrium with local resources of food and shelter. In the joint national survey by the British Trust for Ornithology and the Mammal Society in the 1km squares of the Breeding Bird Survey, 45 species of mammal were recorded in 1995-2000 (Newson & Noble, 2002). Over the six years the population of red foxes was stable as the fourth most numerous and widely distributed species: the highest counts were for rabbits (1040), brown hare (526), grey squirrel (460), red fox and roe deer each 227 with the fox more widely distributed Reference Newson, S. & Noble, D. (2002). BBS Mammal Monitoring. BTO News 239, 11-12. Glasgow Naturalist. 2002. Vol 24. Part 1. 96 A RICH BOTANICAL SITE AT LEADHILLS P Macpherson 1 5 Lubnaig Road, Glasgow G43 2RY Just north of the village of Leadhills, Lanarkshire (VC 77) there is an artificial elongated mound between the road and the bum. It was presumably a slag heap fonned when lead was mined in the area. It is approximately 250 yards long and at the north end 25 ft higher than the road but with a 35 ft slope down to the burn. The road rises alongside the mound and is more-or-less at the same level at the southern end. The plateau is 30 yards wide at the north end and 15 yards at the south, and almost horizontal On the plateau there are small colonies of Frog Orchid {Coeloglossum viride), Moonwort {Botrychium lunaria). Field Gentian (Gentianella campestris), and a very small form of Lesser Meadow-me (Thalictrum minor si) which has not flowered over a four year period. In addition there are the dandelions Taraxacum argutum and T . subnaevosum (det. A.J. Richards & A. A. Dudman). Wall Whitlowgrass (Draba muralis) is to be found on the stony slopes and Downy Oat-grass {Helic to trichon pubescens) on the flat ground at the north end of the mound. All round the area there are carpets of a dwarf form of Water Avens {Geum rivale), the subject of a previous report (Macpherson 2000). It is, therefore, one of the best small sites for rare plants in Lanarkshire. The Frog Orchid is known from only one other extant site; Moonwort has nine other quadrant records, three of which are on abandoned coal bings; Field Gentian has only one other modem record, a flat area between adjacent coal bings; Lesser Meadow-rue is presumed to be native on rocks at the Falls of Clyde where the plants are much larger; T. argutum is the only VC 77 record and T subnaevosum the fourth. Wall Whitlowgrass is currently known only from a railway embankment at Elvanfoot and the Downy Oat-grass from eight quadrants. Reference Macpherson, P (2000). Abnormal water avens morphology- lead indueed? Glasgow Naturalist 23, 53- 54. Glasgow Naturalist. 2002. Vol 24. Part 1. 96-97 LESSER HAIRY-BROME AT THE FALLS OF CLYDE P & LMD Macpherson & J Waddell 1 5 Lubnaig Road, Glasgow G43 2RY Bonavista, Heatheryett Drive, Galashiels, Selkirkshire TDl 2JL In a contract survey of the vegetation of the Scottish Wildlife Trust (SWT) Falls of Clyde Wildlife Reserve, Averts (1997) reported the occurrence of Zerna (Bromopsis) benekenii (Lesser Hairy-brome) on gentle sloping woodland floor and steep banks in the neighbourhood of Corra Castle, Lanarkshire (VC 77). This plant had not been known there before and its presence in such an area was very surprising as the Falls of Clyde has probably been the most botanised part of Lanarkshire. Further, the SWT reserve has been the subject of a number of previous plant surveys. In 1999 JW located the site and sent a specimen to a Botanical Society of the British Isles grass referee (RM Payne) who confirmed that it was, indeed, B. benekenii. In 2000 a further plant from the site was sent as proof to Prof CA Stace so that it might be included for the vice-county in the Vice-comital Census Catalogue. This specimen was, in turn, passed on to a brome specialist (LM Spalton) who agreed with the identification. A search of the surrounding area has since been made (PM & LMDM) and two further colonies discovered which we consider to be B. benekenii. At one site of approximately 6x5 metres there were about 20 plants and at a larger one of 14 x 5 m almost 50 were counted. Associated species noted were Great Wood-rush (Luzula sylvatica). Wood Sedge (Carex sylvatica). Tufted Hairgrass 96 {Deschampsia cespitosa) and Bearded Couch (Elymus caninus). Some plants are close to a path and in the past have just been assumed, without thought to be B. ramosus (Hairy Brome) which is of widespread occurrence in the reserve and for which there are 27 quadrant records in the vice- county. We have consulted all the Scottish Floras and Checklists at our disposal and found records only in the Checklist of the Plants of Perthshire (Smith et al. 1992). They report that the plant occurs in Mid and E. Perth VCs 88 & 89, and instance such well known sites as Keltneybum, Birks of Aberfeldy, gorge near Craighall and Killiecrankie The plant is included in Scarce Plants in Britain (Stewart et al 1994), a Nationally Scarce Species in Britain being defined as being recorded in 16-100 10km squares. The publication gives five records for Scotland (VCs 88 and 89) and 29 for England and Wales. The author of the report in that publication (Newton) states that B. benekenii is largely confined to woods on shallow chalk, limestone or other calcareous soils in steep valleys, growing in small to medium patches and best on a gentle slope. He stated further, that it is a little known and probably under-recorded species which several competent recorders have found difficult to distinguish from B. ramosa with which it sometimes grows. Stace (1997) has also written that it is probably an overlooked species. In general, these criteria apply to its Falls of Clyde occurrence. In a Scottish context it is the rarest species in the reserve. The Rev John Lightfoot travelled in Scotland in 1772 and subsequently wrote Flora Scotica published in 1777. He referred to the "famous falls" and "celebrated falls" and repeatedly mentioned Corra Linn (with different spellings!), listing some of the plant rarities. This area, together with the falls at Bonnington and Stonebyres became well known for their flora in the 19'*’ century (Mackechnie 1958). The precise locality has, therefore, attracted attention, both from the scenic and botanical aspects for well over 200 years. We will now look more closely at colonies of “5. ramosus"' in similar sites in Lanarkshire! Acknowledgements We are grateful to Messrs Payne and Spalton for confirming the identification. References Averis, ABG (1997). The Vegetation of the Falls of Clyde Wildlife Reserve, Contract Survey for the Scottish Wildlife Tmst. Lightfoot, J (1777). Flora Scotica, London. Mackechnie, R (1958). Plant Recording in Clydesdale, Glasgow Naturalist: 18, 3-14. McMicheal.AC (un-dated- late 19* century). Notes By the Way: A Descriptive, Historical and Biographical Account of Lanarkshire, Henry, Ayr. Newton, AL in Stewart et al. (1994). Scarce Plants in Britain, JNCC, Peterborough. Stace, CA (1997). New Flora of the British Isles (2"“* edition), Cambridge University Press, Cambridge. Stewart, A, Pearman, DA & Preston.CD (1994). Scarce Plants in Britain, JNCC, Peterborough. Glasgow Naturalist. 2002. Vol 24. Pt 1. 97-98 PREDATOR/PREY RELATIONSHIPS IN AN URBAN ENVIRONMENT. A.Meadows, P.S.Meadows, &W.Orr Division of Environmental and Evolutionary Biology, Institute of Biomedical and Life Sciences, University of Glasgow, G12 8QQ The pyramid of numbers in any ecosystem is such that there are many organisms at the bottom of the pyramid - the photosynthetic primary producers, progressively fewer as one moves upwards through the secondary producers - the herbivores, and very few at the top of the pyramid - tertiary producers or the carnivores (Elton, 1927; Wynne Edwards, 1962). In a natural ecosystem away from the immediate impact of man the primary producers are the grasses, flowering plants and trees, the secondary producers are often grazing organisms such as deer, gazelles, wild goats, and giraffes (Krebs, 1972; Ricklefs, 1973). The top-level carnivores include the lions, wildcat, eagles and so on. Man imposes a pattern on this by agricultural practice that is expressed to an even greater degree by the construction of villages towns and cities. The city environment is the oddest ecosystem for any wild plant or animal. Much of it is made of buildings, there are many roads, and the gardens are often so well organised that they are semi-deserts as far as many of our natural flora and fauna are concerned. The top predators are clearly ourselves, but a number of other animals that sit at or near the top of the ecological pyramid have managed to intrude. These include urban foxes, cats - feral or domestic, magpies, and to a lesser extent dogs. So any records of top predators and their interactions with other animals in towns and cities are really very important in enabling us to understand how our somewhat unusual city environment functions in an ecological sense. We record here two such instances. The first was an avian interaction. It involved three species of birds - the Kestrel, the pigeon and the magpie. The second involved a mammalian avian confrontation. It involved magpies and squirrels, which occurred four years ago.. On the morning of a Saturday in April, 2003, we witnessed an incident outside our ground floor kitchen window at The Mews, 2 Prince Albert Road, Dowanhill, Glasgow. A large bird hit another bird in flight, and then landed in the overhang outside our front door. It was possible to view and photograph the bird through the glass of the front door. It was a Kestrel with a newly killed young pigeon in its claws. The pigeon was photographed. It had the whole of its under surface ripped, with exposure of the two pectoralis major muscles. These are the two muscles attached between the sternum and the humerus bone of the wings, one on each side. They produce the downbeat of the wing that allows a bird to fly. For those nonvegetarians 97 amongst us, they are also very good eating. Clearly the impact of the Kestrel with the pigeon in flight had killed the pigeon instantaneously The Kestrel left the dead pigeon momentarily, jumped onto a wooden half barrel containing earth, and began to clean its beak on the wood. At that point it saw us and flew off, leaving the dead pigeon on the ground. About fifteen minutes later another bird was seen through the kitchen window. This time it was a magpie that had alighted and begun to eat the dead pigeon. The magpie was encourage to leave, and the dead pigeon put in a polythene bag and hidden under a wood display under the overhang. The bag remained there until darkness fell. However it disappeared overnight. Presumably this was a fox or feral cat. The kestrel incident, besides being extremely interesting in terms of an urban siting of a highland bird in the city, shows how top predators such as the kestrel, the magpie, and presumably a fox or cat, compete for meat in an urban environment. Interestingly enough, kestrels are now fairly common in Glasgow, and kestrels and sparrow hawks are known to nest on the University campus (personal communication - James Munro). The second example is interesting because it is an interaction between a mammal and a bird. During October 1998 on the University of Glasgow campus, two magpies were observed attacking two squirrels. The attack developed as follows. The two magpies attacked a single squirrel in a nest on a whitebeam outside the West Medical Building on the main university campus. The nest was about 1 5 to 20 metres above the ground. The attack involved considerable noise, which first drew the attention of the observer to the event. The noise consisted of screeches and flapping wings. A second squirrel then appeared, apparently from the nest, and joined in the fray. The confrontation between the magpies and the squirrels lasted for about 15 minutes. The magpies eventually ceased attacking and flew off The squirrels appeared to go back into the nest. The nest was probably a crows nest. The question is what were the squirrels and magpies fighting over, in other words what was in the nest? It is possible that the magpies were using the nest to rear their young, whereupon the squirrels were presumably attempting to prey on the young of the magpie. The other alternative is that the nest may have been used as a dray by the squirrels, in which case the magpies were attempting to prey on young squirrels, and the two adult squirrels were protecting their offspring. Neither alternative is entirely convincing, as autumn is not a time that is normally associated with the breeding of magpies or squirrels. These two sets of observations on top predators and their activities in a city environment show how important aspects of an animal’s predator/prey status can be easily recorded. They also provide good evidence of the way in which the species interact with each other, and indicate that much research is needed on the role of animals such as these in urban environment. References Elton, C. 1927. Animal Ecology. Sidgwick & Jackson, London. Krebs, C.J. 1972. Ecology. The Experimental Analysis of Distribution and Abundance. Harper Row, New York. Ricklefs, R.E. 1973. .Eco/ogy. Thomas Nelson, London. Wynne-Edwards, V.C. 1962. Animal Dispersion in Relation to Social Behaviour. Oliver & Boyd, Edinburgh and London. Glasgow Naturalist. 2002. Vol 24. Part 1. 98 GREAT SPOTTED WOODPECKERS FEEDING ON THE NECTAR OF RED-HOT POKERS John Mitchell 22 Muirpark Way, Drymen, G63 ODX During the warm and sultry weather experienced in early July 2001, A. & D. MacFadyen drew my attention to a family group of Great Spotted woodpeckers Dendrocopos major repeatedly visiting a tall clump of South African red-hot pokers Kniphofia uvaria in a west Stirlingshire garden. Both adults and a least one juvenile would individually alight on one of the rigid stems of the plant, just below the large flowering head. Binoculuar observations at close quarters showed that the woodpeckers were drinking droplets of nectar from the pendant perianth tubes of the open yellow flowers. Each bird would spend several minutes working its way around a flower head before moving on to the next, occasionally pausing to pick-off and eat an insect which had also been attracted to the feast. Subsequent enquiry produced a similar record of Great Spotted woodpeckers nectar-feeding on Red-hot pokers at Blackball, Edinburgh, in the summer of 1999 (D. R. McKean, pers comm.) Great Spotted woodpeckers drinking the sap oozing out of the bark of trees in spring are well documented in northern Europe (Cramp et al. 1985), but taking advantage of the availability of sugar-rich nectar from a cultivated herbaceous flower in summer is behaviour that appears to have been little observed in the species. References Camp, S. et al. (1985). The Birds of the Western Palearctic Vol. IV: Terns to Woodpeekers. Oxford University Press. Glasgow Naturalist. 2002. Vol 24. Pt 1. 98-101 HAECKELIAN RADIOLARIAN MATERIAL AND THE MICROSCOPICAL SOCIETY OF GLASGOW P. Geoffrey Moore’ and Richard Sutcliffe^ ‘University Marine Biological Station, Millport, Isle of Cumbrae, KA28 OEG ^ Open Museum Nitshill, Woodhead Road, South Nitshill Industrial Estate, Glasgow, G53 7NN It is a pleasure to put on record the recent receipt of a box of historic microscope slides most kindly donated to the University Marine Biological Station 98 Millport (but see below) by Mrs Elizabeth Fletcher, widow of Professor Bill Fletcher (late of the Biology Dept, Strathclyde University). The slides were all prepared by the renowned Professor Ernst Haeckel (1834-1919) of Jena, who is remembered for his 'gastraea- theorie' (1874) in which all metazoan life was traced to a universal gastrula-like ancestor. The earliest substantial treatise on Radiolaria was the comprehensive report by Haeckel (1887), based on samples obtained from the Challenger Expedition (1873-1876). His monograph remained, until very recently, the major source of information on radiolarian diversity and taxonomy (Anderson, 1983). Not much is known of the links between Haeckel and the Marine Station. He was due to have been the dignitary to have officially opened the Scottish Marine Station [the progenitor of the Millport Station, then based at Granton, nr Edinburgh] in April 1884 but he was ill on the day and, due to his indisposition. Dr (later Sir) John Murray stepped in to do the honours for him (Marshall, 1987; Moore, in press). The wooden slide box (Fig. 1), and each of the 33 slides (number 31 in the sequence 1-34 being missing), which Bill Fletcher had in his safe keeping, however, is clearly stamped with the imprint of The Microscopical Society of Glasgow. (Fig. 2) It contains what may be important specimens of Radiolaria, including Challenger material. In 1931, The Microscopical Society of Glasgow was subsumed into the Glasgow and Andersonian Natural History and Microscopical Society - now the Glasgow Natural History Society (Sutcliffe, 2001). It seemed only right and proper therefore to return these slides to Glasgow, having noted the Millport link. They have been added to the Glasgow Museums collection (Registered number Z.2001.2) We append a list of this material so scholars elsewhere may be aware of its existence in Glasgow. The collection consists of stained balsam mounts (although there seems to be much fungal penetration of the preparations) with printed labels to the left side "Radiolarien- Collection von Professor Ernst Haeckel. Jena. 1890." (Fig. 3), and to the right as follows: Rad. Coll. Nr.l. Polycyttarien-plankton (Pelagisch. ) Mediterraneum. Messina. Haeckel. Rad. Coll. Nr.2. Polycyttarien-plankton (Pelagisch.) Nordl. Atlant. Oc. Canaria. Haeckel. Rad. Coll. Nr.3. Polycyttarien-plankton (Pelagisch.) Nordl. Atlant. Oc. Bermudas. Haeckel. Rad. Coll. Nr.4. Polycyttarien-plankton (Pelagisch.) Sudl. Atlant. Oc. Trinidad. Rabbe. Rad. Coll. Nr.5. Polycyttarien-plankton (Pelagisch.) Sudl. Ind. Oc. Madagascar. Rabbe. Rad. Coll. Nr.6. Polycyttarien-plankton (Pelagisch.) Nordl. Ind. Oc. Ceylon. Haeckel. Rad. Coll. Nr.7 .Polycyttarien-plankton (Pelagisch.) Sudl. Pacif Oc. Elisabeth-I. Rabbe. Rad. Coll. Nr.8. Polycyttarien-plankton (Pelagisch.) Nordl. Pacif. Oc. Japan. Chalk 229. Rad. Coll. Nr.9. Acantharien-plankton (Pelagisch.) Sudl. Pacif Oc. Patagon. Chalk 302. Rad. Coll. Nr. 10. Acantharien-plankton (Pelagisch.) Nordl. Atlant. Oc. Far-Oer. "Triton" Rad. Coll. Nr.lk Phaeodarien-plankton (Pelagisch.) Nordl. Atlant. Oc. Far-Oer. "Triton" Rad. Coll. Nr. 12. Phaeodarien-plankton (Pelagisch.) Nordl. Pacif Oc. Sandwich. Chalk256. Rad. Coll. Nr. 13. Phaeodarien-plankton (Pelagisch.) Sudl. Pacif Oc. Galapagos. Rabbe. Rad. Coll. Nr. 14. Tiefsee-Kerat. Spongien-Skek Psammopemma radiolarium. C. Pacif Chalk 272. (Pelagisch.) Rad. Coll. Nr. 15. Tiefsee-Kerat. Spongien-Skek Cerelasmagyrosphaera. C. Pacif. Chalk 271. Rad. Coll. Nr. 16. Tiefsee-Kerat. Spongien-Skek Psammophyllum annectens. N. Pacif. Chalk 244. Rad. Coll. Nr. 17. Tiefsee-Kerat. Spongien-Skek Stannophyllum zonarium. C. Pacif. Chalk 271. Rad. Coll. Nr. 18. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 225. W.Pacif 4475 Fd. Rad. Coll. Nr. 19. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 226. W.Pacif 2300 Fd. Rad. Coll. Nr.20. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 265. C. Pacif 2900 Fd. Rad. Coll. Nr.21. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 266. C. Pacif 2750 Fd. Rad. Coll. Nr.22. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 268. C. Pacif 2900 Fd. Rad. Coll. Nr.23. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 270. C. Pacif 2925 Fd. Rad. Coll. Nr.24. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 271. C. Pacif 2425 Fd. Rad. Coll. Nr.25. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 272. C. Pacif 2600 Fd. Rad. Coll. Nr.26. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 273. C. Pacif 2350 Fd. Rad. Coll. Nr.27. Tiefseeschlamm. Radiol. Ooze. Chalk Stat. 274. C. Pacif 2750 Fd. Rad. Coll. Nr.28. Tiefseeschlamm. Rother Thon. Chalk Stat. 241. N. Pacif 2300 Fd. Rad. Coll. Nr.29. Tiefseeschlamm. Rother Thon. Chalk Stat. 244. N. Pacif 2900 Fd. Rad. Coll. Nr.30. Tiefseeschlamm. Rother Thon. Chalk Stat. 253. N. Pacif 3125 Fd. Rad. Coll. Nr.32. Tiefseeschlamm. Radiol, ooze. Egeria. IV. 1887. Ind. Oc. 271 1 Fd. Rad. Coll. Nr.33. Tiefseeschlamm. Radiol, ooze. Egeria. V. 1887. Ind. Oc. 2779 Fd. Rad. Coll. Nr.34. Fossile Radiolarien Tertiiiar- Mergel von Barbados (Miocaen. Antillen.) The slides were presumably produced as standard sets to be sent around interested scholars of the day. An incomplete set of almost identical slides exists within Glasgow Museums’ collections. Unfortunately numbers 3, 5, 11, 18, 24, 25 and 31 (of this set) are missing. These have no known connections with the Microscopical Society of Glasgow and their origin was not recorded. The only other set of microscope slides believed to exist from the Microscopical Society of Glasgow 99 Figure 1 Wooden slide box containing 33 slides. Figure 2 Interior of slide box, stamped Microscopical Society of Glasgow. Figure 3 Typical example of one of the slides. Figure 4 Printed label Glasgow Microscopical Society. Figure 5 Some examples of slides showing different methods of preparation. 100 are also in the Glasgow Museums collection (Registration number Z. 1 983 . 1 80). These comprise a set of 24 miscellaneous slides, all marked Glasgow Microscopical Society. (Fig. 4) They are also housed in a similar wooden slide box. They were purchased for the Museum from a James Pollok in 1983, and were said at the time to have originally belonged to Mr Pollock’s grandfather. This was probably Charles Frederick Pollock M.D., F.R.S.E. (b. c. 1854), who was one of the original Vice Presidents of the Microscopical Society of Glasgow. (King, 1936). They include such diverse items as the wings of a wasp, pollen grains, chalk and even chemical crystals from claret wine! All the slides are individually labelled with a section and number, and may represent examples of slides which were passed around from member to member. The slides exhibit different methods of preparation (Fig. 5) and obviously came from several different sources, named preparers including F.T. Barrett; Homell Biological Station, Jersey; Norman (J. T. Norman , c.1814 - 1893); and E. Wheeler (London). Only two slides are dated (both 1887), but all the slides appear to be of a similar date, from around the time the Society was formed (in 1886). More examples from this set were illustrated in Sutcliffe (2001). References Anderson, O.R., 1983. Radiolaria. Springer-Verlag, New York, 355pp. Haeckel, E.,1874. The gastraea-theory, the phylogenetic classification of the Animal Kingdom and the homology of the germ-lamellae. Quarterly Journal of the Microscopical Society, 14, 142-165. 223-247. Haeckel, E., 1887. Report on Radiolaria collected by H.M.S.Challenger during the years 1873-1876. In, C. W. Thompson and J. Murray (Eds), The Voyage of HMS. Challenger, H. M. S. 0., London, 18, 1-1760. King, L.A.L., 1936. The Peter Goodfellow Lecture - A short account of the Microscopical Society of Glasgow. Glasgow Naturalist 1 2(2), 46-49. Marshall, S.M., 1987. An account of the Marine Station at Millport. University Marine Biological Station Millport, Occasional Publication No.4, 133pp. Moore, P .G., in press., Capt. Alexander Turbyne and the origins of the Marine Station at Millport. The Linnean Sutcliffe, R., 2001 Glasgow’s Natural History Societies - an update. Glasgow Naturalist 23(6) 62-67. Glasgow Naturalist. 2002. Vol 24. Part 1.101 LILY BEETLE {LILIOCERIS LILII), IN GLASGOW Richard Sutcliffe Open Museum Nitshill, Woodhead Road, South Nitshill Industrial Estate, Glasgow, G53 7NN Two specimens of the scarlet (or red) lily beetle {Lilioceris lilii (scopoli)), were observed mating on royal lilies, Lilium regale in a garden in Sween Avenue, Glasgow (NS5859) on 6th May 2002 by Kenneth Boyle. Voucher specimens were collected on 12th May 2002 and were deposited in Glasgow Museums (Day Book number DB.7540), and the Hunterian Museum, Zoology (Entry No. Zoo/ 18/2002). This non-native beetle, found throughout Eurasia from the Atlantic to the Pacific, the Middle East and North Africa (Halstead, 1989), has become a serious problem in southern England. Both adults and larvae damage lilies {Lilium spp.) and fritillaries (Fritillaria spp.). There were isolated records of the beetle in southern England and Wales in the nineteenth century (Stephens, 1839), but it did not establish itself There were further occurrences in the London area, Flintshire, Carlisle and Cheshire from 1939 (Fox-Wilson, 1943; Southgate, 1959) and it continued to spread in the 1950s. It had become firmly established in Surrey, Berkshire and Hampshire by the late 1980s (Halstead, 1989). By 2001 it could be found locally in much of southern Britain, as far north as Warton, near Camforth, Lancashire. The history of the beetle in Britain is described by Cox (2001). The beetle has become more widespread in England in recent years. This record appears to be the most northerly record of the species so far in Britain, and the first for Scotland. Since the appearance of an article on the lily beetle in the Glasgow Herald (11 June, 2002) two additional reports have been made in the same area of Glasgow, one in Cathcart and another in Netherlee. This could indicate that a local population may already be or might become permanently established. Acknowledgements Thanks to lain Gibson, Kenneth Boyle, and Geoff Hancock for bringing the record to the author’s attention. References Cox, M.L., (2001). The Status of the Lily Beetle Lilioceris lilii (Scopoli, 1763) in Britain (Chrysomelidae: Criocerinae) The Coleopterist 10, 5-20. Fox-Wilson, G., (1943). The lily beetle Crioceris lilii Scopoli: its distribution in Britain (Coleoptera). Proceedings of the Royal Enomological Society of London (A) 18, 85-86. Halstead, A.J., (1989). On the move? The Garden 114, 321-323. Southgate, B.J., (1959). The present status of the lily beetle Lilioceris lilii (Scop.) in Great Britain (Col: ChrysomQlxAdiQ). Entomologist’s Gazette. 10, 139-140. Stephens, J.F., (1839). A Manual of British Coleoptera, or beetles. London, Longman, Orme Brown, Green and Longmans. 101 Glasgow Naturalist. 2002. Vol 24. Pt 1. 102-103 RABBIT CALICIVIRUS DISEASE ON AILSA CRAIG, AYRSHIRE IN 2003 B Zonfrillo (1), H Thomson (2) & R A Stewart (1) 1. Division of Environmental & Evolutionary Biology, Graham Kerr Building, Glasgow University, Glasgow G12 8QQ. 2. Glasgow University Department of Veterinary Pathology, Veterinary School, Bearsden Road, Glasgow, G61 IQH Following the eradication of Brown Rats Rattus norvegicus from Ailsa Craig in 1991, Rabbit Oryctolagus cuniculus numbers were noted to have declined considerably through eating some of the distributed Warfarin bait. However, they survived the baiting operations whereas the rats did not and over the next decade their numbers increased once again to near their former population level of some thousands. Pictures of the changes in vegetation were recorded in the paper on the island’s flora (Zonfrillo, 1994). Until the first months of 2003 rabbits were again a major detrimental influence on the island’s vegetation. In spring 2003 the island’s vegetation was again looking lush and thriving and noticeable was the lack of the usual “carpets” of rabbit droppings on the pathways. Sightings of rabbits were also lower than could have been expected. Something had clearly reduced the numbers of rabbits at least on the lower sections of the island. During summer 2003 increases in the flowering plants around the lighthouse “flat” area of the island suggested that locally, rabbits had gone completely. Several wizen rabbit corpses were strewn around this area and no live rabbits were observed. On the upper slopes the situation was similar with many corpses but still a few live rabbits around. No deliberate attempt had been made to eliminate rabbits from the island and thus whatever was affecting the rabbit numbers had been contracted under natural circumstances. By early July 2003 rabbits were seen in only two areas of the island - under the west cliffs and near the summit on the south side. In the latter site BZ and R S found on 4 July a fresh dead rabbit. The liver and spleen were removed for subsequent examination. HT examined the tissue and noted focal hepatic necrosis in the rabbit with micro thrombi in the hepatic blood vessels. There was also massive necrosis in the spleen. The observed damage was typical of Rabbit Haemorrhagic Disease or as it is more specifically known Rabbit Calicivirus Disease (RCD). This disease is a caliciviral infection and is specific to rabbits only. Examination of the dead rabbit on the island showed no sign of typical Myxomatosis, a disease that occurs on Ailsa Craig about once a decade (BZ pers. obs.) or any other physical injuries. By August 2003 no rabbits could be found in any of the previous two areas where noted and it was clear that most if not all had been eliminated. Rabbits were mentioned as being present on Ailsa Craig as early as 1688, where, as on many islands, they were deliberately introduced to provide food for residents or stranded fishermen in times of bad weather (Lawson, 1888). Detrimental effects of rabbits on the island include soil erosion, vegetation modification and the banishment of some edible plants to the sea cliffs as relict populations e.g. the nationally rare Tree Mallow Lavatera arborea. RABBIT CALICIVIRUS DISEASE. Caliciviral infections such as RCD are a relatively modern phenomenon in wild rabbit populations. Their origin is believed to be from oceanic sources such as shellfish infected by sewage (Smith et al, 1998). RCD was first reported in China in 1984 and soon after in Europe and then Mexico. By 1991 it had reached Australia. Over a few months, it killed around sixty-four million farmed rabbits in Italy alone. The disease has now affected rabbits in over 40 countries and on four continents. Unlike Myxomatosis that in the 1950s, killed 99 per cent of the rabbit population: the kill rate today is often less than 50 per cent. RCD appears to be 100% lethal, at least on constrained isolated island populations and, in captivity, on rabbit farms. In Europe, the disease appears to follow a two-year cycle. Young rabbits up to five and sometimes eight weeks old that contract the virus do not die from the disease, but develop antibodies and become immune. They survive to become the breeding population in the following year. Maternal antibodies to the virus can be passed to young and confer immunity. However, this immunity is temporary, lasting some twelve weeks. The next generation of young rabbits become susceptible and rabbit calicivirus can spread through the population once again (Munro et al. 1994). Comprehensive tests in Australia of RCD showed every indication that rabbit calicivirus is specific only to the European rabbit. Hares Lepus sp. are not affected. Other research elsewhere into RCD tested 43 different animal species for susceptibility to rabbit calicivirus. The virus did not grow in any of them (Munro et al, 1994). Rabbits usually die 24 hours after the disease is contracted. IMPLICATIONS FOR THE SCOTTISH FAUNA How RCD got to Ailsa Craig, the most southerly of Scottish islands, is a matter for speculation. It was certainly not deliberately introduced but may have arrived through fleas on migrant birds or perhaps even on the boots of human visitors. Rabbits contract the disease through sniffing droppings or contact with infected rabbits or from vectors such as fleas or mosquitoes. If this disease spreads throughout the rest of the Scottish rabbit population on islands or mainland it may have serious implications if rabbit numbers are greatly reduced or completely eliminated. In some cases these may be beneficial as on Ailsa Craig with improved vegetation and knock-on effects on lepidoptera and other invertebrates and the species that feed on them. Elsewhere, predators in some 102 areas feeding almost exclusively on rabbit e.g. Golden Eagles Aquila chrysaetos. Buzzards Buteo buteo, Great Black-backed Gull Lams marinus (on Ailsa Craig), Red Fox Vulpes vulpes and Stoat Mustela erminea to name but a few, may have difficulty surviving. Ecological impact of RCD on native fauna may result in predation on alternative, supplementary or opportunistic prey. If abundance of prey species changes, then it will be recorded in a predator's diet. 'Prey switching' only occurs if the relative availability of a certain prey species is not reflected in the composition of a predator's diet (Murdoch and Oaten. 1975). Predatory species currently locked in to a mainly rabbit diet will therefore “switch” if rabbit numbers rapidly decline. Three main responses may arise from a predator that a relies upon a prey species such as rabbit for a main component of its diet throughout the year: a. Behavioural changes, where habitat use and size of foraging range widens, thus increasing effort. b. Dietary response, when the loss of main prey is balanced by taking alternative species; c. Reduction in numbers, when food availability reduces predator populations through starvation that in turn promotes their wider dispersal, increases mortality and results in overall poorer breeding success. At this early stage the future effects of the calicivirus on rabbit numbers in Scotland should be monitored, as should changes in predator numbers or behaviour patterns. Native fauna and flora may benefit or be at risk from a rapid decline in rabbit numbers. Time will tell if long-term immunity to RCD does not emerge and species recovery therefore becomes unlikely. Consequences for some species of the present Scottish fauna may be profound. REFERENCES Lawson R (1888). Ailsa Craig : Its History and Natural History. Paisley. Munro R K and Williams R T Eds. (1994). Rabbit Haemorrhagic Disease: Issues in Assessment for Biological Control, Bureau of Resources Sciences, Australia. Murdoch W and Oaten A (1975). 'Predation and population control.' Jowr/ju/ of Applied Ecology, 12: 795- 807. Smith AW, Skilling D E., Cherry N, Mead J H and Matson D O (1998). Calicivirus Emergence from Ocean Reservoirs: Zoonotic and Interspecies Movements. Emerging Infectious Diseases. 4.(1) 13 -20. Zonfrillo B (1994). The Flora of Ailsa Craig. Glasgow Naturalist. 22 : 4. 307 - 344. 103 -■ -• • : :^rL:M ih -■‘A‘ :«C'4 -.% i|f v^ * ■■■^* t « i .*W' m ^ -A * "'"{JI " V ' ^ '-'’^Ji . ‘ 'T* 104 Glasgow Naturalist. 2002. Volume 24. Part 1. Pages 105-117. BOOK REVIEWS Compiled by Ruth Dobson A GUIDE TO BIRD WATCHING IN THE CLYDE AREA Edited by Cliff Baister and Marion Osier Colour photographs on covers by Phil Newman, black and white artwork byThelma Sykes. Sturdy 'ring-bound' volume, 145 pp £10.00 Scottish Ornithologists Club, Clyde Branch 148pp. drawings and sketch maps, 2001. No ISBN £10.00. Available from Cliff Baister, 20 Chapelton Avenue, Bearsden, Glasgow, G61 2DQ. £11.50 (inch p&p) also from some wildlife outlets in the area. If you want to watch birds in the Clyde Area this is the book for you! Cliff Baister has drawn on the SOC Clyde Branch for extensive ornithological information on the Area which covers:- City of Glasgow, Loch Lomond, Lanarkshire, Renfrewshire, Dunbartonshire and West Stirlingshire. After a general discussion of the area covered, the book deals with the individual sites. Each site has a clear map showing where to park, the best route to take and the best viewing locations. The text relating to the site describes the locale and explains how to get there by car or by public transport. Once there, you are advised on the best paths to follow and if hides have wheelchair access and if there are closing times on the reserves. There follows 2 sections on what birds you may see in Spring/Summer and in Autumn/ Winter. There are about 80 sites referred to although not all have the full description. The book acknowledges sponsorship of RSPB, SNH, and The Glasgow Natural History Society. Ian C McCallum AMPHIBIANS AND REPTILES: A NATURAL HISTORY OF THE BRITISH HERPETOFAUNA Trevor J.C. Beebee & Richard A. Griffiths Harper Collins, London, 2000, 270pp softback with over 70 black and white illustrations. ISBN 0 00 220084 8, £19.99. This new volume follows Malcolm Smith’s The British Amphibians and Reptiles (1951, New Naturalist number 20) and Deryk Frazer’s Reptiles and Amphibians in Britain (1983, New Naturalist number 69). Do we need a new book for such a small part of the British fauna - only 12 or 13 terrestrial species, one or two marine visitors and an assortment of semi-established introductions? Beebee and Griffiths justify this book by the wealth of new information on the British herpetofauna since Frazer’s work (their 300 or so references are predominantly post- 1980) and by a change of emphasis; they concentrate more on ecology, life histories and conservation than the previous authors. Beebee and Griffiths are well qualified for the task: they are authors of a large number of the cited papers; as they acknowledge, they are both essentially amphibian specialists, but they have consulted widely on their treatment of the reptiles. This is a very authoritative book and very wide ranging. Though it concentrates on ecology and conservation, the treatment of physiology is up to date and refreshing. I liked particularly their account of ectothermy and thermoregulation: they note that the term ‘cold-blooded’ is highly misleading and that ectothermic animals may have considerable advantages over the so-called ‘advanced’ endotherms. The basic structure of the book is similar to Frazer’s. There is a core of five chapters devoted to the main groups in turn: newts, frogs and toads, lizards, snakes and chelonians. Beebee and Griffiths have five additional chapters (compared to Frazer’s two) covering amphibians and reptiles in Britain, historical aspects of herpetology (including folklore), basic biology of amphibians and reptiles, aliens and conservation. The conservation chapter, in particular, is a considerable expansion on the earlier treatment, representing the greatly increased importance of conservation in recent times. From a Scottish point of view, the distribution maps, from the Biological Records Centre, make interesting viewing. Though the book emphasises the modem threats facing reptiles and particularly amphibians, a comparison of Frazer’s maps and the new ones suggests most species are commoner than they were 20 years ago. Especially in southern Scotland, most species, particularly palmate newts, common frogs and toads, adders, slow-worms and common lizards are recorded from more squares than in Frazer’s maps. Of course, records reflect recorders rather than species abundance, but the increased number of records in southern Scotland is very striking. I only hope that they are correct. Interestingly, Bowles (2001) does report increased numbers and more widespread distribution at least in the case of the common lizard (Lacerta vivipara) from his own personal experience. All in all, this is an excellent addition to the New Naturalist series, not as a replacement for the previous books, but as an update and extension of our knowledge. It is good value for money (only £9 more than Frazer - though softback, rather than hardback). My only quibble is that, unlike Frazer, Ireland has been omitted from the distribution maps. The last time I heard, Ireland was still part of the British Isles...? Reference Bowles, F.D (2001) Elusive lizards? Scottish Wildlife 44, 16-19. J.R. Downie 105 MINDING ANIMALS: AWARENESS, EMOTIONS AND HEART Marc Bekoff Oxford University Press, Oxford 2002, 230 pp., hardback with a few black & white illustrations. ISBN 0-19-515077-5. £18.99 Marc Bekoff is a distinguished ethologist (that is a biologist who studies animal behaviour in the spirit of Niko Tinbergen and Konrad Lorenz) who has spent many years investigating the development of individual differences in behaviour. He has concentrated on the role of play in this process, often working with social carnivores such as wolves and coyotes. Like many people studying animal behaviour (and especially the fascinating and enigmatic topic of animal play), he has beome increasingly interested in and impressed by the mental capacities of his subjects. Minding Animals is a synthesis of his views on this topic. The book has two broad, loosely related themes. Firstly, it discusses the nature of the mental processes of animals and, arising from this, the proper way to do science. Secondly, it considers the many ways in which humans abuse animals (including destroying their natural habitats) and what could be done about this. Frankly, I found those parts of the book relating to the first of these themes almost impossibly irritating to read and this has coloured my view of the whole book. To give readers of the Glasgow Naturalist a chance to judge for themselves whether to read this book, in this review 1 have concentrated on summarising the material covered as far as possible without comment, just pointing out briefly at the end what I found so annoying about it. Minding Animals starts with a glowing forward by Jane Goodall (whose famous work on the social behaviour of chimpanzees has been hugely influential). In his own preface, Bekoff explains how he came to be interested in ethology, animal cognition and welfare and previews the material to be covered. Chapter 1 {Chasing coyotes and moving yellow snow) gives a series of examples of the complexity of the behaviour of animals. Bekoff argues from these that we cannot make a clear distinction between human and non-human animals in terms of their cognitive processes and capacity for emotional experience. He cites statistics on the use and abuse of animals (in the food industry and in scientific research), calls for science to be more socially responsible and gives a brief historical account of the ethological approach to behaviour. He stresses the importance of careful comparative studies of the behaviour of wild animals (as practiced by the joint Nobel prize winners Lorenz, Tinbergen and Von Frisch) when compared to more intrusive laboratory-based approaches to the subject. Chapter 2 {Representing and misrepresenting animals) discusses human attitudes towards animals, how animals are represented in television programmes and advertisements (for example) and how this influences the way we treat them. In this chapter Bekoff argues that anecdotes about one-off incidents and anthropomorphism (ascribing human motives, feelings and emotions to animals) have a useful role in science. Chapter 3 {The richness of behavioural diversity, rightly subtitled “a potpourri’') gives another series of case studies of more-or-less complex behaviour patterns in animals (from mate choice in insects to self- medication in chimpanzees). Current understanding of the mental processes of animals is covered in chapter 4 {Animal minds and what's in them), concentrating on cognitive ethology (“the comparative evolutionary and ecological study of animal minds and mental experiences”). In chapter 5 {Animal emotions) Bekoff briefly discusses the nature of emotions and gives more case studies of events that he considers as indicative of strong emotions in animals (not just more simple ones such as fear and anger, but also complex emotions such as joy and embarrassment). He counters the arguments of those who have reservations about ascribing to animals emotions similar to those that humans experience. Chapter 6 {Play, cooperation and the evolution of social morality) draws on Bekoff s extensive experience of studying social play in young carnivores and develops his theory that during play young animals learn codes of social conduct that influence their subsequent interactions with other animals (or lay the “foundations of fairness”). Chapters 7 and 8 {Animal welfare, animal rights and animal protection and Human intrusion into animals ’ lives) discuss some of the various ways we use animals (from hunting them, to keeping them in zoos and dissecting them in practical classes), the rights that animals have and people’s attitudes to exploiting animals. Special attention is given to the use of animals in scientific research. After considering why we need to study animals, Bekoff explains how scientific techniques (even relatively non-invasive ones such as those used on wild animals by fieldworkers) can alter the behaviour of the animals concerned and so compromise any results. Chapter 9 {Science, nature and heart) looks in a more general way at how people influence animals (and consequently nature as a whole). It covers extinctions, trade in endangered species, destruction of natural habitat (I did not know that mining for coltan, which is used to make mobile phones, has decimated populations of gorillas and elephants) and abortive reintroduction programmes. The final chapter {Animals and theology) is designed to pull everything together; the subtitle “Stepping lightly with grace, humility, respect, compassion and love” sums up the points Bekoff is making here about how we should treat our planet and the animals that inhabit it with us. There are a number of good things about Minding Animals. It is well written, in a highly personal style with very little jargon, which potentially makes it pleasant to read. Bekoff also avoids the assumption that all scientists are male. He takes a 106 clear look at the practice of science in general and the study of behaviour in particular, honestly and critically identifying ethical and scientific problems raised by work in this area, including his own. He calls for us to make science fun and stresses the importance of educating young people to care for animals and the environment (and to his great credit has been actively involved in this, putting his time and energy where his mouth is). In this context, he eloquently acknowledges and appreciates the fact that today’s young people seem to be better at this than many of their parents' generation. Finally, he displays a passionate love for animals and concern for their welfare and for the damage that our species is doing to their habitats. This is all entirely admirable, so what did I dislike so much about it? First of all, I found it very poorly structured. The text is broken up into short sections with their own titles, some of which work well (“Elephants aren’t couches”) and others (in my view) do not (“Patient and compassionate altruism; indifference is deadly”). These sections are often repetitive and poorly organised, in the sense that Bekoff keeps coming back to the same points and examples and jumps between different points so that the flow of his argument is obscured. In addition, while purporting to be arguing logically on the basis of facts, at least when supporting the points he wishes to make, Bekoff can be careless about what the evidence does and does not show, and often does not give the reader sufficient information about specific examples to judge for his/herself To give just one of many examples, in chapter 3 we are told that subordinate monkeys who can perform a learned task effectively when on their own, do so less well in the presence of a dominant companion. This is interpreted as showing that they are deliberately “playing dumb” to avoid trouble, whereas they could simply be too stressed to function well. This misuse of evidence (as I see it) would not matter if the book were simply a personal statement of belief in and concern for the rights of animals, but it does matter since the book is also supposed to be about science. Of course we should recognise the complexity of animal behaviour and be prepared to study and understand it, but over-interpreting data does not help. Another thing that puts me off this book is the way that Bekoff treats those who disagree with him, for example on the subject of anthropomorphism and the mental states of animals giving them derogatory names; thus people who are not convinced by the value of cognitive ethology are called “slayers”. He sets up straw men, quoting out of date positions and implying that views are more narrow and rigid than they actually are. Overall, he comes across as somewhat self-congratulatory; for example, many people study chemical communication, so just what is so outrageously special about his study that involves “moving yellow snow” to see how coyotes use urine marks? He is also patronising; for example, the title Minding Animals is quite (in the British sense of “fairly” rather than the North American sense of “very”) a clever play on words, but we really do not need to have it explained to us once, let alone twice. Setting aside my churlish view, in Minding Animals Marc Bekoff touches on many very interesting and important issues and the book is informative and challenging to read, whatever one’s views on anthropomorphism. At any rate, the cover picture is beautiful. This shows a mother polar bear and her two cubs, snuggled in together with beatific expressions on their faces. For all that I have said, one can see where he is coming from. Felicity Huntingford BIRD MIGRATION: A GENERAL SURVEY Peter Berthold Oxford University Press, Oxford 2001, 253 pp with many figures. ISBN 0 19 850786 0 (Hbk), 0 19 850787 9 (Pbk) £27.50. This is the second edition of Peter Berthold's review of bird migration studies, published eight years after the first. The literature on migration is formidable, and it is a brave author who attempts to provide an up-to-date review of recent research. This is an excellent attempt at that task, and like the earlier edition has major sections on the methods used to study behaviour, the various forms that migratory journeys take, the physiological control of migration, and orientation mechanisms. There is a shorter section on the problems of conserving migratory species and a neat chapter that talks the reader through the various remarkable events that an individual warbler will go through during its annual migratory journeys - this brief review of the whole subject I found the most rewarding in the book. There is still a great deal that we do not understand about migration, especially how some bird species are able to navigate so accurately. We really have only the most rudimentary knowledge of how birds might be able to achieve this. The whole book is an excellent review of recent scientific research. My only criticism is that it is rather dull. There is a real sense of wonder and mystery in bird migration. Reg Moreau once calculated that each autumn 5000 million individual small passerines cross the Mediterranean from Europe to overwinter in Africa. The thrill we experience when we see our first swallow of the summer is largely in recognition of the truly awesome journey that this small bird has made from South Africa home to Scotland. It is a pity that scientific writing has so often had the sense of wonder squeezed out of it. David C. Houston 107 COLLINS FIELD GUIDE: CATERPILLARS OF BRITAIN AND EUROPE David J. Carter, illustrated by Brian Hargreaves HarperCollins, London, 2001, 296 pp., hardback with numerous colour plates and monochrome end plates. ISBN 0 00 219080 X, 16.99. Reading the publishers' advertisement one might think that this is a new book which was published during June 2001. However examination of the volume itself soon dispels this notion beeause it is elearly dated 1994 and is simply a eorrected version of the original A Field Guide to Caterpillars of Butterflies & Moths in Britain & Europe which first appeared in 1986 and which was favorably reviewed by the present writer in Glasgow Naturalist 21, 1986 p. 227. There is a differently styled eover and slightly modified title but the text appears identieal to that of the original publication except for slight ehanges in the bibliography and changes in many latin names to accord with modem usage. These changes in names ean be readily detected in the main text beeause amended entries are printed in a slightly less bold type than the originals. Revised names in the eaptions to the plates also appear in a different type faee but there are no differenees in the fonts of the food-plant lists or in the general index. There are some mistakes in the application of these correetions e.g. the Emperor Moth, fomerly Saturnia pavonia is corrected to Pavonia pavonia in the text but appears as Saturnia in the index and in parts of the food-plant lists; similarly the old name Clossiana appears in the index instead of Boloria as in the text. These criticisms apart, the book still remains the most aeeessible and useful book of its kind and, paeked full of information on the life-histories, distribution, food-plants and diagnostic characteristics of the species considered and with its superb eolour illustrations, is highly reeommended to all with an interest in eaterpillars. Ronald M. Dobson GLOBAL WARMING. THE SCIENCE OF CLIMATE CHANGE Frances Drake Edward Arnold, London. 2000. 273 pp. Paperback. £14.99 ISBN 0 340 65302 7 We wish we had been able to write this book ourselves. It is coneise, simple and beautifully written. In an elegant introduetion, Frances Drake reminds us that global warming may be the largest experiment ever undertaken by humankind. His style is such that the book can be read by anyone who has an interest in global wanning and yet it has infomiation in it that is new to the speeialist. There are eight ehapters, starting with basic physical concepts and ending with the need for consensus - surely now even more relevant after the terrible destmction of the World Trade Centre in New York on 1 1th September 2001. In chapter one, as elsewhere, the diagrams are elear and extremely good. Franees Drake quickly and expertly describes the climate system, before guiding us through the concepts of energy, radiation, and the Earth's radiation and energy budget. All of these are central to an understanding of the background to global warming. We also diseovered for the first time that there are three types of scattering of light - Raleigh, Mie and Tyndall. We should have known this, since we have leetured on related subjeets. The climate system, which is covered in chapter two, is central to an understanding of global warming. Frances Drake describes the general circulation of air and wind in the atmosphere, the hydrosphere - the seas and fresh waters of the world, the signifieance of gases, oceanie eireulation, winds and surface currents, and the deep ocean eireulation. This latter is not well known to the non-speeialist and is still not fully understood. The seeond part of the chapter eovers subjects which are eentral to short term ehanges in weather as well as to longer time-scale global warming effects. These are oeean atmosphere interaetions, the El Nino Southern Oscillation (ENSO) and its opposite La Nina in the Paeific Ocean, and the North Atlantie Oseillation (NAO) in the Atlantic Ocean. The last part of the ehapter deals with the importance of ice snow on land and at sea, and then the biosphere. Being biologists we would like to have seen more here, but one eannot have everything. Then follows an introduetion to the central theme of the book, equilibria, feedbacks and global warming. They are clearly described and set the seene for the remainder of the book. One of the fascinating aspects of global warming and for that matter cooling, together with associated changes in climate, is that they are happening the whole time. Furthermore they have happened since the earth was first formed. Few people appreciate this or its significance. In other words, ehange is with us the whole time, and it is ourselves, humankind, who have been laeking in understanding. The next two chapters, chapters three and four, deal with this subjeet and make eompelling reading. Chapter three is entitled 'Past Greenhouses and Icehouses' and ehapter four 'Historical Climate Change'. After a quick tour through the major planets in the solar system, the author introduees us to James Loveloek's Goldiloeks planet, the Gaia hypothesis and Daisyworld. Daiseworld is a planet eolonised only by blaek and white daisies. Changes in their abundance dictate the planet's temperature - a wonderful example of biological feedback. However the idea has critics. For a detailed analysis read Drake! We then read about the geologieal elimate reeord and palaeotemperatures, finishing chapter three with the quaternary glacial ice ages and their explanation in astronomical terms - 108 specifically the Milankovitch Cycle. This is gripping stuff, and makes one understand how our early forebears during the ice ages were governed by extraterrestrial events! The historical record that Drake considers in chapter four is factually robust. There are many quantifiable changes. Most people know that the River Thames regularly froze in the eighteenth century and that Dutch paintings of that era show heavily snowed scenes that were obviously regular events. This was the little ice age between 1550 and 1800 during which 'Frost Fairs' on rivers were frequent winter events. We are then brought up to the present time with an account of changing climate in the nineteenth and twentieth century. The author explains clearly the external (extraterrestrial) and internal forcings that change climate, including minima and maxima of solar activity. The 1 1-year solar cycle is fairly well known, but it is much too short to account for changes in climate over a century or more. There appear to be 180, and possibly 400 and 800 year cycles that may fulfill the role. Atmospheric turbidity is also important. This is the amount of dust in the atmosphere, and amongst other things is modulated by large volcanic events such as Krakatoa in 1883. Overcast skies and reduced atmospheric temperatures of a few tenths of a degree follow such events. The last part of chapter 4 is concerned with climate and human history. We thought that this might overlap with what had gone before, but not a bit of it. Here are some fascinating historically accurate case studies including that of the Norse Greenlanders and the western and eastern settlements of the Vikings in Greenland. The settlements were founded in 982 and 985 respectively, when the climate was significantly warmer than now, by up to four degrees. The subsequent history and disappearance of these small colonies is extremely interesting. Present day observational evidence for global warming is widespread, and as a result there are many models attempting to predict future trends. There is also no doubt about the occurence of global warming since the middle of the nineteenth century. These subjects occupy chapters five and six. The evidence centres on what have now become known as the greenhouse gases - that blanket the earth to such an extent that less heat than normal is irradiated out from the earth. They include carbon dioxide which is natural and of considerable importance. Others include methane, nitrous oxide and the halocarbons. The real question is how much of these effects are anthropogenic, that is are caused by man. A significant part of them are. One of the continuing disagreements between field and experimental scientists on the one hand, and theoretical scientists on the other, is the usefulness of mathematical models in helping one to understand environmental phenomena. Our own views are somewhere in the middle. Mathematical models can be of great help if handled sensibly. Parameters need to be carefully constrained, and initial conditions defined clearly. Chapter six reviews the model evidence for global warming. Atmospheric general circulation models have received considerable attention, and have merit. Other modeling methods are also covered, such as time-dependent climate modeling. Surprisingly, the chapter is fairly straightforward. This is a tribute to the ability of the author, whose summary statement on page 198 expresses our own view. “Depending on you stance on global warming climate models either provide the best picture of climate change yet available or are a mirage of scientific fact, providing only the illusion of certainty rather than anything concrete”. Indirect impacts of climate change are perhaps the most difficult to assess objectively, and may be the most important in the long term. Drake considers this rather difficult area in chapter eight, and draws attention to indirect impacts on human populations and on the biosphere. These latter include socio- economic effects, agriculture and health. As the author points out, even our own western technologically advanced societies in the west are not immune from the problems. He draws attention to the need for cost benefit analysis, integrated assessment models, and mitigation options. This is all rather heady stuff, but as usual Drake makes it easy to understand for the intelligent layman. He finishes the book with a chapter entitled ‘The Need for Consensus’. What better way to end - particularly after nine eleven, and the need for cross-cultural dialogue in science the arts and even religion between the nations of the world. A superb book indeed. Peter Meadows and Azra Meadows THE NEW ENCYCLOPEDIA OF REPTILES AND AMPHIBIANS Tim Halliday and Kraig Adler (editors) Oxford University Press, Oxford (2002). 240pp. Hardback. Extensively illustrated with colour photographs, maps and line drawings. ISBN 019 852 507 9. £25.00 This new encyclopaedia is an update of a previous volume (Halliday & Adler, 1986), which is never referred to, but which partly explains the credits to some contributors who have been dead for some time, notably Angus Bellairs. In addition to Tim Halliday and Kraig Adler, both internationally recognised herpetologists, 36 other herpetologists have contributed to the writing, and an army of artists, photographers and editors to the illustrations and lay-out. This is a beautiful and impressive volume. In recent years, biologists have become alarmed at the catastrophic declines in numbers of both amphibians and reptiles. As might be expected of a book edited by Tim Halliday (he is international director of the Declining Amphibian Populations 109 Task Force), conservation issues are given full prominence here. At the same time, there has been an explosion in our knowledge of these animals and even an explosion in the number of recorded species. Until recently, it was generally considered that biologists had recorded most of the more conspicuous species in the world, especially the vertebrates, and that the gaps were mainly in the invertebrates, especially insects. However, as Hanken (1999) reported one of the fastest growing species lists is for the amphibians. The old encyclopaedia (1986) reported a total of 4015 species; this new version gives 5,400, an increase of 34% in 16 years; and since this volume went to press Meegaskumbura et al (2002) reported over 100 new species of frogs from Sri Lanka alone. The increase in known species of reptiles is a smaller percentage (19%) to 7776, though still considerable. As the editors point out, it is something of a historical anomaly that reptiles and amphibians are so frequently considered together : in many ways, they are zoological opposites, especially when we compare the water relations of frogs and lizards. However, the comparisons can be very instructive in showing how diverse the evolutionary adaptations can be to mainly terrestrial life of mainly small, low-energy-consuming, mainly predatory vertebrates. As you might expect of an encyclopaedia, the main structure of the book is an account of the diversity of the two groups, concentrating heavily on living representatives (fossils receive very brief accounts only). Each order (for amphibians : caecilians, urodeles and anurans) receives a full account of general characteristics and biology, then a biodiversity section to family level. Although many individual species are referred to and illustrated, the book (wisely, I think) does not attempt a species or even generic list. There is a little inconsistency in the coverage given to different orders : caecilians, for example, are not given distribution maps for different families, as most other orders are, but this is understandable, given our relative ignorance of this group. In addition to biodiversity, there are numerous "boxes" and double-page spreads on topics of interest such as turtle nesting, pollution effects, snake venom, temperature dependent sex determination and much more. The book concludes with a glossary of terms, a bibliography and an index. Many people might regard this as mainly a "coffee- table" book, because of its wealth of beautiful illustrations. But I hope more will read the text, both for information and to raise awareness of the interest and diversity of these two fascinating animal groups and their increasingly worrying plight. References Halliday, T.R. & Adler, K. (1986) The encyclopaedia of reptiles and amphibians. Allen and Unwin, London. Hanken, J. (1999) Why are there so many new amphibian species when amphibians are declining? Trends in Ecology and Evolution 14, 7-8. Meegaskumbura, M. et al (2002) Sri Lanka : an amphibian hot spot. Science 298, 379. J R Downie WETLAND ECOLOGY - PRINCIPLES AND CONSERVATION Paul Keddy Cambridge University Press, Cambridge, 2000. (Cambridge Studies in Ecology) pp. 614. ISBN (hbk) 0 521 78001 2 £90.00 (pbk) 0 521 78367 4 £32.95. Surely everyone with even a passing interest in output of the BBC Natural History Unit knows that a wetland is that habitat that is formed where the land and the water meet. The very essence of wetlands is that they are the gradation zone from land to water the type of land and water that merge to fonu the wetland to a very large extent determine its nature. Thus wetlands mean very different things in different parts of the world. The similarities between a Lewisian peat bog and an Indonesian mangrove for example seem superficially tenuous but there is no doubt that wetlands are common, highly productive and thus extremely important habitats. Interestingly according to one figure presented in this book, the whole of Scotland, (with the exception of Aberdeenshire) should be regarded as a wetland. The author (based in Louisiana) has clearly visited Scotland during the rainy season! One of the principal aims to this volume is to provide an adequate, all encompassing definition and a minimalist classification system for wetlands. In this aim the author devotes over 30 pages of closely typed script but eventually appears to admit to only limited success! In his second broad aim, to demonstrate the communality of underlying ecological mechanisms between the enormous range of wetland types the author has more success. Drawing on a very wide and varied literature, the author gives a community ecologists view of how wetland systems function. General principals such as zonation, succession, diversity, disturbance and competition are covered logically, clearly and in very considerable depth. Although drawing heavily on wetland examples, these chapters are of equal interest to those interested in other habitat types. In the final three chapters the author turns to wetland conservation and restoration. As with other topics this is dealt with in depth. On the way to a general conservation and management discussion, the author elucidates functional response relationships and functional classification systems of organisms amongst other relatively complex ecological theories. 110 Extensively referenced throughout, this work is not for those with a casual interest in the ecology of wetlands, rather it is aimed at the professional researcher and post-graduate student with a good basic grounding in ecological theory. As such it makes a useful addition to the Cambridge Studies in Ecology Series. Colin Adams FOSSILS AND EVOLUTION T.S. Kemp Oxford University Press, Oxford, 1999, 284 pp. numerous drawings and diagrams. ISBN 0 19 850345 8 (Hardback); 0 19 850424 1 (Paperback). Price £14.99 “What are fossils good for?” This question is posed in the opening line of the book. Are they just “a few of the hard bits” of past life that, while of interest as curios, can tell us little, or are they able to make an important contribution to evolutionary theory. This publication sets out to show that evidence from fossils together with knowledge gained from living organisms can be used in partnership. Fossils are important in these studies and without them our knowledge of would be much poorer. Indeed they are our only evidence for the existence of various extinct groups of organisms such as trilobites or dinosaurs. Fossils illustrate the great diversity of past life and the complex pattern of change through geological history that has lead to the life of today. The subject is set out in a logical and easily followed manner. The book is divided into two parts. The first deals with principles. Following a scene-setting discussion of fundamental ideas Kemp gives a succinct run through of evolutionary theory and of taxonomy. A chapter outlining incompleteness in the fossil record and how to allow for it rounds off this section The second part deals with practices. Five chapters concentrate on what the author considers as central areas of current palaeobiological research and these are tackled using the principles outlined earlier. The areas chosen are fossils and phylogeny; speciation; rules and laws of taxonomic turnover; mass extinctions: and the origin of new higher taxa. This is a textbook aimed at students in courses on evolution and palaeontology. While, as the back cover states, it will be of interest to amateurs, it is not a light read. The book provides an excellent synthesis of the current state of evolutionary theory and taxonomy. It brings out the importance of fossils and succeeds in answering the opening question posed by the author. Alastair Gunning CORAL REEF FISHES OF THE INDO- PACIFIC AND CARIBBEAN Ewald Lieske and Robert Myers Revised Edition, 2001 Collins Pocket Guide, Harper Collins London. ISBN 0 00 71 1 1 1 1 8. 400 pp. over 2,500 coloured illustrations sbk. This volume is a corrected and slightly expanded version of a first edition, published in 1994. In the mean time the original Lieske and Myers has become the standard handbook for the globe- trotting marine biologist or SCUBA diver interested in fish and fish-watching on coral reefs. It follows the now almost standard arrangement for field guides of having sets of full colour illustrations of typically 10-14 related species on the right hand side of each double page spread, and short accounts of each of these species on the left hand-side. The pictures are by and large of very good quality, conveying for most species its typical poise, as well as its colour in life; and the matching text is informative, giving for most species, typical habitat and feeding behaviour, as well as zoogeographical range. Amazingly it covers almost all the species (>2,100) one is likely to see on coral reefs anywhere, in the Indo-Pacific, and also the Caribbean, and given the books pocket guide size and weight, this makes it an indispensable travelling companion for the natural history minded SCUBA diver, or the SCUBA diving biologist. All told, Lieske and Myers’ volume, as it now is, represents a significant achievement, both by the authors, and by the ichthyologists who have informed it. This work corrects scattered errors in the first edition, the most serious of which was the transposition of two of the plates of coloured illustrations. I have yet to find any inaccuracies in this latest edition, and given the evident advisory input of Jack Randall, of the Bishop Museum in Hawaii, with his encyclopaedic knowledge of reef fish taxonomy, I doubt there are many. The advance which this represents is hard for non- specialists to appreciate. The previous and first attempt at a field guide to all coral reef fishes, also published by Collins, was that by Carcasson, available during the 1980s. This was a useful but by no means infallible guide, and was quietly allowed to go extinct. Even this work was an enormous advance on the state of knowledge 30 years before. The problems then of separating species, and determining if they were new to science, or matched the limited descriptions of workers from a century before, is best illustrated by the pitfalls encountered by J.B.L. Smith, the eminent South African ichthyologist, and discoverer of the Coelacanth, whose work on Indian Ocean reef fishes laid the foundations for modern knowledge. Criticism of some of his identifications was reputedly responsible for his suicide. But that some of his determinations turned out to be wrong is scarcely surprising. Among reef fishes, not only may male, female and juvenile have 111 different coloration, but since sex change is common, all sorts of intermediate forms may be found. Further, while species may vary in exact colouration between regions, to the extent that they have often been described as different species, closely related but distinct species may be less obviously distinguishable. An added problem with marine fishes is that in families such as parrotfishes the colours fade almost immediately on collection, with the result that the original descriptions of many species actually bare little resemblance to their appearance in life. Only the development of underwater photography to an everyday tool has enabled appearance in life of such species to be illustrated with confidence. A limited number of corrections apart, this revised edition of Lieske and Myers differs from its predecessor mainly in including an additional 44 species. The tactic that has been used is to add one or more new species to the bottom of those left- hand pages where space in the first edition allowed, and to re -jig the corresponding page of illustrations to squeeze in the extra fish. While doubtless this means that some previously omitted, but not unusual, species have now been included, these extra species have only appeared within those relatively few families for which space was available. A few other small criticisms come to mind. The handy maps of the Indo-Pacific and the Atlantic on the inside covers of the first edition, invaluable in interpreting the text descriptions of zoogeographic range (is Rarotonga east or west of Vanuatu?) are missing in the revised edition. And the diagram of reef morphology that appears in Black and White in the first edition, but in colour in the revised one, has so obviously been crayoned in as to make the earlier version look the more professional. All told there are so few differences between the two volumes that 1 myself have not rushed to acquire the new one. But 1 would recommend the book as now revised whole- heartedly to anyone with a more than passing interest in reef fish. The one proviso is that most fish inclined divers would be well advised to also carry a more specific guide to the reef fishes of the area that they are visiting at any one time. In Lieske and Myers, as in with many comprehensive guides to the species of whole continents, let alone most of the globe, the reader is often faced with plates of numerous veiy similar species, that make it a daunting task to track down the one at hand. In practice few of these species will co-occur in any one area, and many non-experts will find it more useful to have to hand a guide that focuses on the species of their chosen destination. Rupert Ormond MOTHS Michael E. N. Majerus The New Naturalist No. 90, Harper Collins, 2002. 16 colour plates, over 180 black and white photographs or diagrams, 310 pp. ISBN 0 00 220141 0, £34.99. Each volume of the New Naturalist is awaited eagerly by people of all kinds of natural history persuasion in addition to a strong band of book collectors. It would be natural to compare this new volume on moths with the earlier book of the same title by E.B. Ford (1955). Ford (1946) also wrote the very first volume of the New Naturalist series on butterflies and many of a now ageing generation of naturalists claim to have been influenced in their choice of study or pastime by reading it and subsequent titles in this seminal series. Hopefully this book might have a comparable effect on new students. There is also a widely praised book on moths of more recent origin (Young, 1997) with which to compare this new offering. Majerus acknowledges Ford as someone who influenced his own career but has not simply updated the earlier approach to the subject animals. Ford did not flinch from introducing quite technical aspects of biology. In his case it was in the burgeoning area of fundamental genetics. For the new century there is more emphasis on ecological genetics and population biology. Thus Majerus pitches in to a discussion on the Hardy- Weinberg law and a number of other aspects used in university-based studies but perhaps not generally employed by field naturalists. In not succumbing to a ‘dumbing-down’ of these subjects the inclusion of a fairly comprehensive glossary is welcome if not essential. The context of all these is firmly placed in studying the whole organism. Thus the practises of running light traps and wielding nets plus captive breeding are still tools for developing greater understanding of basic biological systems. The considerable effort required for this is necessary for knowledge to advance. This is an idea which needs to be re-inforced in many younger people who believe that gaining access to the internet is a primary source of information. Chapters include folklore, life history and anatomy, evolution, sex lives, host plants, flight dispersal and distribution, death and defence. Of particular interest is a chapter on melanism. It provides some facts to counter recent claims that the classic case of industrial melanism in the peppered moth, used as a textbook example of evolution in action, is based on invalid (or even falsified) experimental methods. The publicity surrounding this, approaching the level of drama in some quarters, has been seized upon by creationists as “evidence” to disprove Darwinian evolutionary processes. Such events are to the detriment of science and the teaching of science in society. 12 The colour plates are small, not of the highest quality of reproduction and some have been badly cropped. The insect depicted in Fig 10.1, p.259, is not the well-known yponomeutid pest of cabbages, the diamond back moth, but a species of a different family, Tortricidae probably Epinotia bilunana, an innocent birch feeder. The people who buy this volume, apart from book collectors, are going to get a huge amount of information derived from the author’s great experience. Facts are presented and ideas for further investigation suggested. It is possible that the work of Young (1997) appeals more to the person who has already developed an interest in moths, dealing with many topics under multiple headings and often including practical advice for study. Each New Naturalist offering seems to strike new chords with readers regardless of their own subject bias. The library of the dedicated field naturalist includes many titles on many subjects but copies of the New Naturalist volumes will surely always form the core. During the life of the series this is the second book on moths, with over 2,500 different species in Britain, and quite different from the first. Butterflies (60+ species) have yet to be revisited but there have been three treatments of amphibians and reptiles, a group with just over a dozen species! There does appear to be an imbalance and duplication in respect of coverage and many topics have not been addressed at all such as native woodlands, urban wildlife, alien animals, beetles, history of natural history, estuaries and so on. Perhaps this is one of the joys of natural history - it is endless in its possibilities. Long may the New Naturalist books appear to enlighten us. REFERENCES Ford, E.B. (1946) Butterflies, The New Naturalist No. 1, Collins, London, pp 368. 56 colour plates, 65 black and white plates or diagrams. Ford, E.B. (1955) Moths, The New Naturalist No. 30. Collins, London, pp 266. 77 colour plates, 90 black and white plates or diagrams. Young, M. (1997). The Natural History of Moths. T & AD Poyser Natural History, London. 27 1 pp. 1 6 colour plates, 70 black and white drawings or diagrams. E.G. Hancock THE SHIELDING AND DROVE WAYS OF LOCH LOMONDSIDE John Mitchell Produced by Jamieson and Munro in association with Stirling Council Library Service. ISBN 1 870 542 43 6, 2000, 31 pp. black and white photographs, £2.95 Obtainable from Shiela Miller, Stirling Council Library Headquarters, Borrowmeadow Industrial Estate, Stirling Council, FK7 7TN. This attractive and very informative pamphlet gives the history of shieldings and identifies their sites in the area. It likewise deals with the movement of Highland Cattle along the drove ways to fairs in the Central Belt and gives the locations of such roads in the area. Although detailed map references are added, sketch maps would have been an extra help. Ruth H, Dobson TREES OF BRITAIN AND NORTHERN EUROPE Alan Mitchell, illustrated by John Wilkinson Harper Collins, London. New Edition 1988, reprinted 2001, pp. 228, over 600 tree illustrations in colour. (Collins Pocket Guide). ISBN (pbk) 0 00 219857 6, £14.99 This book was fully reviewed in the “Glasgow Naturalist” 21, pt 4, 1988, the only obvious difference between the two printings being the design on the cover and the price (twice that of 1988), while colour reproduction has been improved somewhat. It continues to be a very good book for use in the field with compact but comprehensive information, numerous colour illustrations and a light but hard wearing cover. It is still good value for £14.99 Ruth Dobson THE BROADS Brian Moss New Naturalist No. 89. Harper Collins, London, 2001.392 pp., 29 colour and numerous black & white illustrations including maps. Softback ISBN 0 00 712410 4, £19.99; hardback ISBN 0 00 220163 1, £34.99. Professor Moss's welcome book The Broads takes quite a different approach to the subject than its 1965 predecessor with the same title in the well respected "New Naturalist' series. Gone, for example, are the old-style check lists of species - which in most cases were probably far from complete - to be replaced by a more detailed environmental account of this outstanding waterscape in East Anglia. The author discusses all aspects of the history of the Broads, from their creation to the present time. The ills which have beset these unique wetlands brought on by the effects of modern day living, together with legislative and management solutions to the problems, are thoroughly aired. Up-to-date and extremely readable, this book is unreservedly recommended. As any enthusiast of the "New Naturalist' series will confirm, the earlier version of The Broads by E.A. Ellis is now a much sought after collector's item, with anything up to three figures asked for a clean second-hand copy still wrapped in its dust jacket. So, although the price of the hardback of this new work may seem a trifle expensive, throw caution to the wind and go for it. John Mitchell 113 COLLINS BIRD GUIDE Killian Mullarney, Lars Svensson, Dan Zetterstrom Peter J. Grant Harper Collins, London, 2000. (Large Format Edition) ISBN 0 00 7100 82 5, (hbk) 398 pp. Over 3,500 painted illustrations, £29.99 An A4 sized version of the smaller original guide reviewed in The Glasgow Naturalist three years ago. Essentially this is a field guide that would require a vehicle to transport it around. What is to be gained in producing an over-sized field guide to Europe's birds? Actually quite a lot - the illustrations are bigger and therefore more detailed and shows better the artists work, and the text has also been revised and enlarged. Unfortunately some species treatments have not been brought up to date -such as the "Mediterranean Shearwater" - now regarded as two species. This apart, the illustrations are excellent and this larger edition perhaps serves as more of a reference work for the home library than a field guide. It would no doubt also benefit those who suffer with failing eyesight. At £29.99 it is quite expensive but probably represents good value at today's prices. Bernard Zonfrillo SMALL FRESHWATER CREATURES Olsen, Sunesen and Pedersen Oxford Natural History Pocket Guide, Oxford University Press, Oxford 2001, 229 pp over 800 colour illustrations. Hardback, ISBN-0- 19-850798- 4. £12.50. If, like myself, you find it difficult to pass by a pond, or even a ditch, without peering into the water to discover what small water creatures are lurking on or under the surface, this is the book for you. The paintings of insects and their larvae of about 4 to a page have plenty of explanatory text. The itemisation is surprisingly extensive, covering moths whose larvae feed on water plants, flies, wasps, spiders, worms, leeches, sponges, amphibians, and snakes etc. Perhaps more interesting are the specialised habitats which are described in detail, such as, animals on floating leaves, animals on the surface film and animals on stones in streams. Although some locations are given, the coverage of the book is Northern Europe which could lead to misidentification. I think an improvement would be for the description to be followed by abbreviations indicating countries where the species are found. If you are trying to identify a fly or beetle you may well not find it listed as the total number of animals listed is only 500. Ian C McCallum SMALL WOODLAND CREATURES Olsen, Sunesen and Pedersen Oxford Natural History Pocket Guide, Oxford University Press, Oxford. 207pp over 900 colour illustrations. Hardback, ISBN- 0-19-850797 6, £12.50. This is an attractive field guide with paintings of more than 900 creatures you are likely to come across in a walk through the woods. All the usual creatures are covered ;- butterflies, moths, beetles, wasps, spiders, slugs, snails, ants, dragonflies, flies, hoppers, bugs etc. There are excellent sections on environments. Anyone who collects fungi is soon aware that fungi carry its own fauna, when various beetles and maggots start to appear from the collection. There are other environmental sections on ant heaps, Dutch elm disease, carrion and galls. Perhaps the section that will be of most interest to people is the section on invertebrates which attack people. The only comment I would make which also applies to the companion volume is that it would be helpful if country locations were indicated by abbreviations after the entries. Ian C McCallum ANIMAL TRACKS AND SIGNS Band Preben and Dahstrom Preben Oxford University Press, Oxford, 2001. 264pp, hardback with numerous colour photographs, colour illustrations, drawings and diagrams. ISBN 0 19 850 796 8, £10.00. This is a well written guide to the signs left by the animals and birds which frequent Europe. These signs range from gnawing, burrowing, territorial marking, droppings, pellets and animal remains. The difference between “barking” and “fraying” by deer was clearly explained. Particularly fascinating was the chapter on droppings, something that even those who seldom leave an urban environment will find useful, enabling, for example, in identification of excrement of the house mouse. This book is a real “eye-opener” to the presence of life that often goes unnoticed. The illustrations, photographs and diagrams are of high quality and the cross- referencing within the text is extremely useful as is the index. The book is pocket size and is ideal for taking on field trips and will be a valuable tool for both the general reader and the experienced naturalist. Margaret M.H. Lyth 114 CONSERVATION BIOLOGY Andrew S. Pullin Cambridge University Press, Cambridge, 2002, 345pp., softback with numerous illustrations. ISBN 0 521 64482-8, £27.95. Andrew Pullin has taught Conservation Biology at the Universities of Keele and then Birmingham for a good many years. This book is constructed from that experience and is intended as an introductory text for undergraduates. He is quite explicit about attempting to provide a competitor to the prevailing texts (Primack; Meffe and Carroll) whose emphasis is predominantly North-American. The book concentrates on conservation biology as an applied science that has grown out of ecology. Pullin writes "The relationship of conservation biology to ecology is rather like that of emergency surgery to health care... It is a crisis discipline in which action must be taken despite lack of sufficient knowledge. Because to wait would mean certain destruction". This means that conservation is necessarily value laden, but Pullin regards this as true of most applied sciences. Despite this, the book deliberately only touches on the legal, political, economic and social aspects of conservation. Pullin claims that these aspects are "often covered inadequately in conservation biology and I did not want to repeat the mistakes". The book has 15 chapters, divided into three sections: 1- two introductory chapters on biodiversity and the characterisation of ecosystems; 2- four chapters on the threats to biodiversity: habitat disturbance, habitat loss, non-sustainable resource use; 3- nine chapters on conservation biology: its development as a science, and the various aspects of its implementation such as species and habitat protection, ex situ and in situ measures and restoration ecology. A key chapter (in my view) is on Conserving the Evolutionary Process. Early on, Pullin nails the myth that there is an ecological stability that we should aspire to maintain. But not only do populations fluctuate and migrate: species also naturally become extinct, and new species evolve. Pullin sees a long-term aim of Conservation Biology as the conservation of the evolutionary process itself Each chapter closes with a summary, a set of discussion points, a list of further reading sources and a set of web-sites. Each chapter includes one or more Boxes (in blue) where particular examples are given in fair detail. There are many illustrations, including some rather beautiful ones of habitats but, particularly usefully, lots of data figures, tables and distribution maps. The book ends with a reference list to all cited papers and books, and an adequate index. The book is very clearly written and well aimed at its target audience. The examples used are world- wide, but UK students will find much more that is familiar here than in the competitor North American texts. Part of Pullin's own expertise is in invertebrate animal conservation, and this brings in some nice and less familiar examples. 1 would like to make two general criticisms. First, the examples are often given to a rather low level of detail and, although there are sometimes cited avenues for further exploration of the topic, this is not always so. For example, the declining amphibian story is given in a Box with a single reference (Lips, 1998) and no web-site: it would have been easy to give the Declining Amphibian Populations Task Force (DAPTF) web-site. And another: the key captive-breeding story on the black-footed ferret is summarised, but has no references at all: good reviews are available. The American texts do better in this respect. Second, and more fundamentally, by excluding politics, economics etc., the book pays insufficient heed, in my view, to the importance of people. Without the support of people, conservation efforts are likely to fail and some of the biggest conflicts in world-wide conservation derive from attempts to exclude people from areas of high conservation value. The conservation biology course I teach puts conservation ethics and the role of people at the start, and major international conferences such as the Rio Earth Summit (1992) and the recent replay in South Africa explicitly link environmental issues to development. Overall, I found this book a very good introduction, and well worth recommending to students. The omissions may well result from the publisher's concern to keep costs down. I hope the book will succeed well enough that a second edition can deal with some of the deficiencies. Roger Downie COLOUR IDENTIFICATION GUIDE TO THE GRASSES, SEDGES, RUSHES AND FERNS OF THE BRITISH ISLES AND NORTH WESTERN EUROPE Francis Rose Penguin books Ltd, Harmondsworth, Middlesex, 1989, reprinted 2000.240pp. hardback with colour plates showing over 350 species. ISBN 0 670 80688 9, £45.00 This book complements The Wild Flower Key previously written by the author in that it covers the groups of vascular plants not dealt with in that work. The aim of the original publication was described as being a guide to plant identification in the field, with and without flowers of the plants of the British Isles and Northwestern Europe. The work continues to have appreciable sales. The present publication deals with grasses, sedges, rushes, wood-rushes, clubmosses, quillworts, horsetails, adder's-tongues and ferns. Only a few hybrids are included, the author considering that it was not possible to deal with these in the space available and refers the reader to other publications. 115 It is in hardback fonn and running to 240 pages, is not particularly suitable for use in the field. Main and often subsidiary keys are given to aid the identifieation of plants in each group or sub-group and have been well thought out. The descriptive detail is of high standard and the illustrations are excellent. For the sedges it is particularly useful to have these in colour. Detailed drawings of particular parts are always added where these are important for identification. The amount of work involved in producing the book has been immense. I have used the publication as an aid to identification throughout the 2001 recording season and found it to work well in practice. It is definitely a useful addition from the point of view of plant identification and is well produced. The only drawback to widespread sales of this book will be the price, but I consider that it is worth the extra to have the illustrations in colour. During 2001 I had occasion to meet the author and without informing him why I had the book, had him autograph the Society's copy! P MacPherson TREES: THEIR NATURAL HISTORY Peter Thomas Cambridge University Press, The Edinburgh Building, Cambridge CB2 2RU, 2000. 286pp., softback with many monochrome drawings and a few black & white photographs. ISBN 0-521 - 45963-X, (pbk) £16.95, (hbk) 0 521 45351 8, £50.00 The aim of the author was to create a book that would describe how trees work, grow, reproduce and die. He has achieved his aim, writing with considerable clarity about the biology of trees, starting with a brief look at the fossil record of conifers and hardwoods. He defines his terms clearly at the outset of the book and the text is supplemented with an adequate number of clear diagrams, summary tables and black and white photographs. The bulk of the book is devoted to the biology of the main organs of a tree, namely, roots, trunk, leaves, flowers, seeds and fruits. These topics are dealt with in a most comprehensive and lucid manner, bringing together a huge number of interesting facts about trees that would otherwise only be found seattered through many botanical and arboricultural textbooks. The main diffieulty is that of the title. One might expect that a book purporting to deal with the natural history of trees would contain details of their ecology. No reference will be found to this huge aspect of the natural history of trees. Neither the index nor the text contains any reference to the ecological status of different species, such as whether they are light demanders or shade bearers. Aspects of ecology relating to succession such as pioneer, serai or climax species are not approaehed. An excellent chapter, however, is devoted to trees and the environment, containing comprehensive references to a plethora ofbiotic and abiotic factors impinging upon the life of trees. This book is a valuable source of reference for anyone interested in trees and how they work. Bob Gray PHOTOGRAPHIC GUIDE TO THE BUTTERFLIES OF BRITAIN & EUROPE Tom Tolman Oxford University Press, Oxford, 2001, 320pp, over 500 colour illustrations ISBN 0 19 850606 6 £16.50 softback ISBN 0 19 850607 4 £35.00 hardback This book aims to help the user to find and identify butterflies from anywhere in Europe, including the Canaries, Azores, Madeira and all the Aegean islands. The colour photographs are of very good quality, and despite their small size are clear and backed up by descriptive, although limited text. This appears on the whole to be accurate and informative, and should allow identification of most butterflies. Not all species are illustrated - images of most of the butterflies from the Azores and Madeira are unfortunately conspicuous by their absence. I like the way there is a different coloured edge to the pages for different families, allowing the reader to quiekly locate for example the Pieridae (green) or Lycaenidae (blue), without searching through lots of pages to find them. The Maps are very clear, with even restricted distributions showing up well, and areas where butterflies may be encountered as migrants, shown in a different colour. Unfortunately a few maps show an over-generalised distribution, e.g. the Pearl-bordered Fritillary, which appears to occur throughout the whole of England, Wales and Scotland, and is described as ‘widespread, locally common’, which it is not. The map for the Brimstone shows the butterfly occurring throughout the Scottish Lowlands. Unfortunately this is not the case, as the text correctly states, it is absent from Scotland. Despite these few errors, if you are going to be travelling around Europe and want to identify the butterflies you see, I would recommend this book as a useful guide. Richard Sutcliffe 116 THE VARIETY OF LIFE Colin Tudge Oxford University Press, Oxford, 2002, 684 pp., paperback with numerous monochrome drawings, ISBN 0 19 860426 2, £14.99. This is a soft back reprint of the original which was favourably reviewed they the undersigned in the Glasgow Naturalist, v. 23 pt 5 (p.73) in 2000. The reduction in price from £35 will be greatly welcomed by prospective buyers, but is should be noted that the minor defects point in the review have not been corrected. Ronald M. Dobson SEABIRD NUMBERS AND BREEDING SUCCESS IN BRITAIN AND IRELAND, 1999 A.J. Upton, G. Pickerell & M. Heubeck Joint Nature Conservation Committee, Peterborough, 2000.ISBN 1 86107 507 3, 60 pp. numerous graphs and tables. £10.00 The Seabird Monitoring Programme, under the auspices of the JNCC, and with the collaboration of RSPB and SOTEAG has for a number of years now amassed data on breeding seabirds in the UK and Ireland. Several wildlife groups. Trusts, Government and Non-Government Organisations, as well as amateur and professional ornithologists, feed data into the system from a plethora of sample plots on islands, cliffs, beaches and harbours around the entire coast. Out of it all comes a fairly comprehensive picture of how well our seabirds are faring. This in turn may reflect the condition and productivity of the sea in relation to the seabird's feeding eeology. Nearly all seabirds are monitored and the data and variability of success between different parts of the country makes fascinating reading. For anyone remotely interested in seabirds or the marine ecosystem this is essential reading and excellent value. Bernard Zonfrillo AN ESSENTIAL GUIDE TO BIRD PHOTOGRAPHY Steve Young Guild of Master Craftsman Publications, East Sussex, 2001, 166 pp. soft back with over 300 coloured photographs. ISBN ! 86108 193 6, £16.95 If you are looking for an informative and attractive book about how to begin taking bird photographs, then this work by Steve Young is well worth examining. The author is a professional photographer from Liverpool who was inspired by Eric Hosking’s famous book An Eye for a Bird. Young has gained much experience since he took up bird photography in the 1970s and sets out to instruct and encourage the beginner in this specialist branch of photography. In recent years there have been many books published on bird photography. What is different about Young’s book is that he is willing to include a number of his failures as examples of what can go wrong and he demonstrates how to rectify the problems. To this end the book is copiously illustrated in high-quality colour throughout with over three hundred individual bird pictures plus examples of suitable equipment. There is, however, considerable duplication of the bird species illustrated. Common birds are well represented. An impressive series of behavioural studies of the familiar mute swan serve as encouragement to the beginner who may not have the time or equipment to tackle more secretive and rarer subjects. Steve Young himself travels the length and breadth of the country in pursuit of rare migrants and pictures of many of these unusual and spectacular birds are included. I was pleased to notice that the author discourages photography of birds at the nest claiming, correctly in my opinion, that this approach endangers successful breeding. The author also provides plenty of tips to improve your filed craft. This new guide will be a worthy addition to any photographer’s library. If you are a keen birder this impressive collection of bird pictures is worth having at any price. T. Norman Tait 117 OBITUARY Dick Hunter (1906=2002) Dick Hunter who died, aged 95, on Sunday 24th February 2002 at 9.00 pm at Drumchapel Hospital, was an Honorary Member of both the Glasgow and Edinburgh Natural History Societies. He was also an active member of the Ramblers' Association, the Scottish Wildlife Trust and a prolific watercolour artist. Dick was born in the village of Slamannan in Stirlingshire. His mother, of whom he spoke fondly, brought up a family of seven. Life was hard, although it was only on looking back that he realised this. He often spoke of his school days and would tell of the children going to school in the summer with bare feet. He also spoke of some children going to school in the winter with bare feet. He had happy memories of his childhood and could recount vividly conversations he had had when he was five years old. When, for instance, his sister would clypeus to his mother that he had stopped stirring the porridge. As a young man he worked in the coalmines. Even in later years he was happier in a confined situation rather than on an exposed open hillside. About this time his mother died and his father remarried. After the remarriage, Dick went to live with a married sister in the Coatbridge area. These were the days of the depression and he would say that his sister should have been made Chancellor of the Exchequer because of her ability to run the household on a pittance. At this time he was a keen cyclist and covered the length and breadth of Scotland. After a spell working in a brickworks in Bonnybridge he joined the army in 1939 and was posted to India and Burma where he was a radio mechanic. He would say that he was not a very good soldier. One day after having a cup of tea in a cafe he left without his rifle. He returned two days later to find his rifle still propped up in the comer where he had left it. His memories of Burma were of the exotic vegetation. He always wanted to revisit Burma when he became a proficient botanist, but never did. Demobilised, he returned home, constructed tennis courts, and then maintained bowling greens until he retired when he was 75. During his tennis court days one of his assistants 'knew ' the flowers. This sparked an interest in him to study botany and from his mid 40's developed a passionate interest in plant life. He was a regular at many of the Field Centres where he attended courses on flowers, grasses, mosses, sedges, fungi and trees. He joined the Glasgow and Edinburgh Societies and became a very popular and competent excursion leader. Apart from his official outings he delighted in taking groups of friends around Mugdock looking at mushrooms, sedges, plants, and mosses. A large number of people benefited from his good humour, patience, and expert tutoring on these outings. He was probably happiest in Mugdock where he had intimate knowledge of the locations of the rarer fungi. Most of us remember his gleeful chuckle when he discovered some hidden treasure. He delighted in introducing people to the fascinating world of fungi and explaining how to identify the different species. Occasionally I would ask him to explain the diagnostic features of a particular species. He explained that he recognised a specimen as he would a person. You did not need to know in detail a person's height colour of hair etc. to recognise him; and it was the same with plants. Occasionally 1 have seen him discussing in a very learned way with visiting botanical dignitaries on some finer point of mycology. Not bad for some one who started work in the mines! When he came across a fungus he could not identify he was known to take a spore print, make a watercolour painting of the fungi and sent them off to Professor Roy Watling, who, as always, would be helpful and reply to Dick with the answer. On his 90th birthday the local Ramblers Group hosted a party for him in Dougalston Golf Club Restaurant on the 13th April 1996, when local artist Priscilla Dorward presented him with a splendid portrait of himself The Glasgow Natural History Society also honoured his 90th birthday, at a celebration dinner at the Burnbrae Hotel when the late Camilla Dickson made a cake for him with 90 candles on it. After blowing out the candles he commented 'If I had known that I was going to live as long, I would have taken better care of myself Dick was cremated at Clydebank Crematorium on Wednesday 6th March at 2.30 pm. The service was a Humanist one in accordance with his wishes. He was one of Nature's gentlemen. I. C. McCallum and J.H. Dickson 119 PROCEEDINGS 2001 The Chaimian, place*, number present, lecturers name and title of lecture are given for most meetings. *BOB Boyd Orr Building GKB Graham Kerr Building WILT Western Infirmary Lecture Theatre 1 1th January Bob Gray, GKB, 35, 32nd Paisley International Colour Slide Exhibition, Natural History Section, Compiled by members of Paisley Colour Photographic Club and presented by Wineford Brown and projected by Jim Campbell. 9th February Prof Jim Dickson, BOB, 235, “Plants and the Tyrolean Iceman: New Discoveries”, speaker. Professor Klaus Oegg (joint with Glasgow Archaeological Society). 13th February Bob Gray, GKB, 45, “The Rhynie Chert”, speaker. Dr Lyall Anderson. 27th February Bob Gray, GKB, 26, 71st AGM. Reports were presented on activities during 2000, and elections held, resulting in appointments as shown opposite. Membership stands at 254, made up of 194 oridinary, 29 family, 17 associate, 4 school and 10 honorary members. There were 3 Council Meetings and the executive met informally as required. AGM followed by a video taken by the Macpherson family at the launch of the Changing Flora of Glasgow & a talk -“Waders & Wildlife of Arctic Alaska”, by Jane Reid. 13th March Bob Gray, GKB, “The Plants of the Falkland Islands”, speaker, Kerry Dalby. 3rd April Bob Gray, GKB, 48, “Oxford University Expedition to Western Himalayas 2000”, speaker. Dr George McGavin. 15th May Bob Gray, GKB, 30, “Wildlife Crime Detection”, speaker, John Simpson. 15th-16th June BOB, 150th Anniversary Conference; “Alien Species: Friends or Foes”. 17th June Summer Social, Forth & Clyde Canal Trip, Kirkintilloch Followed by meal at The Stables. 18 Excursions took place during the year. 28th September Bob Gray, GKB, welcomed members to the Exhibition Meeting with Wine and Cheese. 9th October Bob Gray, GKB, 23, “Pandora’s Box - Developing a British Fern Collection”, speaker Prof Alastair Wardlaw. 24th October Dr Jan Lindstorm (of DEEB, University of Glasgow), WILT, 255, “Co-operation in Mammals”, Prof T H Clutton-Brock, DEEB/GNHS Annual Lecture, Blodwen Lloyd Binns Lecture. 9th November Bob Gray, GKB, 31, “Lime Trees at the Border”, speaker, Prof Donald Piggott. 13th December Bob Gray, Glasgow University College Club, Christmas Dinner followed by talk - “Seals and Seabirds of the South Atlantic” given by Dr Dominic McCafferty. OFFICERS AND COUNCIL SESSION MMI 2001 President: Robert Gray, B.Sc., M.I.Biol. Vice Presidents: Edna Stewart, B.Sc. Ian C. McCallum, C.Eng., M.I.C.E., F.I.H.T. Ruth H. Dobson, B.Sc., M.Sc. Councillors: Eric Curtis Prof James Dickson, B.Sc., Ph.D., F.L.S., F.R.S.E Russell Semple Carole McLay David D Wylie, B.Sc. General Secretary: Kirsty A Kennedy-Wylie, B.Sc., M.Sc. Treasurer: Morag Mackinnon, B.A., B.Sc. Librarian: Joan Chapman Editor Roger Downie, B.Sc., Ph.D. Section Convenors: Keith Watson, B.Sc., M.Sc. (Botany) Richard Weddle, B.Sc. (Computer) Julian Jocelyn (Geology) Ian C. McCallum, C.Eng., M.I.C.E., F.I.H.T. (Excursions) Sandy McNeil (Ortnithology and Photography) E. Geoffrey Hancock, B.Sc., F.M.A. (Zoology) Assistant Secretaries: Catherine D. Aird, M.A., A.L. (Library) Fiona Giffard, B.Sc. (Minutes) John Lyth, B.Sc. (Publicity) Hazel Rodway (Social) Newsletter Editor: Prof Norman R. Grist, B.Sc., M.B.Ch.B., F.R.C.P., F.R.C.Path. Editorial Board: The Editor Allan McG. Stirling Prof James Dickson B.Sc., Ph.D., F.L.S., F.R.S.E. Iain C. Wilkie, B.Sc., Ph.D. BLB Administrators: Peter Macpherson, F.R.C.P., F.R.C.R., D.T.D.C., F. L.S. 120 THE GLASGOW NATURALIST - Advice to Contributors. The Glasgow Naturalist publishes articles, short notes and book reviews. Book reviews are commissioned by the Librarian. Books reviewed are kept in the Society's Library. Articles and short notes should be sent to The Editors, Azra and Peter Meadows, The Glasgow Naturalist, Graham Kerr Building, University of Glasgow G12 8QQ. Short notes are edited by Mr A. McG. Stirling. Articles and short notes are refereed. Acceptance of articles and short notes are the responsibility of the Editors. The journal is published yearly. The subject matter of articles and short notes should concern the Natural History of Scotland in all its aspects, including historical treatments of natural historians. 2. Short notes should not normally exceed one page of A4 single-spaced. They should be headed by the title and author's name and address. Any references cited should be listed in alphabetical order under the heading References. There should be no other sub-headings. Any acknowledgements should be given as a sentence before the references. Short notes may cover, for example, new stations for a species, rediscoveries of old records, additions to records in the Atlas of the British Flora, unusual dates of flowering, unusual colour forms, ringed birds recovered, weather notes, occurrences known to be rare, interesting localities not usually visited by naturalists, and preliminary observations designed to stimulate more general interest. 3. Articles should be more substantial than short notes. The maximum length should not exceed approx 6000 words including references and equivalent space for tables and references. Longer articles should be discussed with the Editors before submission. They should be headed by the title and author's name and address. Any references cited should be listed in alphabetical order under the heading References. The text should be divided into sections with sub-headings as appropriate . (e.g. Abstract, Introduction, Methods, Results, Discussion, Conclusions, Acknowledgements, References) 4. References in articles and short notes should be given in full (please do not abbreviate journal titles) according to the following style: Pennie,I.D. 1951. Distribution of Capercaillie in Scotland. Scottish Naturalist 6?), ■ Wheeler, A. 1975. Fishes of the World. Ferndale Editions, London. Grist N.R. & Bell, E.J 1996. Enteroviruses. Pp. 381-90 in Weatherall, D.J. (editor) Oxford Textbook of Medicine. Oxford University Press, Oxford. 5. Nomenclature of vascular plants should be as in Stace, C.A. (1997). The new Flora of the British Isles, (Second Edition) Cambridge University Press, Cambridge. Normal rules of zoological nomenclature apply. Please use lower case initial letters for all common names e.g. wood avens; blackbird, unless the common name includes a normally capitalised proper name e.g. Kemp's ridley. Where giving distribution information by vice-county, use the following style: VC 30. 6. Submitted manuscripts (two copies) should be typed double-spaced on A4 paper. Typesetting is greatly assisted if the manuscript can be supplied on a microcomputer diskette. Authors are therefore strongly encouraged to produce manuscripts using a word processor (preferably on a PC-compatible microcomputer). However, to assist amateur naturalists, the Editor can make arrangements to have hand- written manuscripts or typed manuscripts transferred to disc. 7. Tables are numbered in Arabic numerals e.g. Table 1: they should be double-spaced on separate sheets with a title and short explanatory paragraph underneath. 8. Figures, Line drawings and photographs are numbered in sequence in Arabic numerals e.g. Fig. 1. If an illustration has more than one part, each should be identified as 9a), (b) etc. The orientation of the figure and name of the first author should be indicated on the back. They should be supplied camera-ready for uniform reduction of one-half on A4 size paper. Line drawings should be drawn and fully labelled in Indian ink or dry-print lettering or laser printed. A metric scale must be inserted in micrographs etc. Legends for illustrations should be typed on a separate sheet. The Editors are able to accept a small number of high quality colour photographs for each issue. Please consult the editors before submitting the paper. 9. Proofs should be returned to the Editor by return of post. Alterations should be kept to the correction of errors. More extensive alterations may be charged to the author. 10. Offprints.Ten offprints and one complimentary copy of the Journal are provided free of charge. Further copies may be purchased, provided that they are ordered at the time the proofs are returned. 11. Review. All submissions are reviewed by the Editors or their appointed referees. They are also assessed by the Editors for ethical considerations. Publication may be refused on the recommendation of the Publications Committee. Front Cover (photograph provided by J. Mitchell) An early French impression of water meadow management, an agricultural practice now lost to Scotland. Back Cover (photograph provided by J. Maxwell) A female willow tit emerging from the nest cavity with ringed male waiting to feed the nestlings (top); a blue tit approaching an "appropriated " stump cavity with caterpillars (bottom left); female willow tit pauses between feeds (bottom right). SMrrHSONIAN INSTtTUTION LIBRARIES imHIHlIHI 3 9088 01205 4375 e.li I c&X h^e Glasgow Naturalist Volume 24 Part 2 2004 Journal of THE GLASGOW NATURAL HISTORY SOCIETY GLASGOW NATURAL HISTORY SOCIETY (formerly The Andersonian Naturalists of Glasgow) The object of the Society is the encouragement of the study of natural history in all its branches, by meeting for readirlg and discussing. papers'and exliibiting specimens and by excursions for field work. The Glasgow Natural flistory Society meets at least once a month except during July and August, in the University of Glasgow, the Glasgow Art Gallery and Museum, or Hillhead Library. The present rates of subscription per aimum are; for Ordinary Members. £17: Family Members. £3 extra; Junior Members (under 21) £8; School Members. £1 . Payment by Duect Debit is encouraged. Further information regarding the Society's activities and membership application forms are obtainable from the General Secretary. The Glasgow Natural Flistory Society, c/o DEEB. Graham KeiT Building. University of Glasgow, G12 8QQ, Scotland. The Glasgow Natural flistory Society website is http :/Avww. gnlis.freeuk.com/. w hich also contains details of the Glasgow Naturalist. The Glasgow Naturalist The Glasgow Naturalist is published by the Glasgow Natural Flistory Society. ISSN 0373-241X. Price £4.25 plus p.& p (as applicable at the time of posting). The Glasgow Naturalist is free to members. Editors; Azra Meadows and Peter Meadows. Graham KeiT Building. University of Glasgow G12 8QQ (Tel: 0141 330 6622/6624. Email: gbza3 1 r/ udcfgla.ac.uk) Contributions are invited, especially when they bear on the natural history of Scotland. Full details of how to contribute articles or short notes are given at the end of the volume. A limited number of advertisements can be accepted and enquiries should be sent to the Editors. This publication is included in the abstracting and indexing coverage of the Bioscience Information Seivice of Biological Abstracts and the Botanical Society of the British Isles Abstracts. The following back numbers are available for purchase in their separate parts: Vols.II - VII (1890-1918); Vols. XIII - XXIII (1937-1999). Of the earlier Journals the only parts available are: Proceedings and Transactions of the Natural History Society of Glasgow Vol. II pt. 2; Vol. VI pt. I; Vol. VII. pt. 3; VoL Vlll.pts. I &2. Enquii ies regardmg prices of and orders for any of the above, or for reprints or photocopies, should be addressed to the Librarian: Mrs Joan Chapman. 121 Randolph Road. Jordanliill. Glasgow' G1 1 7DS. Scotland. Publications of Glasgow Natural History Society Alien Species: friends or foes? Edited by J.R.Dowiiie (2001). Proceedings of the GNHS 150* Amhversary Conference. Price £10.00 plus p & p. Bound copies of the follow ing may be obtained from the Librarian at the address above and at the prices show n: The Flora of the Clyde Area (Original printing). J.R. LEE, Price £1 1.00 to members of GNHS and to the book trade; £1 3.50 to others (p. & p. £1 .00 extra). This is still the only work of its type and is in diminisliing supply. A few unbound copies are available; £5 (p.&p. £1 extra). The Flora of Ailsa Craig. B. ZONFRILLO. 1994. Price £2.50 plus p.& p. The Natural History of the Muck Islands. N. Ebudes: Introduction and Vegetation with a List of Vascular Plants. R.H. DOBSON & R.M. DOBSON. 1985. Price £1.00 plus p. & p. Seabirds and Wildfowl. R.H. DOBSON & R.M. DOBSON. 1986. Price £1.00 pins p. & p. Landbirds. R.FI. DOBSON. 1988. Price £1.00 plus p. & p. The following bound reprints from the Glasgow Naturalist may be obtained from the librarian at the above address and at the prices shown. Additions to the Flora of the Clyde Area. .Tolm R. Lee (1953). £1 (p.&p.) Archives For archive information and special collections of the GNHS contact Karol Magee. Tel. 0141 287 2907. Society Microscopes The Society incoi-porates the Microscopical Society of Glasgow . Microscopes may be bonowed by members and are cunenth kept in the room of E.G. Hancock. Curator of Entomology, in the Graham KeiT Building. University of Glasgow . Front Cover (ialls of Knopper gall w asp {Audriciis qtiercuscalictis) glowing from the cups of acorns on Oak (Quercus robur). Man' Hall Hotel parkland. Erskine, September 2002. Insert: adult asexual female Knopper Gall w asp {Andriciis querciiscalicus) February 2003 - x 7. Photogiaphs © Norman Tait. Back Cover Top. Water Fern (Azolla fdictdoides) South Haugh. Flamilton. September 2003. Photogiaph © Peter Macpherson. Bottom. Japanese Knotweed (Fallopia japonica war japonica) Millport. Isle of Cumbrae. May 2004. Photogiaph © Azra and Peter Meadows. The Glasgow Naturalist Volume 24 Part 2 2004 CONTENTS EDITORIAL Azra Meadows and Peter Meadows FULL PAPERS The distribution of Cochyliinae (Lepidoptera: Tortricidae) in Scotland. Keith Bland 5-10 A survey of feral goats on the Oa Peninsula, Isle of Islay, Western Isles, Scotland, July 2003. Peter B. Copley & Donald James MacPhee 11-20 Hvdroporns scalesiamis (Coleoptera, Dystiscidae) new for Scotland. G.N. Foster 21-23 The effect of water content and compactness of soil on the survival of the New Zealand flatworm A rthurdendyits triangulates . Peter H. Gibson & Derek J. Cosens 25 - 28 The effect of temperature on the survival and distribution of the New Zealand flatworm Arthurdendyus triangulates. Peter H. Gibson , Katherine L. Ponder & Derek J. Cosens 29 - 33 Studies on the conservation biology of Irish Lady’s- tresses Orchid, Spiranthes romanzoffiana\ 1) Population sizes, grazing, vegetation height and capsule status at reference sites. Richard Gulliver, Mavis Gulliver, Margaret Keirnen, and Christopher Sydes 35 - 52 Studies on the conservation biology of Irish Lady’s- tresses Orchid, Spiranthes romanzoffiana\ 2) The establishment of 10 exclosures, dung counts and further studies on associated Junctis taxa (species and hybrid rushes) Richard Gulliver, Mavis Gulliver, Margaret Keirnen, and Christopher Sydes 53 - 68 Rediscovering the Firth of Forth Beluga. Andrew C. Kitchener and Jerry S. Herman 69 - 70 Climate change and its effects on catchment hydrology of the River Spey, Scotland, the River Neckar, Germany, and the River Alpenrhine, Switzerland. GerritKlemm 71-78 Gro’wth of Ailsa Craig slow-worms Anguis fragilis: Prey preference and temperature effects. Cara Lavery, J. R. Downie, & Suzanne R. Livingstone 79 - 85 A rich Botanical strip at Shettleston, east end of Glasgow. P. Macpherson & E.K. Lindsay 87 - 88 Distribution and population status of the otter in the Loch Lomond Area. D. J. McCafferty 89-91 The effect of fine detrital material and microbial activity on the permeability of intertidal sediments from Ardmore Bay, Firth of Clyde. Azra Meadows & Peter S. Meadows 93 - 106 Japanese knotweed (Fallopia japonica) occurrence and distribution on the Isle of Great Cumbrae. Azra Meadows & Peter Meadows 107 - 114 The early stages in the development of an ox-bow lake beside the River Enderick. John Mitchell 115-118 Alexander Patience (1865-1954): Glasgow’s little-known Edwardian carcinologist. P. Geoffrey Moore & E. Geoffrey Hancock 119- 129 The spread of Knopper gall wasps (Andricus quercuscalicis) into the Clyde Area. Norman Tait & Pearl Tait 131-132 Records of Coleoptera from Islay. O. Vorst 133 - 135 Abundance and patterns of occurrence in butterflies, Ailsa Craig, Ayrshire. B. Zonfrillo & E. G. Hancock 137 - 140 SHORT NOTES Unusual colour forms of the woodlouse Porcellio scaber (Latreille) on Mull Glyn M. Collis and Dawn Collis 141 An ancient beetle collection saved. Ronald M. Dobson 141 - 142 Large-leaved Avens {Geiim macrophyllum) established at Mugdock Country Park (VC86). Bill Hansen 142 The red-necked footman in West-Central Scotland. John T. Knowler and John Mitchell 142-143 Hawfinches at Talla reservoir. Margaret M.H. Lyth 143 Bats in Clarence Drive, Cleveden, Glasgow West End. David McNaught 143 BOOK REVIEWS Compiled by Ruth H. Dobson (reviewers names in brackets) British Bats (2003) by John D. Altringham (Dominic McCafferty) 145 Guide to the Identification of Soil Protozoa - Testate Amoebae (2003) by K.J. Clarke (Keith Vickerman) 145 Birds by Behaviour (2003) by Dominic Couzens (Bernard Zonfrillo) 145 - 146 Photographing Plants and Flowers (2002) by Paul Harcourt Davies (T. Norman Tait) 146 How to Identify Weather (2002) by Storm Dunlop (Bob Gray) 146 The National Pakrs and other Wild Places of Britain and Ireland (2002) by Jonathan Elphick and David Tipling (Margaret M.H. Lyth) 146 Biodiversity: An Introduction (2004) by Kevin J. Gaston and John L Spicer (Azra Meadows) 147 - 148 Tarns of the Central Lake District (2003) by Elizabeth Haworth, George de Boer, Ian Evans, Henry Osmaston, Winifred Pennington, Alan Smith, Philip Storey and Brian Ware (Daniel Gates) 148 - 149 Niko ’s Nature: a Life of Niko Tinbergen and his Science of Animal Behaviour (2003) by Hans Kruuk (Keith Vickerman) 149 - 150 Collins Wildlife Trust Guide to the Weather of Britain and Europe (2001) by David M. Ludlum (Ian C. McCallum) 150 - 151 Fossils at a Glance (2004) by Clare Milsom and Sue Rigby (J. Jocelyn) 151 ii CasselVs Trees of Britain and Northern Europe (2003) by David More (author) and John White (artist) (Bob Gray) 1 5 1 The Garden Bird Handbook (2003) by Stephen Moss (Joyce Alexander) 1 5 1 Bill Oddie^s Introduction to Birdwatching (2002) by Bill Oddie (Norman R. Grist) 151 - 152 The New Encyclopedia of Insects and their Allies (2002) Edited by Christopher O'Toole. (Ronald M. Dobson) 152 Wild Side of Town: Getting to know Wildlife in our Towns and Cities (2003) by Chris Packman (Margaret M.H. Lyth) 152 Windermere - Restoring the Health of England's largest Lake (2001) by A.D. Pickering (Colin Adams) 152 - 153 Keys of Larvae and Juvenile Stages of Coarse Fishes from Fresh Waters in the British Isles (2001) by A. C. Binder (Colin Adams) 153 Lakeland (2002) by Derek Ratcliffe (John Mitchell) 153 Atlas of Cetacean Distribution in North West European Waters (2003) by J.B. Reid, P.G.H. Evans and S. P. Northbridge (Bernard Zonfrillo) 153 - 154 The Phytoplankton of Windermere (English Lake District) (2000) by C.S. Reynolds and A.E. Irish (Colin Adams) 1 54 People and Woods in Scotland: A History (2002) Edited by T. C. Smout (Joyce Alexander) 1 54 The Story of Life (2003) by Richard Soiithwood (Ronald M. Dobson) 154-155 Britain 's Butterflies (2002) by David Tomlinson and Rob Still (K.P. Bland) 155 Keys to the Case-Bearing Caddis Larvae of Britain and Ireland (2003) by I.D. Wallace, B. Wallace and the late G.N. Philipson (E.G. Hancock) 155 Travellers' Nature Guides (2003) Britain by Martin Walters and Bob Gibbons; Scotland text by Kenny Taylor, France and Greece by Bob Gibbons (Edna Stewart) 155 - 156 Arable Plants - A Field Guide (2003) by Phil Wilson and Miles King (Jean M. Millar) 156 Keys to the Freshwater Microturbellarians of Britain and Ireland with Notes on their Ecology (2001) by J.O. Young (E.G. Hancock) 156-157 BLODWEN LLOYD BINNS BEQUEST REPORTS Glasgow and the Hindu Kush Azra Meadows and Peter Meadows 159 - 161 Breeding Procellariiforms in the Azores Archipelago: abundance and distribution as affected by introduced predators Ana de Leon Marti 1 62 - 163 Predation on tern eggs by European starlings in the Azores Veronica Neves 163-164 OBITUARIES Alan Muirhead Maclaurin (1907-2003) By Ronald M. Dobson 165 Agnes (Nancy) Craib (1912 -2003) By James H. Dickson 166 Irene Nove (1918-2004) By Agnes Walker 167 - 168 Proceedings 2002; Officers and Council, Session LXXIl 2002 169 Proceedings 2003; Officers and Council, Session LXXIII 2003 170 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 1-4. THE GLASGOW NATURALIST EDITORIAL Azra Meadows and Peter Meadows DEEB, IBLS, University of Glasgow, G12 8QQ Emails: and The Glasgow Naturalist journal has a long and distinguished history dating back to the 19'*’ century. It now has a unique and developing role to play in terms of the ecology and biodiversity of terrestrial, freshwater and marine environments in and around Scotland. This includes the following topics, many of which are central issues for Scotland at local, national and international levels. Biodiversity at all its levels including plants, animals and microorganisms. Taxonomic descriptions of new and rare species. Field studies of species and ecosystems. Experimental studies of speeies and ecosystems. Wildlife conservation and management. Environmental management and impact assessment. Historical and archaeological aspects of natural history. Lives and aetivities of eminent natural historians Environmental implications of waste management. Implications of climate change. Environmental geomorphology. Environmental engineering. Eco-sociological issues. Environmental economics and policy. Members of the Glasgow Natural History Society, together with our amateur and professional contributors, are essential to the future success of the Journal, and to its development as a prestige journal originating from the West of Scotland. As editors, therefore, we hope that the Glasgow Naturalist will provide our contributors and our wider reading audience with a quality service, as follows. • Production of a journal that is 2U‘ century orientated in its contents and presentation. • Encouragement of high quality papers on Scottish topics or topics that have relevance to local and national issues in Scotland. • Stimulation of a broadening of the breadth of subject areas of submitted papers, including all aspects of freshwater, marine and terrestrial ecosystems and species in Scotland. • Encouragement of younger biologists, whether amateurs or professionals, including postgraduate and undergraduate students, to submit quality research and review articles and short notes. • Provision of an effective professional standard of refereeing and editing for the journal. . • Provision of leadership for amateur and professional contributors to the journal in relation to subject topic and paper layout for the journal, especially in relation to current electronic facilities. The Glasgow Naturalist receives papers and short notes on a range of subject areas. After examining the subject areas covered by The Glasgow Naturalist for the last decade we felt it timely to assess what the broad areas and their distribution are. We have reviewed the breadth and statistics of subject areas, which have been covered in the Glasgow Naturalist during 1996 to 2001 (Volume 23 parts 1-6) and during 2002 to 2004 (Volume 24 parts 1- 2). We have classified the papers and short notes into four broad subject areas: general, terrestrial, freshwater and marine, whose overall percentages are illustrated in Figure 1 . The contents of the current volume (volume 24 part 2) are shown in table 1. Table 2, at the end of this editorial, provides a detailed breakdown for volume 23, parts 1 to 6 which were edited by Roger Downie, and volume 24 parts 1 and 2 which have been edited by ourselves - the eurrent editors. The overall pattern throughout this period has been very similar. The breakdown of the subject areas of the papers and short notes is interesting. In volume 23 parts 1 to 6, 68% of the papers and short notes were on terrestrial topics, 1 7% on freshwater topics, 8% on marine topics, and 7% on general topics. In volume 24 parts 1 and 2, In volume 23 parts 1 to 6, 76% of the papers and short notes were on terrestrial topics, 8% on freshwater topics, 1 2% on marine topics, and 4% on general topics. Summing over both volumes of the journal (volume 23 parts 1 to 6, volume 24 parts 1 and 2), there were 159 (70%) terrestrial papers and short notes, 35 (15%) freshwater papers and short notes, 21 (9%) marine papers and short notes, and 14 (6%) general papers and short notes. We hope that in future the distribution of papers between terrestrial, freshwater and marine subject areas will become more even. Scotland is recognised globally for its unique freshwater lochs and its magnificent marine coastline, as well as for its dramatic mountain and island scenery. This should be reflected in the journal. 1 □ , n Terrest. FreshW. Mar. Subject Area Figure 1. Percentage of subject areas covered in the Glasgow Naturalist volume 23 parts 1 to 6, and in volume 24 parts 1 and 2. EDITORIAL PROCEDURE For contributors who are interested to know the process of editing contributions for a journal, we have listed the following procedure. 1. Paper received in hard copy and electronically from authors. Authors informed of receipt. 2. Initial acceptance or rejection. 2.1 Paper accepted subject to review by referee. Go to 3. 2.2 Paper rejected. Authors informed. 3. Hard copy of paper sent to referee for review. 4. Hard copy of paper received from referee with comments. 4.1 Referee states that paper is suitable for publication without modification. Go to 8. 4.2 Referee states that paper is suitable for publication, but needs revision. Go to 5. 4.3. Referee rejects paper. Authors informed. 5. Hard copy of paper with referee’s comments, together with electronic version sent to authors for revision. 6. Hard copy of paper with referee’s comments, together with electronic version modified in accordance with referee’s comments, received from authors. 7. Editors check that modified electronic version contains all referee’s comments. 7.1 Modified electronic version needs further revision, and is returned to authors for further correction. 7.2 Modified electronic version is accepted for publication. 8. Modified electronic version formatted for journal, and sent as hard copy to authors for proof reading. Minor alterations only allowed. Category Nos. Full Papers 17 Short Notes 7 BLB Reports 3 Book Reviews 28 Obituaries 3 Table 1. Current Issue of the Glasgow Naturalist Vol. 24 Part 2 2004. 9. Hard copy of authors proofread copy received from authors. Alterations incorporated into formatted electronie version of paper, ready for publication in next issue of the journal. 10. Short notes. Book reviews, BLB reports and obituaries received from collators. 11. Papers, short notes, book reviews, BLB reports, and obituaries collated for next issue of journal, and sent to printers. 12. Final proofs of complete journal issue received by editors from printers, and checked for errors in pagination. 13. Proofs with corrections returned to printers for printing. 14. Print run received from printers. Copies of journal and reprints of papers distributed to members of the Glasgow Natural History Society and authors. In general, following the submission of a paper, the whole process running from item 1 to 14 takes between 1 0 and 20 months. This is the time it takes for a paper to be published in any national or international peer-reviewed journal. CONTENTS OF THE CURRENT ISSUE In the current issue the flora include micro- organisms, orchids, and the Japanese knotweed, beside other terrestrial plants, while the faunal groups include the flatworm, slow worm, butterflies, moths, beetles and woodlouse, hawfinches, goats, cetaceans, otters and bats. Here we list brief accounts of the full papers followed by short notes in alphabetical author sequence as they appear in this volume. The issue opens with a paper by Bland who reappraises the Watsonian 2 vice-county distribution of the Scottish Cochyliinae (Lepidoptera), reporting that out of the 47 species resident in Britain 32 have been recorded in Scotland. This is followed by Copley and MacPhee’s survey of feral goats on the Oa peninsula. Isle of Islay, where they recorded 400 feral goats. The authors predict an increase in goat population in Oa. Foster describes a beetle species Hydroporiis scalesianus inhabiting fens new to Scotland. The two laboratory-based papers that follow are by Gibson et al., on the New Zealand flatworm Arthurdendyus triangidatiis. One paper tests the effect of water content and compactness and the other the effect of temperature on the survival of the flatworm. 30% water by weight was shown to give the optimum survival and the species demonstrated preference to 19°C. We then come to a second pair of longer papers now led by Gulliver and co-workers on the conservation biology of the Irish lady’s-tresses orchid, Spiranthes romanzqffiana. The first paper deals with the population sizes, grazing, vegetation height and capsule status. The study shows grazing on leaves by slugs and vertebrates and on flowering stems by domestic stock and rabbits. The fact that this species can occur in an underground form for up to six years is interesting, together with the presence of mycorrhiza which is a source of fungal organic carbon. The sister paper by Gulliver et al. looks at the establishment of 10 enclosures, their sheep and cattle dung counts and hoof holes, and the frequency of Juncus articulatus and Juncus acutifloris x articulatus. Kitchener and Herman examine the rare sightings of the Firth of Forth beluga Delphinapterus leucas. They have studied the Tumer/Monro skull of a beluga specimen whose whereabouts is unknown. The damaged cranium and the missing teeth are suggestive that this skull is that of the missing Firth of Forth beluga. Klemm’s paper discusses climate change and its effect the hydrology of three European rivers, River Spey (Scotland), River Neckar (Germany) and the river Alpenrhine (Switzerland). The paper predicts more flooding in the Spey and consequential runoff and winter seasonality. In the Neckar the winter and spring runoffs are higher than the summer and autumn runoffs. The River Alpenrhine shows four independent seasons, but would change with increase in warming from a snowmelt fed river to a rain-fed river. The paper by Lavery et al. investigates the growth rate of Ailsa Craig slow- worm Anguis fragilis and its prey preference. The laboratory experiments show that slow-worms grew faster and ate more at temperatures of 27- 28°C and showed a decrease in this activity at 1 8°C. There was no size preference for slugs over a range of 0.5-2. 5g. This is followed by Macpherson and Lindsay’s record of the 266 floral taxa along a strip in Shettleston, east end of Glasgow, and this included seven plants which were the first record for the vice-county. McCafferty has conducted a spraint, track and prey remain survey to assess the distribution and population status of the otter Lutra lutra in the Loch Lomond area. His results show that otters were widespread throughout the loch and that there was an increase in the proportion of tributaries with otters present between the late 1970s and 2002. The paper by Meadows and Meadows demonstrates the effect of fine detrital material and microbial activity on the permeability of intertidal sediments from Ardmore bay in the Clyde estuary. The laboratory experiments show that fine detrital material decreases permeability by about 75%, and that heterotrophic bacteria dramatically decreased sediment permeability by 98% while photosynthetic microbial activity decreased permeability by 38%. In the subsequent paper these authors describe the occurrence and distribution of the Japanese knotweed Fallopia japonica on the Isle of Cumbrae, and discuss the length of time the species may have been on the Island. The paper by Mitchell considers the stages in the development of an Ox-bow lake beside the Endrick River. It includes a list of plant species along the waters edge and the newly colonized aquatic plants, and describes the changes in river flow. We then come to a biographical account by Moore and Hancock on Glasgow’s little-known Edwardian amateur Carcinologist Alexander Patience, and his scientific contributions. This paper examines Patience’s involvement with the Marine Biology Station at Millport and with scientific professionals, and includes a full bibliography of his published works. Tait and Tait describe the spread of the Knopper Gall-wasp Andricus quercuscalicis into the Clyde area, together with the complex life cycle this insect undergoes. Vorst reports records of Coleoptera from Islay. This study yields 71 species from varying habitats such as the coastal dunes, river banks, moorland and sheep carrion. 18 of the 71 species have not been recorded in Islay before, and five of the 71 species are new for the Inner Hebrides. The last full paper in this issue is by Zonfrillo and Hancock and is also an Island study, but this time on butterflies from Ailsa Craig, Ayrshire. The authors report 21 species of butterflies. Five of these breed regularly and 12 are sporadic on the island, while the remaining are regarded as doubtful. Under the short notes section, Collis and Collis report the unusual colour forms of the woodlouse Parcel Ho scaber on the Isle of Mull; Dobson describes an ancient beetle collection which he saved; Hansen records the presence of Large- leaved Avens Geum macrophyllum in Mugdock Country Park; Knowler and Mitchell record the presence of the red-necked footman moth At ohms rubricollis in West-central Scotland; Lyth reports hawfinches at Talla Reservoir, Peebleshire, and McNaught observes bats in the West End of Glasgow. 3 Table 2. Numbers and percentages of full papers and short notes according to their subject area (General, Terrestrial, Freshwater, Marine). Vol. and Part Papers/ Short Notes General Terrestrial Freshwater Marine Edited by Roger Downie Vol. 23 (Parts 1-6) Papers 11 50 14 11 Short notes 1 72 17 4 Total (2=180) 12 122 31 15 % 7% 68% 17% 8% Edited by Azra Meadows and Peter Meadows Vol. 24 (Parts 1-2) Papers 2 18 4 5 Short notes 0 19 0 1 Total (2 = 49) 2 37 4 6 % 4% 76% 8% 12% GRAND TOTALS Vol. 23 and Vol. 24 Grand Total (2 = 229) 14 159 35 21 Grand Total % 6% 70% 15% 9% 4 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 5-10. THE DISTRIBUTION OF THE COCHYLIINAE (LEPIDOPTERA; TORTRICIDAE) IN SCOTLAND. K.P.Bland National Museums of Scotland Chambers Street Edinburgh EHl IJF TORTRICIDAE INTRODUCTION The recent completion of the curation of the tortricoid subfamily Cochyliinae in the collection of the National Museums of Scotland (NMS) is an opportune time for a reappraisal of the Scottish distribution of this group. Thus the following is an attempt at a comprehensive up-to-date account of the Watsonian vice-county distribution of the Scottish Cochyliinae, with the further purpose of making more widely available those records not previously published. All the 47 species of Cochyliinae resident in Britain are represented in the 3,000 specimens of this subfamily in the NMS collection. Thirty-two species have been recorded from Scotland but the authenticity of two of these species requires confirmation. Scottish specimens of this group are present in the collection amassed by A. B. Balfour, J.W.Bowhill, P.W.Brown, l.C.Christie, J.A.Clark, A. A.Dalglish, A.B. Duncan, D.W.H.Ffennell, D.R.Gifford, R.K.Greville, D.J. Jackson, G.A.T.Jeffs, R.F.Logan, C.W.Mackworth-Praed, D.A.B.Macnicol, R.M.Mere, K.J. Morton, E.C. Pelham-Clinton, T.E.D. Poore, A.Richardson and B. W. Weddell. Additional material has been contributed by K.P.Bland, P. A. Buxton, J.L.Campbell, D.L.Coates, Edinburgh University Biological Society, A.G.Long, R.I.Lorimer, I.H.K.Lyster, E.A.M.MacAlpine, P.Marwick, R.& B.Meams, J.M.Nelson, M.G. Pennington, G. Petrie, O.W. Richards, T. Rogers and A.R.Waterston. A comprehensive list of the previously published records of Scottish Lepidoptera is held in the National Museum of Scotland (NMS), Chambers Street, Edinburgh in the “Scottish Insects Record Index” (SIRI). This lists under each species the bibliographic reference and an abbreviated locality, often in the form of the Watsonian vice-county number (Shaw, 1987). SIRI contains some 130 references which include information on the Scottish distribution of the Cochyliinae. These have all been carefully checked back to the original to ensure the accuracy of the distribution data quoted here (Table 1). For completeness the author has also included his own records and all other unpublished records known to him. Watsonian vice-counties (Table 2) divide Britain up into approximately equal sized areas that people can identify with. Current unitary authority boundaries are based more on areas of equal population and so have little value for wildlife recording. Full details of the addition records are shown below. The earliest record is shown, where multiple records exist. Records accompanied by specimens have in some cases been given priority. Cochyliinae Hysterophora maculosana (Haworth, 1811) Vc. 72 Grove, Dumfries. 9. vi. 1977 A.B.Duncan. Vc. 88 Killin,Perths. 14.vi.l985 M.R. Young. Vc. 97 Fort William, Inverness. 26.V.1957 E.C. Pelham-Clinton. Vc. 98 Port Appin, Argylls. 26.V.1956 E.C.Pelham-Clinton. Vc. 105 Strath More, W. Ross. 10.vi.l963 E.C.Pelham-Clinton. Vc. 106 Alcaig, Ross. 18.vi.l988 E.C.Pelham- Clinton. Phalonidia affmitana (Douglas, 1 846) Vc. 73 Caulkerbush, Kirkcud. Larvae 27. xii. 1970 (reared) E.C.Pelham-Clinton. P. manniana (Fischer von Roslerstamm, 1839) Vc. 79 Selkirk. 28.vi. 1941 B.W.Weddell. Vc. 82 Tyninghame, E. Loth. 5.vi.l933 A.B. Balfour. The specimen is very worn and without an abdomen and so is unidentifiable. It bears an identification label '"Phalonia manniana FR.” in Alice Balfour’s handwriting. Gynnidomorpha minimana (Caradja, 1916) Vc. 80 Whitlaw Moss, Roxburgh. 3.vii.l981 K.P.Bland - erroneously published as in Vc.79. No records from Vc.79 are known to me. G. permixtana ([Denis & Schiffermuller], 1775) Vc. 98 At the 1957 Annual Exhibition of the South London Entomological Society, Pelham- Clinton (1958) exhibited specimens of Phalonia walsinghamana Pierce (= G. minimana) from Port Appin. This record was repeated by Bradley, Tremewan and Smith (1973). The identity of these specimens was corrected to G. permixtana (Pelham-Clinton, 1982). G.vectisana (Humphreys & Westwood, 1845) Vc. 73 Auchencairn, Kirkcud. 2003 E.A.M. MacAlpine. Cochylimorpha alternana (Stephens, 1834) Vc. 92 There are 3 specimens labelled “Aberdeen” in the J.A.Clark collection which was received in 1906. This is a surprising record and the number and condition of the specimens suggests they may have been mislabelled. C. straminea (Haworth, 1811) Vc. 73 Merse Head, Kirkcud. 20.vii.l996 J.MacKay. Vc. 84 Queensferry, W. Loth. Pre 1857 R.K.Greville. Vc. 96 Kincraig. 5.vii.l950 P. Harwood (Macnicol coll.). Vc. 97 Spean Bridge, Inverness. 30.vi.l965 E.C.Pelham-Clinton. Vc. 105 Kishorn, W.Ross. 18.vii.l984 P.W.Brown. 5 Agapeta hamana (Linnaeus. 1758) Vc. 72 Connansknowe, Dumfries. 26.vi.1998 R.Meams. Vc. 79 Midgehope Marsh, Selkirks. 3/4.viii.l979 K.P.Bland. Vc. 84 Winchburgh, W. Loth. 2.viii.l969 E.C.Pelham-Clinton. Vc. 95 Garten Railway Line, Elgins. 14. vi. 1975 M'.R. Young. Vc. 96 Newtonmore Golf Course, E. Inverness, l.vii.l992 M.R.Young. Vc. 97 Kilchoan, Ardnamurchan. 28.vi.1965 E.C.Pelham-Clinton. Vc. 105 Kishorn, W.Ross. I.vii.l985 P.W.Brown. A. zoegana (Linnaeus, 1767) Vc. 72 Castlehill, Dumfries. 7.viii.l973 A. B. Duncan. Vc. 73 Kirkconnell, Kirkcud. 31.vii.l977 A. B. Duncan. Vc. 83 Burdiehouse, Midloth. 7.vii.l958 D. A.B.Macnicol. Vc. 84 Winchburgh. W.Loth. ll.viii. 1965 E. C.Pelham-Clinton. Vc. 90 Lunan Bay, Angus. 7.viii. 1996 B.Goater. Aethes cnicana (Westwood, 1854) Vc. 73 Lochaber Loch, Kirkcud. 2.vii.l996 J.MacKay. Vc. 75 Kennedy’s Pass, Ayrshire 19.vii.l984 I. C. Christie. Vc. 78 Menzion Farm, Tweedsmuir. 23/24.vii.l979 K.P.Bland. Vc. 79 Thomielee, Selkirks. 19.vi.l981 A.Buckham Vc. 85 Tayport, Fife. 4.vii.l970 E.C.Pelham- Clinton. Vc. 89 Glenshee, Perths. 1976 E.F. Hancock. Vc. 96 Rothiemurchus. vii.1896 K.J.Morton. Vc. 98 Inverlochy, Argylls. 26.vi.1943 C. W.MacKworth-Praed. Vc. 100 Shiskine, Isle of Arran. 26.vi.1999 M.R.Young. Vc. 101 Tayvallich, Kintyre. 5.vii.l974 E.C.Pelham-Clinton. Vc. 102 Colonsay. Date unknown. T.C.Dunn. Vc. 105 Rassal, W. Ross. 25. vi. 1991 P.W.Brown. Vc. 106 Swordale, E. Ross. Il.vii.l909 D. J. Jackson. Vc. 110 Isle of Lewis. 1901 H. McArthur (Mackworth-Praed coll.). A. piercei Obraztsov, 1952 Vc. 73 Balmae, Kirkcud. 18. vi. 1999 J.MacKay. Vc. 74 Stranraer, Wigtown. 22. vi. 1998 R.Meams. Vc. 75 Bennane Head, Ayrshire. 4.vi. 1987 I.C. Christie. Vc. 82 Petersmuir, E.Loth. 28.vi.1935 A.B. Balfour. Vc. 84 Queensferry, W. Loth. pre 1857 R.K.Greville. Vc. 88 Kinloch Rannoch, Perths. v.1927 T.E.D. Poore. Vc. 96 Speybridge, Inverness. 9.vi.I968 E. C.Pelham-Clinton. Vc. 97 Arisaig, Westemess. 2.vi.l941 C .W . Mackworth-Praed. Vc. 101 Tayvallich, Kintyre. 12. vi. 1981 I. C. Christie. Vc. 105 Kishorn, W.Ross. 6. vi. 1 988 P.W.Brown. Vc. 106 Swordale, E. Ross. 20.vi.l908 D. J. Jackson. A. rubigana (Treitschke, 1830) Vc. 72 Parkgate, Dumfries. 9.vii.l982 E. C.Pelham-Clinton. Vc. 73 Kirkdale, Kirkcud. 10.vii.l999 R. & B. Meams. Vc. 79 Glen Kinnon, Selkirks. 31.vii.l999 A.Buckham. Vc. 81 Gordon, Berwicks. 18.vii.l953 E.C.Pelham-Clinton. Vc. 84 Winchburgh, W. Loth. 8.vii.l970 E.C.Pelham-Clinton. Vc. 85 Aberdour, Fife. 27.vii.1969 E.C.Pelham- Clinton. Vc. 89 Kinnaird, Perths. 2/4. vii. 2000 J. A.T.Woodford. Vc. 90 The Scorrie, Glen Clova. 14. vii. 1995 K. P.Bland. Vc.104 Isle of Canna.29.vii. 1971 J.L.Campbell. Vc. 106 Nigg Sutor, E. Ross. 9.vii.l910 D. J.Jackson. A. smeathmanniana (Fabricius, 1781) Vc. 72 Kirkton, Dumfries. 7. vii. 1996 R.Meams. Vc. 73 Merse Head, Kirkcud. 20.vii.l996 J. MacKay. Commophila aeneana (Hubner, 1800) Vc. 92 There are 2 specimens labelled “Aberdeen, 1910 Purdey coll.” In British Museum (Natural History) (Bradley, Tremewan & Smith, 1973). This is a surprising record that has never been repeated. A single specimen was taken on a window in Berwick-on-Tweed (Vc.68) in 1887 (Bolam, 1929) suggesting that it is an occasional vagrant. Eupoecilia angustana (Hubner, 1799) Vc . 72 Kettleton, Dumfries. 13.vii.l978 A.B. Duncan. Vc. 73 Kirkconnell, Kirkcud. 4.viii.l976 A.B. Duncan. Vc. 78 Whim Moss, Peebles. 12.viii.l986 K. P.Bland. Vc. 79 Selkirk. 16. vi. 1943 B.W. Weddell. Vc. 84 Winchburgh, W. Loth. 20.vii.l979 E. C.Pelham-Clinton. Vc. 94 Cullykhan, Banff 22.vii.1993 M.R.Young. Vc. 97 Arisaig Islands. 7.vi.l941 C. W. Mackworth-Praed. Vc.lOl Taynish, Kintyre. 11. vi. 1983 K.P.Bland. Vc.105 Beinn Eighe, W. Ross. 1953 O.W. Richards. Vc.l07 East Sutherland. Date unknown. H.N.Michaelis. E. ambiguella (Wijibner, 1796) Vc. 83 Logan (in Lowe & Logan, 1852) reported a possible specimen of Eupcecilia ambignana Steph. from Duddingston (Vc. 83), but later corrected its identity to E. atricapitana St. (Logan, 6 1853). Wilkinson (1859) presumably overlooked this correction and states that E. ambiguella “has also occurred near Edinburgh”. Cochylidia subroseana (Haworth, 1811) Pre 1 895 Scottish records of EupoeciUa subroseana refer to the richly coloured form of Falseimcaria ruficiliana (see White, 1869; Barrett, 1869; Griffith, 1884; and Bankes, 1893). C. implicitana (Wocke, 1856) Vc. 73 Barlocco Bay, Kirkcud. 6/7. viii. 2003 R.& B. Meams. C. rupicola (Curtis, 1 834) Vc. 73 Abbeybumfoot, Kirkcud. 22.vi. 1996 R.Meams. Falseuncaria ruficiliana (Haworth, 1811) Vc. 73 Hannaston Wood, Kirkcud. 22.vi.1996 K.P.Bland. Vc. 95 Aviemore. 1 2-2 l.vi. 1909 E.R.Bankes (Brit.Mus.(Nat.Hist.) coll.). Vc 96 Aviemore area, Eastemess. 17.vi.l967 E.C.Pelham-Clinton. Vc. 97 Lochailort, Westemess. 4.vi.l941 C . W. Mackworth-Praed. Vc. 101 Ronachan, Kintyre. lO.v.1953 E.C.Pelham-Clinton. Vc. 102 Colonsay. Date unknown. T.C.Dunn. Vc.105 Kishorn, W. Ross. 25.vi.1980 P.W.Brown. Vc. 107 Invershin, Suth. 19.V.1921 F.G. Whittle (Brit.Mus.(Nat.Hist.) coll.). Cochylis atricapitana (Stephens, 1852) Vc. 73 Carsfad, Dairy, Kirkcud. 4.vi.l993 J. MacKay. Vc. 74 Tors, Wigtown. 13. viii. 1983 A.B. Duncan. Vc. 75 Ailsa Craig, Ayrshire. 21.V.1983 I. C. Christie. Vc. 76 Paisley, Renfrews. 12. V. 1983 J. Morgan. Vc. 81 Gordon, Berwicks. 14. vi. 1952 E.C.Pelham-Clinton. Vc. 84 Winchburgh, W. Loth. 4.vi.l964 E.C.Pelham-Clinton. Vc. 89 Aldclune Meadow, Perths. 19.vi.l988 K. P.Bland. Vc. 90 The Strone, Glen Clova. Larvae. 4.viii. 1992 (reared) K.P.Bland. Vc. 95 Aviemore. 23.vi.1908 E.R.Bankes (Brit.Mus. (Nat. Hist.) coll.). Vc. 97 Sanna, Ardnamurchan. 6. viii. 1956 E.C.Pelham-Clinton. Vc. 98 Benderloch, Argylls. 15.vi.l974 E.C.Pelham-Clinton. Vc. 101 Southend, Kintyre. 2. vi. 1985 l.C. Christie. Vc. 103 Carsaig, Mull. 14.vi.l989 l.C. Christie. Vc. 105 Wester Ross. Date unknown. J. M.Chalmers-Hunt. C. nana (Haworth, 1811) Vc. 73 Beeswing. 1980 E.F. Hancock. Vc. 79 Selkirk, l.vii. 1931 B.W.Weddell. Vc. 81 Gordon, E.C.Pelham-Clinton. Berwicks. 30.vi.l951 Vc. 84 Blackburn, E.C.Pelham-Clinton. W. Loth. 9.vi.l970 Vc. 85 Markinch, Fife. l.vi. 1980 E.C.Pelham- Clinton. Vc. 89 Glen Garry, Perths. 22.vi.1975 E.C.Pelham-Clinton. Vc. 97 Lochailort, Westemess. 15.vi.l941 C.W. Mackworth-Praed. Vc. 98 Lephinmore, Argylls. 20.V.1952 E.C.Pelham-Clinton. Vc.lOl Taynish, Kintyre. 14. vi. 1985 l.C. Christie. Vc.105 Kishorn, W. Ross. 26.vi.1985 P.W.Brown. Vc.107 Shin Falls, Suth. 2/3.vi.l990 K.P.Bland. C. pallidana Zeller, 1 847 Vc. 73 Portling, Kirkcud. 7. vi. 1984 A.B. Duncan. C. roseana (Haworth, 1811) G.H.Stainton (1845) reported this species from Airthrey (Vc.86). The record seems most unlikely and H.T. Stainton seems to have realised the error, for the record is under E. subroseana (=Falseuncaria ruficiliana) in the manual (Stainton, 1859). It is hoped that entomologists will in future keep this distribution data up-to-date by publishing, or communicating to the author, any new vice-county records they have or get. ACKNOWLEDGEMENTS The author is very grateful to the many fellow entomologists who have kindly made their records available to him. REFERENCES Bankes, E.R. (1893) EupoeciUa subroseana: a query! Entomologist 's Record 4 (4), 99-100. Barrett, C.G. (1869) Notes on some British species of EupoeciUa. Entomologist’s Monthly Magazine 5, 244-246. Bolam, G. (1929) Lepidoptera of Northumberland and the Eastern Borders. History of the Berwickshire Naturalists ' Club 27, 1 15-142. Bradley, J.D., Tremewan, W.G. & Smith, A. (1973) British Tortricoid Moths. Cochylidae and Tortricidae: Tortricinae. Ray Society, London. viii+ 251 pp. Dandy, J.E. (1969) Watsonian Vice-counties of Great Britain. Ray Society, London.33pp + 2 maps. Griffith, A.F. (1884) Tortrices, &c., in South Wales and Sutherlandshire. Entomologist’s Monthly Magazine 20, 259-260. Logan, R.F. (1853) Additions to the Lepidoptera of Midlothian. The Naturalist (B.R. Morris edit.) 3, 69-71. Lowe, W.H. & Logan, R.F. (1852) The Lepidopterous Insects of Midlothian. The Naturalist (B.R. Morris edit.) 2, 141-149. [Pelham-Clinton, E.C.] (1958) The Annual Exhibition - Records of Exhibits. 26"’ October 1957. Proceedings and Transactions of the South London Entomological and Natural History Society 1957, 36-37. Pelham-Clinton, E.C. (1982) Correction to published records of Phalonidia minimana (Caradja)(Lepidoptera: Cochylidae). Entomologist’s Gazette 121. 7 Shaw, M.R. (1987) Scottish Insect Records. Entomologist's Record 99, 37-38. Stainton, G.H. (1845) Capture of Lepidopterous Insects in Scotland in 1845. The Zoologist 3, 1090- 1091. Stainton, H.T. (1859) A Manual of British Butterflies and Moths. Vol.2. John van Voorst, London, xi + 480pp. Table 1. The known Watsonian vice-county distribution of the species of the subfamily Cochyliinae (Tortricidae) in Scotland. Log Book Species Scottish Vice County 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 PHTHEOCHROA 921 inopiana (Haw.) s s HYSTEROPHORA 924 maculosana (Haw.) A s s s s s s S s a s PHALONIDIA 932 affinitana (Douglas) A 926 maimiana (F.v.R.) A A? s s GYNNIDOMORPHA 930 alismana (Rag.) S s 927 minimana (Carad.) s A 928 pennixtana (D. & S.) s s 929 vectisana (H.&W.) s a COCHYLIMORPHA 935 altemana (Steph.) 936 straminea (Haw.) s a s S S S s S S S A s s s s S s AGAPETA 937 hamana (Linn.) A s s s s s a s s S S A s s s s s s 938 zoegana (Linn.) A A s s s s s A A S s s s s a s AETHES 945 cnicana (Westw.) a S a s S a a s S s S S A S s s a s s 950 francillana (F.) s 941 hartmanniana (Cl.) s s 942 pierce! Obraz. s a A A s S s S A S A s A s s 946 rubigana (Treits.) a a s s a s A s s A A S s S a a s 940 nitilana (Hb.) s 947 smeathmanniana (F.) a a s s S S S s s 939 tesserana (D. & S.) s COMMOPHILA 952 aenaena (Hb.) EUPOECILIA 954 angustana (Hb.) A A s s s S a A s S s S A S S S S s s s 955 ambiguella (Hb.) s? COCHYLIDIA 956 implicitana (Wocke) a s 959 rupicola (Curtis) A s FALSEUNCARIA 960 nificiliana (Haw.) a s s s s s S S S s s S COCHYLIS 966 atricapitana (Steph.) a A A a s A s s A S s s a a s 964 dubitana (Hb.) s S s s s S s s 965 hybridella (Hb.) s s S s 968 nana (Haw.) s a s A A s S A A s s s A s S 967 pallidana Zell. A S s 962 roseana (Haw.) s? White, F. Buchanan (1869) Notes on the Lepidoptera inhabiting Rosshire. Entomologist’s Monthly Magazine 5, 281-285. Wilkinson, S.J. (1859) The British Tortrices. John van Voorst, London, viii + 328pp. Log Book no. Species Scottish Vice County 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 PHTHEOCHROA 921 inopiana (Haw.) s HYSTEROPHORA 924 maculosana (Haw.) s s A A s s s A A S PHALONIDIA 932 affmitana (Douglas) s 926 manniana (F.v.R.) GYNNIDOMORPHA 930 alismana (Rag.) s 927 minimana (Carad.) 928 permixtana (D. & S.) S s s 929 vectisana (H.&W.) COCHYLIMORPHA 935 altemana (Steph.) A? S ' 936 straminea (Haw.) s s A a S s s s S s A S s AGAPETA 937 hamana (Linn.) s a a a A s s s s S S A S S S 938 zoegana (Linn.) AETHES 945 cnicana (Westw.) S s s s A A s a a a S S A A s A s S 950 francillana (F.) 1 941 hartmanniana (Cl.) s s s s s S 1 942 piercei Obraz. s s S A A S s A s s A A S S s 1 946 rubigana (Treits.) s s S s s A A S s I 940 rutilana (Hb.) s 1 947 smeathmanniana (F.) s s S s s s s S 939 tesserana (D. & S.) s s COMMOPHILA 952 aenaena (Hb.) s? EUPOECILIA 954 angustana (Hb.) s S a s s A S s s a s s s A S a S s S s S 955 ambiguella (Hb.) COCHYLIDIA 956 implicitana (Wocke) 959 rapicola (Curtis) S s s FALSEUNCARIA 960 ruficiliana (Haw.) s a A A s S A a s s A s S S s s COCHYLIS 1 966 atricapitana (Steph.) s s a S a a s A A a s S ; 964 dubitana (Hb.) s 1 965 hybridella (Hb.) 968 nana (Haw.) s s s s A A s s A s A a 967 pallidana Zell. ; 962 roseana (Haw.) S or s - Published records quoted in SIRJ. A or a - Additional records not in SIRI, i.e. currently unpublished. Letters in bold capital script indicate specimen of this provenance is present in NMS. 9 Table 2. Watsonian vice-counties for Scotland. Vc Number Vice County Vc Number Vice Coimty 72 Dumfriesshire 93 North Aberdeenshire 73 Kirkcudbrightshire 94 Banffshire 74 Wigtownshire 95 Moray 75 Ayrshire 96 East Inverness-shire (with Naim) 76 Renfrewshire 97 West Inverness-shire 77 Lanarkshire 98 Argyll Main 78 Peebleshire 99 Dumbartonshire 79 Selkirkshire 100 Clyde Isles 80 Roxburghshire 101 Kintyre 81 Berwickshire 102 South Ebudes 82 East Lothian 103 Mid Ebudes _ 83 Midlothian 104 North Ebudes 84 West Lothian 105 West Ross 85 Fife & Kinross 106 East Ross 86 Stirlingshire 107 East Sutherland 87 West Perthshire (with Clackmannan) 108 West Sutherland 88 Mid Perthshire 109 Caithness 89 North Perthshire 110 Outer Hebrides 90 Angus (& Forfar) 111 Orkney Islands 91 Kincardineshire 112 Shetland Islands 92 1 South Aberdeenshire 10 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 1 1-20. A SURVEY OF FERAL GOATS ON THE OA PENINSULA, ISLE OF ISLAY, WESTERN ISLES, SCOTLAND, JULY 2003 Peter B. Copley' and Donald James MacPhee^ ' Biodiversity Conservation Programs, South Australian Department for Environment and Heritage, GPO Box 1 047, Adelaide, S 5001, Australia. E-mail: coplev.peter@saugov.sa.gov.au ^Author to whom enquiries should be directed: Dunlossit Garden Cottage, Port Askaig, Isle of Islay, Argyll, PA46 7RE, United Kingdom; E-mail: macphee-islav@supanet.com ABSTRACT An observational survey of feral goats was conducted on The Oa peninsula of the Isle of Islay, Scotland, between 6"’ and 17“’ July 2003. Approximately 400 (+/- ca30-40) goats were recorded in 45 groups varying in size from 1 to 44 individuals. Most groups were recorded within less than 500 metres of the coast where they regularly found shelter amongst rock-falls and in various caves in the cliffs. White, black and brown were the most frequently recorded coat colours, followed by a combination of white/beige. Horn characteristics of adult billies, evidence of tagged individuals, and accounts of local farmers, suggest that this goat population has arisen from several sources on several occasions. The total number of goats recorded during this survey is similar to a count obtained in 2000, but considerably larger than all previous counts obtained for the area between 1981 and 1998. Despite an annual cull of between 40 and 50 goats being conducted for landholders on the northern half of The Oa since 1997, the population still appears to be increasing. This paper provides a relatively detailed baseline against which future systematic counts of goats on The Oa may be compared. INTRODUCTION Feral goats, Capra hircus, originating from local domestic animals, have been established along the coastal cliffs and adjacent moors and grasslands of the Scottish island of Islay for at least a century, and probably for considerably longer. These goats occur in four discrete areas of the island. The largest sub- population is on The Oa peninsula (locally referred to as “The Oa”) at the island’s southern extremity. The next largest is on the north-eastern portion of the island, east and north-east of Gortantaoid Point and mostly north-west of Bunnahabhain. Another occurs on the Rhinns of Islay - the island’s south-western peninsula. A fourth small sub-population occurs in the Smaull Farm to Sanaigmore (to Ardnave Point) area on the north-western comer of the island. In addition, a population of feral goats occurs on the small island of Texa that lies about 600m off the south coast of Islay and 4km to the east of the Oa peninsula. Following the recent acquisition of Kinnabus Farms to add to it’s other reserve holding of Upper Killeyan Farm on The Oa peninsula, the Royal Society for the Protection of Birds (RSPB) had gained ownership and management responsibility for approximately the south-western half of the peninsula. The Oa Reserve has been established primarily for the conservation of choughs, Pyrrhocorax pyrrhocorax, which have their largest breeding population in Britain here on the Isle of Islay. The survey described below was undertaken to determine the current distribution and relative abundance of feral goats on The Oa, as a basis for detenuining future management requirements and options on this newly enlarged, 1,883 hectare reserve. Information was also sought to determine current distribution and abundance of goats on other parts of Islay and on the nearby island of Texa. METHODS The survey area The Isle of Islay (lat. 55° 45’ 20” N; long. 06° 15’ 20” W) is the most southerly of the Western Hebridean Isles of Scotland and lies approximately 23km west of the Scottish mainland (see Fig. 1). It is a large island, measuring approximately 40 km from north-to-south and 30 km from east-to-west, but is almost dissected, as shown in Fig. 1, along its north-south axis by Loch Gruinart from the north and the wider intmsion of Loch Indaal and Laggan Bay from the south. Islay experiences an Atlantic seaboard climate described as “hyperoceanic, humid, temperate - 01 H3 Tl” (see MacKay, 1996). Figure 2 plots mean monthly rainfall and mean monthly maximum temperatures, as recorded at the meteorological recording station on Islay between 1983 and 1987 - the recording station closed at the end of 1987. Mean annual rainfall for the period was 1,043 mm (range 975-1 103mm) over the five-year period. The months from September - January tend to receive more rain than the other months, but the pattern is very variable. Temperatures are lowest between November and March, although the pattern for this is also variable. Sea fog often obscures visibility at altitudes above about 100m even during summer months (including on one day during this survey). The high sea-cliffs rising l-200m above sea level add a significant level of wind-chill to minimum temperatures. Winter gales are common, with strong winds predominantly from the SW quarter. 11 The Oa peninsula is the oval-shaped, southern-most promontory of the Isle of Islay and is situated immediately to the west and south-west of the town location relative to the Scottish mainland of Port Ellen (Fig. 3). It measures nearly 8km east-to- west, 8.5km north-to-south, covers approximately 47 km^, and has a (‘smoothed’) coastal circumference of ca 26km. The Oa is mostly a gently undulating, hilly landscape with the highest point, Beinn Mhor, rising to 202m. It is covered mostly by heather moorland and improved and semi-improved pasture, and fringed by coastal cliffs, steep grassy slopes and numerous rocky coves (see Ordnance Survey “Pathfinder” map 439 (NR 24/34/44) “Port Ellen” for details). The Oa reserve consists of approximately 800ha of dry heath / acid grassland (42%), 650ha of coastal grassland / heath (35%), 350ha blanket bog (19%), 65ha of improved grassland (3%), 5ha of arable land and 13ha of standing water (RSPB, unpublished report 2002; but see also Madders, et al. 1998). Scattered around the coastline are many caves of varying dimensions, mostly former sea-caves with entrances at, or near, the cliff-base and indicative of former relatively higher mean sea levels. The Oa reserve has been used for grazing of mixed, traditional suckler-type cows and of Highland black- faced sheep. Such grazing will be retained by RSPB to maintain and improve the grassy pastures as feeding habitats for chough (see Madders, et al. 1998; Finney and Jardine 2003) and other farmland birds of conservation concern. However, numbers of both sheep and cattle have been reduced significantly on The Oa in recent years and future management may require changes in grazing regimes (grazing species, density, frequency, timing) in key management zones to achieve particular conservation outcomes. The role that grazing and browsing by feral goats may play in this is yet to be determined. Survey Methods The coastal perimeter of The Oa peninsula was surveyed on foot, searching for individuals or groups of feral goats both on the cliff-slopes and inland. High vantage points with clear views over large areas were sought first to note the positions of distant groups and how best to approach these. Visibility was generally very good for locating goats, due to the vast areas of very low vegetation and the starkly contrasting colours of the goats’ pelage. The significant exceptions to the good visibility were numerous, mostly fairly small, areas obscured by topographic features such as ridges and gullies. These were searched fairly systematically, although some small areas were inevitably missed. Also, a few areas of tall bracken were searched carefully as small numbers of goats were occasionally hidden within them. One of us (DJM) is familiar with these areas of cover often used by goats on The Oa, and has regularly stalked and culled small numbers of goats there for the local farmers and for occasional trophy hunters. When goats were located, an initial count was obtained. Then group composition, based on the numbers of adult males (billies), adult females (nannies) and kids was recorded, with ‘yearling’ and younger kids separated where possible. Where distant groups disappeared from sight before all group characteristics could be recorded, total counts were obtained as a priority and then numbers of ‘obvious’ adult males were noted. The composition of a small number of groups was therefore insufficiently known and a minimum number of adult males were recorded. These groups are marked with an asterisk in the summary table (Appendix 1). In these situations all other goats were recorded as “adult females +/- kids”. Consequently, the numbers of adult males is likely to be a slight under-estimate and the numbers of adult females +/- kids a slight over-estimate. An attempt was also made to identify particular groups based on coat colours and coat patterns, and on the size and shape of the horns of particular adult males. No attempt was made to classify the age of individual goats based on horn ring counts (after Bullock and Pickering 1984). This would have 12 18 16 14 12 10 8 6 4 2 0 E Month Figure 2. Climatic conditions. Isle of Islay. required far longer than time permitted for this general overview survey. Coat colours recorded were simplified to black (B), white (W), brown (Br) and beige (Be), and predominant combinations of these (e.g. B/Br, BAV, etc.). Although scoring of these colours and combinations was, at least in part, subjective, it nevertheless provided a useful cue for recognition of particular groups. The whole survey was conducted from 6* to 1 7''’ July 2003. One or two people with binoculars and a telescope, and two to four other observers, walked close together at all times, except when goats needed to be ‘flushed’ from cover back in the direction of the main recorder(s). The weather was clear on all but one foggy day when only a short stretch of coastline around Glen Astle was searched. Wherever possible, searching was carried out into the wind, so that goats could be approached as closely as possible. On the evening of 16 July, four people conducted a survey of the feral goats on the uninhabited isle of Texa. The goats have been present on this low, hillocky island of approximately 60ha for many decades. The numerous hillocks and, in some places, tall bracken, obscured visibility of several areas of the island, especially in the north. But the hillocks also provided excellent vantage points for observing goats once they were located and ‘driven’ into open areas. A fmd-flush-and-count method was therefore used for this survey. Approximate numbers of goats present in other areas of Islay were obtained from knowledgeable local residents, to provide as complete a view of feral goat numbers and distribution across the island contemporaneous with that obtained in this more detailed survey on The Oa. RESULTS The locations, relative sizes and composition of groups of feral goats encountered during this survey on The Oa are shown in Figure 4. From this it is clear that the goats occur predominantly along the coast and for only short distances (up to 1km) inland. It is also evident from this plot that the groups are clumped in particular areas. Minimum numbers of goats observed, by sex and by relative age (young kids up to about 4 months old, ‘one-year-old’ kids actually about 6-8 months old, and adult females and adult males) are summarised per group recorded in Appendix 1 . A total of 432 goats were recorded in 46 groups, although one of these groups of 12 individuals was definitely seen and scored twice. Thus a maximum of 420 goats were observed in 45 groups. These groups included: • at least 1 66 billies within 27 groups, • a maximum of 204 nannies within 34 groups, • at least 49 “year-old” kids within 28 groups • only three young kids within three separate groups. 13 Group size (see Fig. 5) ranged from one to 44 individuals (mean 9.8 goats; n=45). Twenty-three percent of groups were all-male groups and ranged in size from three to 11 individuals (mean 6.0; n=10). Thirty-five percent of groups were female +/- kid groups and ranged in size from one to 23 individuals (mean 5.1; n=16). Forty-two percent of groups were mixed sex groups, ranging in size from five to 44 individuals (mean 15.8; n=19). Interpretation of data The number of adult females recorded was greater than for adult males (0.55:0.45), although there was almost certainly an over-count of females and a corresponding under-count of males, due to the rapidity with which three groups disappeared from sight and the consequent method used to record group composition. When these groups are removed from this part of the analysis, the sex ratio becomes 0.48:0.52. As the counts in the present survey were performed over several days and there appeared to be some fluidity between groups (e.g. some groups divided or amalgamated as they grazed or as they returned to their shelter sites late in the day), the count data presented are likely to include some ‘double- counting’ of individuals. This is most likely to have occurred within the area between Dun Athad, just east of The Mull of Oa, and Stremnishmore, because different counts were made along partially- overlapping areas of this section on at least three separate days, over an eight day period. However, it is thought that this is unlikely to represent more than about 30 to 40 individuals of the 60 counted in this area on 7* July (see Appendix 1). In addition, a group of 1 2 is also known to have been counted twice between The Mull of Oa and Lower Killeyan (as already mentioned), but this has been accounted for in the total. Any double-counting is also likely to be countered by the possible lack of sighting of a group of 40-60 goats that are considered by one of us (DIM), and by Hamish McTaggart the owner of Kintra Farm (pers. comm.), to be ‘resident’ on Kintra Farm. On the day that Kintra Farm was surveyed, and also on the day before, the property had been quartered on quad bikes to find and muster all sheep for their annual wool clip. As a consequence of this, the local group(s) of goats may have moved elsewhere, and only eight goats were seen on this property despite considerable search effort. Numbers seen immediately to the south of here were not particularly high, and were not higher than might have been expected from local accounts. The coat colours and patterns of the goats were many and varied and careful noting of these would greatly enhance future identification of individuals, or at least some individuals within particular groups or areas. While the scoring of coat colours in this survey was simplified to either predominant colour or a combination of the two predominant colours, this was still a useful practice for attempting to distinguish groups, especially when combined with each animal’s sex. The relative frequencies of predominant coat colours by sex are summarised in Fig. 6. White is the most frequent coat colour among both male and female goats on The Oa. Black coat colour and brown were next most frequently recorded, followed by the combinations of white/beige and black/brown. A few individuals also had very distinctive coat patterns such as square patches or circular ‘bulls- eyes’ that allowed them to be identified immediately. Mature male goats on The Oa exhibit a variety of horn shapes. Many have the relatively simple, backward-curving, scimitar-type of horn. Others have an additional outwards curve, while still others curve, or flair, outwards at least twice, with some spiralling of the horns. These characteristics add considerably to the observer’s ability to identify individual animals. b) Comparison of counts of feral goats on The Oa with other areas on Islay Table 1 below summarises counts, or recent estimates, of feral goat numbers on other parts of Islay, and on the isle of Texa. It also compares the current survey counts with those of previous counts conducted on The Oa. While the current survey is almost certainly the most thorough undertaken on The Oa, it is worth noting the much higher numbers recorded in this area over the last decade. Also of note are the lower counts on The Oa in 1985, indicating either a population decline following the earlier 1980s counts, or the difficulty of locating all goats along this complex coastal landscape. (c ) The Isle of Texa A total of 84 goats were counted on the isle of Texa during this survey. They were all in a single large tribe consisting of 41 adult males, 32 adult females and 1 1 kids. DISCUSSION a) The Oa Peninsula Population size and group composition In July 2003 there were approximately 400 (+/- ca 30- 40) feral goats living on The Oa peninsula of Islay. This total is very similar to that obtained by Angus Keys (local reserves manager, RSPB, personal communication) during a seabird census around the coastline of The Oa (from a boat) in 2000 (see Table 1). Approximately even numbers of adult males and females were recorded. However, it should be noted that considerably more adult males than adult females (ca 170 cf 80) have been culled from the population between 1997 and 2003 (DJM, unpublished data). Goats have been culled on The Oa each year since 1994. Initially only very small numbers were removed from the population. However, with the exception of 1998 (when 10 goats were culled). 14 between 40 and 50 goats have been culled there annually since 1997. These culls have usually consisted of four to six adult billies and about 35 to 40 adult nannies and kids. Fifty-two first-year kids were identified in the count, representing at least 12% of the counted population, although this may also be an under-estimate because many were of a comparable size with adult nannies. A small number may therefore have been classed as adult nannies when seen only at briefly. A total of 45 groups of feral goats were counted, ranging in size from one to 44 individuals. The mean group size of 9.8 animals is considerably larger than the mean group size of about 5 animals in July recorded on nearby Rum (Shi, et al. (b) in press). However, this latter study involved a much larger sample size over a far greater time span. The study on Rum (Shi, et al. (a) 2003 and (b) in press) showed that group size varies with time of day, generally decreasing as the goats divided into smaller feeding and social groups as they move away from their night caves and increasing again as the goats return to their night shelters. Most groups observed during the survey of The Oa, were recorded between the hours of 10:30 and 18:00. It is therefore likely that fewer, but larger, groups might have been encountered if the survey was undertaken earlier in the morning and later in the afternoon. Group size and composition also vary throughout the year on Rum, with the percentage of mixed-sex groups increasing sharply during the rut in August and September, and being at its lowest from April to July (Shi, et al., (b) op. cit.). Outside of the rut, the frequency of female groups is usually greater than male groups and mixed-sex groups. The relatively high proportion of mixed-sex groups recorded during this survey on The Oa during late July may be indicative of the approaching rut. Distribution on The Oa Most goats on The Oa were observed on, or near the coastal cliffs, and mostly less than 500m inland. Several local residents observed that goats headed for the coastal cliffs at the onset of bad weather. Also, during this survey, most goat groups that were disturbed by our presence, usually headed for the coastal cliffs and rocky bays that were within close proximity. Closer examination of the coast revealed that these favoured areas have substantial rock overhangs and former sea caves (from periods of higher relative sea levels) in which the goats clearly seek regular refuge. All shelters had deep deposits of goat droppings on their floors and strong odours of goats prevailed. Impressive examples of these shelters occur in the two bays to the east of Dun Athad and on the south side of the bay at Glen Astle. The goats also shelter amongst and behind some of the larger rockfalls on cliff-slopes and at the cliff- bases. Coat and Horn Characteristics The diversity of coat colours and patterns, and adult male horn shapes, observed in The Oa goat population suggest that the population has arisen from several sources. Local residents reported that domestic goats were likely to have escaped or been released at various times over the past 100 or more years. Certainly, there appear to be some goats with ‘ancestral’ features, including billies with simple, long, scimitar-shaped horns, that may have arisen from older “Scottish-type” stock (see Whitehead 1972). Others have features, such as wider-sweeping, spiralled horns more characteristic of the more recently introduced angora-type goats. While these latter goat traits may have been introduced to The Oa population on more than one occasion, and over a longer period than just two or three decades, three local residents reported that ‘angora-type’ goats had escaped from a goat farm near Kilnaughton Bay, on the north-eastern portion of The Oa, over the past 20 years or so, and that several had also been deliberately released near The Mull of Oa in about 1995. These accounts are supported by the observation of at least two goats with orange ear-tags seen annually between 1999 and 2002 by RSPB reserve manager, Angus Keys (pers. comm.). Population Trends While the data presented here are insufficient to examine population growth trends, there is circumstantial evidence that The Oa goat population is increasing. Firstly, the highest counts (and by a substantial margin) have been recorded in recent years (see Table 1). Secondly, several local residents made the observation that goats are now seen further inland than previously - up to as much as 2km inland in the summer months - and that they have even been seen occasionally on the western outskirts of Port Ellen in the last two to three years. Farmers on The Oa also claim that more goats are being seen than ever before. Interestingly, this is in spite of the numbers of goats that have been culled on farms on both sides of the peninsula, each year since 1 994. Dunbar, Shi, Buckland and Miller (unpublished observations) and Dunbar, Buckland, Miller and Coldbeck (unpublished observations) have shown that the feral goat population of the Isle of Rum - just 120 km north of Islay - appears to be controlled largely by extreme climatic conditions and that population growth is limited mostly by the amount of shelter available in caves and rock shelters in the coastal cliffs. Dunbar, Shi, Buckland and Miller (unpublished observations) have examined climatic influences on the activity budgets of the feral goats on the Isle of Rum and the consequent implications of this for population dynamics under climate change. From this work, they concluded that a mean monthly windchill-adjusted temperature of 5^C appeared to be a critical threshold in terms of the goats’ ability to survive and that caves and other sheltered sites were 15 an essential component of the animals’ survival strategy under such conditions because they effeetively raised ambient temperatures for the goats. Climatic conditions on The Oa are generally less severe than on the Isle of Rum, due primarily to The Oa’s lower altitude (200m compared with 400m). Mean monthly rainfalls are also considerably less on Islay (about a third less in most months) and mean monthly temperatures are about 1-2°C warmer. It therefore seems likely that the climatic limitations on the goat population on The Oa may be significantly less than on the population on the Isle of Rum and that the population on The Oa may be increasing. The feral goat population on The Oa may therefore provide an interesting basis for a comparative study of population growth rates and controlling influences when contrasted with population data from the Isle of Rum, or elsewhere. Future management of the feral goat population on The Oa peninsula Madders, et al. (1998) have shown that choughs foraging on The Oa peninsula select grassland habitats to feed in. They showed that in December- February choughs forage preferentially in acidic grasslands and improved grasslands of inland areas. However, in July-September, when the choughs are distributed mostly around the coastline (and have nests mainly in coastal caves). Madders, et al. (1998) showed that they forage preferentially in neutral (dune) grasslands and on cliffs and slopes. Bullock, et al. (1983) have demonstrated the importance of grazing herbivores to chough feeding ecology, through maintaining the grass sward at a height short enough to allow choughs access to soil invertebrates, while dung beetles (Aphodius) found in the faeces of cattle and sheep (and presumably goats) represent an important additional source of prey (Madders, et al. 1998). Shorter swards, as well as areas of bare soil, bare rock and dung, also benefit other thermophilic invertebrates, such as the yellow mound ant {Lasius flaviis), which the choughs can feed on D. Beaumont, RSPB reserves ecologist, pers. comm.). Feral goats are now the major vertebrate herbivores with access to the coastal cliffs and the vegetation adjacent to eliff-tops on The Oa peninsula on Islay. Domestic livestock (sheep and cattle) numbers have been reduced significantly on the peninsula in recent years and their access to the dangerous cliffs is now either excluded (in several areas), or is managed carefully to minimise the risk of livestock losses through misadventure. Browsing and grazing by the goats is therefore the main way that the short sward and floristically diverse vegetation of the cliffs and cliff-top pastures may be maintained, and also potentially expanded in future. In the absence of such ongoing grazing and browsing it is likely that such areas may become colonised by taller, denser, woody perennial shrubs sueh as heather, willow and bracken. In addition, the likely high production and at least oceasional mortality of feral goat kids will provide prey and carrion for birds such as golden eagle and raven. Each of these considerations needs to be weighed against other possible conservation outcomes, the potential for degradation of some areas through impacts of too many goats should their population continue to expand, and the affects that annual culling of goats on The Oa peninsula may have on a range of other land management objectives. h) Other Areas By comparison with the population on The Oa, feral goat numbers in the north-east, north-west and south- west of the island appear to be considerably smaller. There is insufficient evidence on which to base any conclusion about population trends in these areas. However, more recent figures for both the north-west and the north-east of the island are higher than previous counts or estimates. Closer assessment is needed. The goat population on the small isle of Texa is interesting for two reasons. Firstly, at the time the survey reported here was undertaken, all 84 goats counted were in a single tribe on the north-western comer of the island. Secondly, these goats apparently supplement their diet with kelp (DJM personal observation). The relatively larger proportion of adult males to adult females in this population (41:32) may, at least in part, be attributed to some recent population management through culling of some adult females. As with The Oa population, at least one female goat in the Texa population was noted to have a red ear- tag, suggesting that someone has introduced goats here in the recent past (or, less likely, that someone has been catching and tagging goats on the island). CONCLUSIONS The present study has raised more questions than it has answered. The feral goat population on The Oa appears to be increasing in both abundance and range, though this will need to be monitored, using this summary paper as a base-line. There appear to be some significant similarities and some significant differences between the goat populations on The Oa and on the Isle of Rum. The Oa goat population may therefore provide an interesting and very instructive comparison with the data obtained on Rum - if researchers choose to establish a research program on The Oa. It is likely that the goat population on The Oa will require ongoing management for both farming and conservation purposes. Any population research that follows this initial baseline survey should help to define what that management should be and where and when it should be applied to best effect. 16 ACKNOWLEDGEMENTS We wish to acknowledge Dave Beaumont (RSPB reserves ecologist) for suggesting this project, and to thank Dave and the local RSPB staff - John McGhie and Gus Keys, in particular - for their support and assistance. We also wish to thank Alistair Carmichael of Coillabus farm, Donald Sinclair (senior) of Lower Coillabus farm, Archie Carmichael of Kinnabus Cottage, Ian Carmichael of Fang Dhu and Hamish McTaggart of Kintra farm, for telling us about their observations of the local goats on The Oa. Malcolm Ogilvie kindly provided information on earlier counts of goats on Islay. Estimates of the numbers of goats in the north-east of the island were provided by Jack Adamson, gamekeeper at Islay House. Professor Robin Dunbar of Liverpool University kindly provided background information about the research work conducted on the feral goats on the Isle of Rum. The maps for this paper were kindly produced by Neil Cowie (RSPB reserves ecologist). Anna Copley, Pippa Copley and Rhona Macphee provided valuable field assistance during the survey work. REFERENCES Bullock, D.J. and Pickering, S.P. (1984). The validity of horn ring counts to determine the age of Scottish feral goats {Capra (domestic)). Journal of Zoology, London 202, 561- 564. Bullock, I. D., Drewett, D. R. and Mickleburgh, S. P. (1983). The chough in Britain and Ireland. British Birds 76,377-401. Finney, S. K. and Jardine, D. C. (2003). The distribution and status of the Red-billed Chough in Scotland in 2002. Scottish Birds 24, 11-17. MacKay, C. R. (1996). Conservation and ecology of Red- billed Chough Pyrrhocorax pyrrhocorax. Unpublished PhD thesis. University of Glasgow. Madders, M., Leckie, F. M., Watson, J. and McKay, C. R. (1998). Distribution and foraging habitat preferences of choughs on The Oa peninsula, Islay. Scottish Birds 19, 282-289. Newton, S. (ed.) (1984). Observations of birds and mammals. Birds of Islay, 1981-83. Brathay Field Studies Report, no. 40. Shi, J., Dunbar, R., Buckland, D and Miller, D. (2003). Daytime activity budgets in feral goats {Capra hircus) on the Isle of Rum: influence of season, age and sex. Canadian Journal of Zoology, 81, 803-815. Shi, J., Dunbar, R., Buckland, D and Miller, D. (in press). Dynamics of grouping patterns and social segregation in feral goats {Capra hircus) on the Isle of Rum, NW Scotland. Mammalia. 17 Figure 3. The Oa peninsula, Isle of Islay, Scotland, showing place names referred to and indicating the distributon and relative size and composition of feral goat groups observed during July 2003. I Reproduced from Ordinance Survey 1:50,000 scale map with permission of the Crown Controller of Her Majesty ‘s Stationary Office. Crown Copyright. Licence AL815519. 18 Figiure 4. Feral goat group size (numbers of goats). The Oa, Isle of Islay - July 2003. Figure 5. Feral goat colour frequencies. The Oa. Isle of Islay - July 2003. Goat coat colour 19 Table 1. Comparison of counts of feral goats made on the Scottish Isle of Islay since 1981. Location Date Count / Estimate Comment The Oa peninsula April 1981 123 Malcolm Ogilvie, (pers. comm.) Aug. 1983 101 Malcolm Ogilvie, (pers. comm.) 1984 100-150 Nevvton (1984) Aug. 1985 59 Malcolm Ogilvie, (pers. comm.); island-wide survey 1998 ca 300 Donald James MacPhee during island-wide deer census by helicopter 2000 ca370 A. Keys, (pers. comm.) July 2003 ca 400 This survey NE Islay Aug. 1985 163 Malcolm Ogilvie, (pers. comm.); island-wide survey July 2003 ca 180 Donald James MacPhee NW Islay Aug 1985 32 Malcolm Ogilvie, (pers. comm.); island-wide survey July 2003 ca 40 Donald James MacPhee SW Islay July 2003 ca 70 Donald James MacPhee 20 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 21-23. HYDROPORUS SCALESIANUS (COLEOPTERA, DYTISCIDAE) NEW FOR SCOTLAND G.N.Foster SAC Research & Development Division, Auchincruive, Ayr KA6 5HW, U.K. ABSTRACT Hydropoms scalesianus Stephens, 1828 is a species typically associated with relict fens. It ranges from Ireland, and Les Landes in south-west France to northern Italy, the Czech Republic, and much of Denmark, Fennoscandia, and has recently been recorded as far as West Siberia. This paper describes for the first time the species presence in Scotland. Adults were were fairly common in Fonah Bog (NO 5350), just west of Balgavies Loch in Angus. The beetles were commonest in a mat of Calliergon moss lying over a floating bed of bogbean {Menyanthes trifoliata L.), bog myrtle (Myrica gale L.) and Sphagnum in a shallow, hard- bottomed depression on alluvial deposits. The habitat together with its flora and fauna, including other species of water beetle, is described. The literature on records from outside Scotland is briefly reviewed. RESULTS Hydroporus scalesianus Stephens, 1828 is a species typically associated with relict fens. It ranges from Ireland, and Les Landes in south-west France to northern Italy, the Czech Republic, and much of Denmark, Fennoscandia, and Russia (Nilsson & Holmen, 1995), recently as far as West Siberia (Petrov, 2002). In Ireland it is still relatively frequent in lake fens and peat bogs from Tipperary to County Down. Fragments of H. scalesianus have frequently been identified in peat deposits because no other Hydroporus species is so small and red. Even though this species has a largely northern distribution it has never been reported in Scotland previously. Adults were fairly frequent in Fonah Bog (NO 5350) on 4 April 2003. The site lies just west of Balgavies Loch in Angus. The beetles were commonest in a mat of Calliergon moss lying over a floating bed of bogbean {Menyanthes trifoliata L.), bog myrtle {Myrica gale L.) and Sphagnum in a shallow, hard-bottomed depression on alluvial deposits. The floating carpet ran into a willow carr on one side and beaked sedge {Carex rostrata Stokes) on the other. A few specimens could be found in the extreme edge of sedge fen. The fauna was dominated by water hoglice or slaters, Asellus aquaticus (L.), and 19 other species of water beetle were found near to but not necessarily in immediate association with H. scalesianus. These were Haliplus ruficollis (De Geer), Hygrotus inaequalis (Fab.), Hydroporus angustatus Sturm, H. erythrocephalus (L.), H. palustris (L.), H. striola (Gyllenhal in Sahlberg), H. umbrosus (Gyllenhal), Agabus bipustulatus (L.), A. congener (Thunberg), A. unguicularis (Thomson), Ilybius ater (De Geer), Rhantus exsoletiis (Forster), Anacaena lutescens (Stephens), Enochrus coarctatus (Gredler), E. ochropterus (Marsham), Hydrobius fuscipes (L.), and Cercyon tristis (llliger). Of special interest was a single male of Acilius canaliculatus (Nicolai) a species previously known from Angus in Restenneth Moss and Fithie Loch, taken by Professor Frank Balfour-Browne in 1908 and 1947 respectively. Several individuals of Hydrochus brevis (Herbst) were found in the sedge fen margin, a new record for Angus. H. scalesianus has Red Data Book (RDB) 2 status in Britain (Shirt 1987), H. brevis having RDB 3 status, and A. canaliculatus provisional RDB 3 status (Hyman & Parsons, 1992). H. scalesianus was originally named in honour of a Mr Scales of Beechamwell, West Norfolk, but it was not until 1977 that the species was rediscovered in that area, in richly vegetated ponds on the Brecks (Foster, 1982). The species was especially well known from Askham Bog, mid- West Yorkshire (Balfour-Browne, 1940) from 1857 until the end of the 19"’ Century. Balfour- Browne (1940) found H. scalesianus infrequently in the Norfolk Broads during his 1904-6 survey, and it was found spasmodically at Sutton Broad (1926) and Catfield Fen (1923 and 1932) until the 1970s, since when it has become more widespread in the fenland surrounding the Broads (personal observation). Balfour-Browne (1940) discussed three further English records for H. scalesianus, from Hebden Bridge, south-west Yorkshire ca 1830, from Boxmoor, Hertfordshire and from the Portsea area of South Hants up to 1880. He thought it unlikely that these records were due to wrong identification, and he has been proved right for Boxmoor, E.G. Elliman’s material in the National Museum of Wales, Cardiff including a specimen from Boxmoor taken in the summer of 1901 (Foster, 1990). The Portsea record has greater credibility following Jeff Robinson’s discovery of a site at Sandford Bridge, Dorset in 1997, and, of course, the Hebden Bridge record is supported by the beetle’s former occurrence at Askham Bog, and the beetle’s recent rediscovery in Yorkshire, beside Hornsea Mere, by Hammond (2002). Horsfield and Foster (1982) reported H. scalesianus in a small, peat-filled kettlehole in County Durham in 1978. Bilton (1984) discovered H. scalesianus in Cumberland, at Biglands Bogs, in 1983. Bilton (1988) was also the first to detect it in Ireland, in 1986. The post-glacial subfossil records are concentrated in the Somerset Levels (Girling, 1984), often in sufficient numbers through peat monoliths to indicate survival over the whole transition from lake fen to raised bog. These records also range 21 from the Neolithic (5170 before present - B.P.) to post-iron Age (ca 1700 B.P.), and records continue to accumulate from other post-Glacial deposits, most notably Lindow Man in Cheshire (Dr M H Dinnin, pers. comm. - see also the BUGS2000 data-base - Buckland et al. 2002). DISCUSSION This compilation of records is a mixture of relict status and discovery, the latter possibly even indicating a recent extension of range. H. scalesiamis is amongst a group of fenland insects largely confined to relict fen and unknown to colonise habitats of man-made origin. At one stage (Foster, 1982) the British distribution suggested association with relict ponds in the periglacial zone of the most recent Ice Age, but this possibility has been disposed of most effectively by the Cumbrian and Scottish finds. As a member of the relict fen group, H. scalesiamis might be expected to be flightless. One specimen has been detected in a flight trap in south-east Sweden (Lundkvist, Landin and Karlsson, 2002), so recent occupancy of relict sites retaining high quality habitat cannot be ruled out. However, the most likely scenario, given the species’ association with relict sites and other species considered to have relict status, is that we are simply accumulating knowledge of a fragmented distribution by deploying a relatively small number of observers. An interesting feature of the present day distribution, as opposed to what is known from the fossil record, is that most modem British sites are a relatively short distance from the sea, suggesting increased dependency on a mild coastal climate. The continued wider distribution within Ireland might lend support to this idea, but it is negated by the loss of the species from fens around the Severn estuary, and, of course, by what is known of the extensive distribution in Continental Europe. ACKNOWLEDGEMENTS I am grateful to Dr J A T Woodford for arranging access to the site, which is part of the Scottish Wildlife Trust Reserve of Balgavies Loch. Mr Brian Nelson updated my knowledge of the status of H. scalesiamis in Ireland. SAC receives financial support from the Scottish Executive Environment & Rural Affairs Department. REFERENCES Balfour-Browne, F. (1940). British Water Beetles. Volume 1. London, Ray Society. Bilton, D.T. (1984). Four water beetles (Col., Dytiscidae) new to Cumberland, including Hydroporus scalesianus Stephens. Entomologist's Monthly Magazine 120, 251. Bilton, D.T. (1988). A survey of aquatic Coleoptera in central Ireland and the Burren. Bulletin of the Irish Biogeographical Society 11, 77-94. Buckland, P.C., Buckland, P.I., YuanZhuo, D. & Sadler, J. (2002). Bugs Coleopteran Ecology Package [CD- ROM]. Foster, G.N. (1982). Notes on rare Dytiscidae (Coleoptera) in Norfolk. Transactions of the Norfolk and Norwich Naturalists' Society 26, 3-10. Foster, G.N. (1990). Hydroporus scalesianus in the British Home Counties in 1901. Balfour-Browne Club Newsletter A6, 13. Girling, M.A. (1984). Aquatic Coleoptera in the fossil insect assemblages from archaeological sites in the Somerset Levels. Balfour-Browne Club Newsletter 30, 1- 11. Hammond, M. (2002). A preliminary survey of the water beetles of Hornsea Mere, East Yorkshire (England), including the rediscovery of Hydroporus scalesianus in Yorkshire. Latissimus 15, 1-3. Horsfield, D. & Foster, G.N. (1983). Hydroporus scalesianus Stephens and Laccornis oblongus (Stephens) (Col., Dytiscidae) in Hart Bog, County Durham. Entomologist's Monthly Magazine 119, 62. Hyman, P.S. & Parsons, M.S. (1992). A review of the scarce and threatened Coleoptera of Great Britain, Part I. Peterborough, Joint Nature Conservation Committee. (UK Nature Conservation, No. 3.) Lundkvist, E, Landin, J & Karlsson, F. (2002). Dispersing diving beetles (Dytiscidae) in agricultural and urban landscapes in south-eastern Sweden. Annales Zoologici Fennici 39, 109-123. Nilsson, A.N. & Holmen, M. (1995). The aquatic Adephaga (Coleoptera) of Fennoscandia and Denmark. II. Dytiscidae. Fauna Entomologica Scandinavica 32, Leiden, E.J. Brill. Petrov, P.N. (2002). The aquatic Adephaga beetles (Coleoptera) of southern Tyumen Region. Bulletin of Moscow Society of Naturalist, Biological Series 107 (3), 31-38 [in Russian]. Shirt, D.B. (Ed) (1987). British Red Data Books: 2. Insects. Peterborough, Nature Conservancy Council. 22 Fig. 1. Distribution of Hydroporiis scalesianus in Britain and Ireland. Large circles represent 10 km squares in which the species has been found from 1980 onwards, smaller circles earlier records of living beetles, and the crosses post-glacial fossil fragments, with the most recent records taking priority 23 24 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 25-28. THE EFFECT OF WATER CONTENT AND COMPACTNESS OF SOIL ON THE SURVIVAL OF THE NEW ZEALAND FLATWORM ARTHVRDENDYVS TRIANGULATES Peter H. Gibson and Derek J. Cosens Institute of Cell, Animal and Population Biology, University of Edinburgh, Ashworth Laboratories, West Mains Road, Edinburgh EH9 3JT ABSTRACT In an Edinburgh allotment from February to June the soil immediately beneath surface debris where the flatworm Arthurdendyus triangulatus was found contained 30% water. When specimens of A. triangulatus were allowed to desiccate at 20°C in the laboratory, 50% survived a body water loss of 23% by weight. Specimens, when presented with a choice of soils with differing water contents in a vermarium, chose soil containing 30% water by weight. In a Petri dish study at 10°C, 50% of specimens survived on soil with 16% water, while 90% survived on soil with 27% water. In another vermarium study A. triangulatus chose loose soil over compacted soil. The water content of the soil is argued to be one of the key factors in determining the distribution of A. triangulatus in Great Britain. INTRODUCTION The New Zealand flatworm Arthurdendyus triangulatus (Dendy) (formerly Artioposthia triangulata, see Jones and Gerard, 1999) is a predator of earthworms and has become endemic to Scotland after being accidentally introduced some 37 years ago (Boag, Yeates, Johns, Neilson, Palmer and Legg, 1995). The general biology of A. triangulatus was reviewed by Cannon, Baker, Taylor and Moore (1999), and Jones and Boag (2001) have given an historical account. It is typically found on the soil surface under items of debris such as wooden planks and plastic bags filled with compost, but principally occupies earthworm burrows within the soil. The under-surfaces of long-standing debris have the environmental characteristics of burrows and are a ready source of earthworms (Lillico, Cosens and Gibson, 1996; Gibson and Cosens, 2000a, b). The debris and burrows, in effect, act as refuges for^^. triangulatus (and other invertebrates) against desiccation to which it is very vulnerable. This paper examines the role of soil water content on the survival and distribution of A. triangulatus. GENERAL PROCEDURES Specimens of A. triangulatus and soil samples were collected from Midmar Drive allotment in Edinburgh, Scotland (Ordinance Survey sheet 66, grid reference NT 252707). The soil in the allotment was a loam derived from a glacial till consisting of a reddish sandy silt-clay of carboniferous sandstone origin containing a proportion of organic material added over the years by gardeners. Specimens of A. triangulatus of 0.5- 1.0 g body weight were collected from under debris and stored in the laboratory on fresh compacted allotment soil in 90 mm diameter soda glass Petri dishes at 1 0°C in a refrigerator. The soil used in the dishes was sieved through a 3 mm mesh. Death of the specimens resulting from experiments was indicated by a lack of movement when they were placed in cold tap water. Living flatworms typically revived immediately. METHODS AND RESULTS What is the normal water content of the allotment soil? From February 1st to June 26th soil samples were taken to a depth of 100 mm from under items of debris (plastic sheeting, wooden planks and stones) lying on the soil surface of the allotment. These samples were compared with 10 soil samples taken nearby from directly beneath turf To establish their water content (0g) by gravimetric analysis, each was placed in a plastic bag which was immediately sealed. In the laboratory 150 g of each sample was dried in an oven at 100°C for 24 h, allowed to cool over silica gel in a desiccator and reweighed. (The dried soil was stored in the desiccator for use in other experiments). The mean water content (0g) of 26 samples of soil collected from under debris was 30.1% (SD = 10.0, Fig. 1) and that for 10 soil samples collected from under turf was 22.0% (SD = 8.1). Fig. 1. The mean percentage water content by weight of soil samples taken from beneath debris in Midmar Drive allotment, Edinburgh, during the spring and early summer (error bars show standard deviations). How are relative humidity, resistivity and the water content of soil related? In the laboratory 200 g of oven-dried soil, sieved using a 3 mm mesh, was placed in a 60 mm deep by 120 mm diameter crystallising dish. After settling the soil into a layer, a hair hygrometer was placed on the surface and the dish was sealed with a glass lid and petroleum jelly. The dish was then placed in a refrigerator at 10°C for 24 h. After removing the dish from the refrigerator the hygrometer was read and removed. The resistivety (ohms) of the exposed soil was measured using an avometer with the two 25 electrodes placed 50 mm apart. This procedure was repeated 12 times for differing degrees of soil dampness within the range 0-50%. The soil was dampened initially by adding 50 mm^ of tap water and mixed thoroughly to give a measure of 2.5%. The other 10 degrees of soil dampness were produced by adding multiples (1-10) of 100 mm^ of tap water and mixing. Additional comparative measurements were made using soil that had been compacted, and with the electrodes separated by multiples of 10 mm up to 90 mm. Where there was only 5% water in the soil at 10°C in the sealed container the relative humidity of the air above the soil was 100%. The soil resistivity measured in ohms gave a calibration curve for the range of water content (Fig. 2) against which the survival of A. triangulatiis was later assessed (Fig. 3). Soil compactness and the distance between the electrodes had no effect on resistivity. Fig. 2. Percent humidity (♦ left axis) and resistivity measurements in ohms (■ right axis) in and above the sieved allotment soil of varying water content in a crystallising dish with an airtight lid kept for 24 h at 1 0°C. What degree of water loss is lethal to A. triangulatus when exposed to air? Thirty specimens were placed in cold tap water for 30 minutes to obtain a standard level of hydration. Each specimen was then blotted with filter paper and allowed to dehydrate at 20°C to a specified weight while on an electronic balance. Two specimens were used for each measurement over a water loss of 10% to 25% of their initial hydrated weight. The specimens were then re-hydrated for 30 minutes, placed on damp allotment soil in Petri dishes in a refrigerator set at 10°C and observed over the following 48 h to determine their survival. All specimens that lost 18% or less of their initial body weight survived the treatment, while those that lost 25% or more died (Table 1). The survival of specimens dehydrated to a percentage between these values was variable. Of the 30 specimens, half recovered from a water loss of 23% of their original body weight. Table 1. Survival after re-hydration of 30 specimens of Arthurdendyus triangulatus allowed to dehydrate in air at 20 degrees C resulting in a loss of between 1 1 and 25% of their initial body weight. % weight loss on % Survival dehydration 11 100 12 100 13 100 14 100 15 100 16 100 17 100 18 100 19 50 20 100 21 100 22 50 23 0 24 50 25 0 Fig. 3. Survival over eight days at 10°C of groups of 10 specimens of Arthurdendyus triangulatus on soil samples containing different quantities of water in Petri dishes. % water in soil What is the level of soil water content at which A. triangulatus will die when on soil beneath a cover? Sixty specimens were kept for eight days in Petri dishes on dampened oven-dried soil. Water was added to 30 g of the dried allotment soil in each dish and mixed thoroughly. The resulting percentages of water in the soils used were: none, 6.6% (20 mm^), 13.3% (40 mm^), 20% (60 mm^), 26.6% (80 mm^) and 33.3% (100 mm^). The soil in each dish was compacted, five specimens were added and the dish sealed with tape. The 12 dishes (two for each dampened soil sample) were each placed in a plastic bag that was also sealed and then 26 kept at 10°C in a refrigerator. The survival of the specimens was checked every 24 h. All specimens on soil with a water content of 33% and above survived while all specimens died where the content was below 7 %. Half of the specimens died on soil with a water content at around 15% (Fig. 3). Specimens were always found on the surface of the compacted soil. Can A. triangulatus drown? Twelve pairs of specimens of A. triangulatus were each submerged in distilled water in 90 mm Petri dishes with lids and were kept at room temperature. Pairs were removed in succession at hourly intervals (1 through to 12), transferred to Petri dishes containing dampened compacted sieved soil and their survival observed over the following 48 h. The experiment was repeated for 24, 36 and 48 hours submersion. A control pair of specimens was kept on dampened soil. Specimens survived submersion for periods up to 48 h. Within an hour of submersion they became flaccid and were incapable of movement but regained their normal appearance and behaviour within 48 h of being out of water. What degree of soil dampness is preferred by A. triangulatus? The dry allotment soil was divided by weight into six parts, one was kept dry and the others had water mixed in to give five degrees of dampness expressed as percentages of water by weight, 9g, (Table 2). One dry part and three with different water contents were packed into separate quarters of a rectangular horizontal veiTnarium with internal dimensions of 600 mm long by 440 mm wide by 10 mm deep. Two artificial earthworm burrows were made diagonally from the comers so that each crossed a soil quarter. They were produced by pressing an 8 mm diameter doweling rod into the soil and then carefully removing it. Eight 0.5 g specimens of A. triangulatus were placed in the artificial burrows: four at the central intersection and one half way along the diagonal crossing each soil portion. The upper glass face of the vermarium was lightly pressed into position on the soil and sealed along the edges with a small quantity of silicone-sealant used in the constmction of aquaria. The vermarium was left in the dark for 48 h at 1 0°C after which the positions of the specimens were recorded. The experiment was replicated five times with different combinations of dampened soil, each degree of dampness being used three times in a combination with a dry soil portion, which acted as a baseline. An identical vermarium in which the four parts of soil had 30% water by weight was used as a control. After 48 h the specimens in the four quarters of the vermarium and at the centre had dispersed. The majority were found in the artificial burrows mnning through soil with 20% to 30% water by weight (Table 2). This gave a significantly different distribution compared with the expected result of two specimens for each quadrant (P < 0.001, x' = 220, df = 5). By contrast, the specimens in the control vermarium with soil of 30% water content moved only a few centimetres from the positions at which they had been initially placed. Table 2. The distribution of specimens of Arthurdendyus triangulatus in a horizontal vermarium containing soil samples of different moisture content in each of the four quarters. The experiment was repeated five times with eight specimens to give a total of 40 specimens. * dry sample present in all tests. % water in No. Distribution Final distrb. the quarters Repl. Start 48 h %of total 0 * 5 5 3 7.5 10 3 3 3 7.5 20 3 3 19 47.5 30 3 3 14 35 40 3 3 1 2.5 50 3 3 0 0 Central 5 20 0 0 Does soil compactness influence the choice of location? The vermarium, of the type described above, was stood vertically and, with its front sealed in place, half filled from the top, which had been opened, with a known weight of dampened soil. The soil was then packed down by repeatedly tapping the vermarium on the floor to give a bulk density of 13 g mm'^. Eight specimens of A. triangulatus were dropped onto the surface of the soil and the space above filled with a known weight of similar soil to give a bulk density of 8 g mm'^. The positions of the specimens were recorded after the vermarium had been sealed and left in the dark for 48 h at 10°C. The experiment was replicated three times and a control vermarium containing only loose soil was used. After 48 h, of the 24 specimens in the experimental vermarium four were found at the interface between the compacted and loose soils and the remaining 20 (83%) were all in the loose soil. None were found in the compacted soil. In the control vermarium the specimens moved in random directions for short distances. DISCUSSION To survive for any length of time, A. triangulatus requires a damp habitat. Specimens dehydrate when exposed to air on a dry surface and die after losing a quarter of their initial body weight. They will, however, live for weeks in Petri dishes without soil but covered with cling film on which water has condensed. When soil in a sealed container had 5% or more water the air above the soil was saturated. Yet, even with air of high relative humidity (Fig. 2), 27 A. triangulatus will die if it remains on soil with a low water content (Fig. 3). Death resulted because the suction pressure, pF, (White, 1987) of the soil removes water from A. triangulatus. For survival, A. triangulatus needs to be on or in soil with a gravimetric water content of 30%. In a vermarium and under debris in the allotment specimens of A. triangulatus selected soil with this water content. Very damp soils do not appear to be detrimental since specimens survived total immersion for at least 48 hours. In Scotland, where rainfall often saturates soils, the surface water typically drains away within that period. We may infer that the soil water content rather than temperature is the more important parameter in determining the geographical distribution of A. triangulatus in Great Britain. The laboratory experiments showed that specimens of A. triangulatus retreat into the soil when the surface temperature was lowered (Gibson, Ponder and Cosens, 2004). In the allotment the temperature at a depth of some 300 mm rarely exceed the upper and lower lethal limits for A. triangulatus. However, under natural conditions the soil could dry out at this depth should the water table fall. In the soil compactness study, where burrows were not present, specimens of A. triangulatus chose loose rather than compacted soil. Presumably this is because movement within loose soil is possible and does not require an ability, which A. triangulatus lacks, to burrow. When burrows are present, specimens of A. triangulatus use them in preference to moving through the soil per se (Lillico et a!., 1996). Whilst there may be a geotactic effect, none was apparent in the present studies or those of other workers (Cannon et al., 1999). Since the numbers of A. triangulatus in the environment have never been reported to increase exponentially we may assume that a large proportion of the population dies before reproducing. Young specimens are most likely to be vulnerable. Whether predation is important in limiting numbers is not known. However, carabid and staphylinid beetle larvae have been reported to feed on A. triangulatus in its natural habitat (Gibson et al. 1997; observation by Anna Gibson, August 2002). Also, when exposed on the soil surface A. triangulatus is eaten by farm-yard ducks and geese (Jones and Boag, 2001) and by frogs in the laboratory (Anna Gibson, personal communication). Although predators may be individually insignificant in controlling A. triangulatus population, they may be collectively effective. Environmental conditions produced by rainfall and temperature will affect the abundance of potential invertebrate predators. Under some conditions or in different seasons or years predators may be numerous and, therefore, have a significant affect on A. triangulatus numbers. Of course, the juvenile stages of A. triangulatus and potential predators (in this case, the beetles and their larvae) must occur in the same place and at the same time. ACKNOWLEDGEMENTS We wish to thank Keith Hood and Simon Lillico for their help in collecting data used in these studies. REFERENCES Boag, B., Yeates, W.G., Johns, P.M., Neilson, R., Palmer, L.F. and Legg, R.K. (1995). Distribution of the New Zealand flatworm (Artioposthia triangulata) in New Zealand and Seotland. Acta Zoologica Fennica 196, 212-214. Cannon, R.J.C., Baker, R.H.A., Taylor, M.C. and Moore, J.P. (1999). A review of the status of the New Zealand flatwonn in the UK. Annals of Applied Biology 135, 597- 614. Gibson, P.H. and Cosens, D.J. (2000a). Density of the New Zealand flatworm, a predator of earthworms, in allotments in Edinburgh between 1993 and 1995. Annals of Applied Biology 137, 347-352. Gibson, P.H. and Cosens, D.J. (2000b). Studies on the local dispersal of the New Zealand flatworm, a predator of earthworms, in allotments in Edinburgh between 1993 and \995. Annals of Applied Biology 137, 353-359. Gibson, P.H., Cosens, D.J. and Buchanan, K. (1997). A chance field observation and pilot laboratory studies of predation of the New Zealand flatworm by the larvae and adults of carabid and staphylinid beetles. Annals of Applied Biology 130, 581-585. Gibson, P.H., Ponder, K.L. and Cosens, D.J. (2004). The effect of temperature on the survival and distribution of the New Zealand flatworm Arthiirdendyiis triangulatus. Glasgow Naturalist In Press. Jones, H.D. and Boag, B. (2001). The invasion of New Zealand flatworms. Glasgow Naturalist 23, 77-83. Jones, H.D. and Gerard, B.A. (1999). A new genus and species of terrestrial planarian (Platyhelminthes; Tricladida; Terricola) from Scotland and an emendation of the genus Artioposthia. Journal of Natural History 33, 387-394. Lillico, S., Cosens, D. and Gibson, P. (1996). Studies on the behaviour of Artioposthia triangulata (Platyhelminthes; Tricladida), a predator of earthworms. Journal of Zoology, London 238, 513-520. White, R.E. (1987). Introduction to the principles and practice of soil science. Blackwell Scientific Publications, London. 28 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 29-33. THE EFFECT OF TEMPERATURE ON THE SURVIVAL AND DISTRIBUTION OF THE NEW ZEALAND FLATWORM ARTHURDENDYUS TRIANGULATES Peter H. Gibson, Katherine L. Ponder and Derek J. Cosens Institute of Cell, Animal and Population Biology, University of Edinburgh, Ashworth Laboratories, West Mains Road, Edinburgh EH9 3JT ABSTRACT In the early spring, temperatures below the soil surface and under items of surface debris such as plastic sheeting were greater than those at the exposed surface. The population density of Arthurdendyus triangulatus on the soil surface immediately under debris was positively correlated with the soil surface temperature. However, temperature alone could not account for differences in the densities of A. triangulatus under the debris. In the laboratory most specimens of A. triangulatus were unable to survive freezing for more than an hour. At low temperatures they became more active at first but then became sluggish. In earth-filled vermaria A. triangulatus typically avoided adverse temperatures by moving down the soil column. It tolerated temperatures up to 23 °C but showed a preference for 19°C. Some three times as many earthworms were eaten by A. triangulatus when the soil was at 10°C than at 3°C. INTRODUCTION The accidental introduction of the New Zealand flatworm Arthurdendyus triangulatus (Dendy) (formerly Artioposthia triangulata, see Jones and Gerard, 1999) into Britain nearly forty years ago presents a potential threat to the numbers and species of earthworms. A. triangulatus has become established throughout Northern Ireland and Scotland (Willis and Edwards, 1977; Boag, Yeates, Johns, Neilson, Palmer and Legg, 1995) and is now found in England (Boag, Neilson, Palmer, and Jones, 1994). It typically occurs in urban areas and has probably been spread as egg capsules in the soil of bedding plants bought at garden centres and market gardens. The general biology and distribution of the species have been reviewed by Cannon, Baker, Taylor and Moore (1999), who concluded that there is ‘need for a better understanding of the environmental conditions under which A. triangulatus survives’. Accordingly, this paper examines the effect that temperature has on the survival of A. triangulatus. GENERAL PROCEDURES Field studies were carried out at Midmar Drive allotment, Edinburgh, Scotland (Ordinance Survey sheet 66, grid reference NT 252707). Specimens of A. triangulatus of varying body weight were collected from under debris for use in laboratory studies. They were kept for up to 7 days on fresh compacted allotment soil in 9 cm diameter soda glass Petri dishes at 10°C in a refrigerator. The soil was a loam derived from a glacial till consisting of a reddish sandy silt-clay of carboniferous sandstone origin. METHODS AND RESULTS What is the typical soil temperature profile in the early spring? From February 10th to March 10th 1995 the soil temperature was recorded every 15 minutes using thermisters connected to a four channel automatic temperature recorder powered by a 12 V battery (Grant Instruments Ltd, Cambridge). Three thermisters were inserted horizontally into the vertical side of an excavated pit at depths of 100, 200, and 400 mm and the pit was then refilled with the original soil. The fourth thermistor was placed on the soil surface. Over 24 h the temperatures within the soil column fluctuated in a regular manner (Table 1, Fig. 1): at the surface there was a range of 6.2°C; at 100 mm 1.5°C; at 200 mm 0.7°C; at 400 mm 0.4°C. Table 1 . Mean maximum and minimum soil temperature (°C) at different soil depths over a period of 23 days from 10 February to 10 March (+SD standard deviation; also see Fig. 1). Surface Min Max -0.35 5.88 ±1.76 ±2.39 200 mm depth Min Max 3.33 4.01 ±0.41 ±0.95 100 mm depth Min Max 1.83 3.37 ±0.80 ±1.52 400 mm depth Min Max 3.70 4.11 ±0.30 ±0.67 Fig. 1. Allotment soil temperature profiles for the 10th Febmary at three depths (since the temperature at 400 mm fluctuated little from 4°C and was so similar to that at 200 mm it is omitted from the graph). The profile is typical for the 29 days of the study (also see Table 1). 29 How do A. triangulatus numbers relate to variation in soil surface temperature? Soil temperature and numbers of A. triangulatus under debris were recorded over 17 days from March 19th. Numbers were monitored under 17 items of debris ( 1 3 wooden planks, 2 plastic sheets and 2 stones) evenly distributed over the soil surface and covering a combined area of 40 m^ within a 400 m" area of cultivated soil. The soil temperature was recorded at midday from a 10 mm depression beneath a flat stone at the centre of the experimental area. Without moving the stone a thermister probe of the Grant recorder was carefully inserted into the depression to record the temperature. At midday the temperature ranged from 2.5 to 8.0°C. The numbers of specimens of^. triangulatus found correlated significantly with the soil surface temperature (Fig. 2) (Speannan’s rank correlation r = 0.67,P<0.001). Fig. 2. Numbers of Arthiirdendyus triangulatus (♦) on the soil surface under debris lying within 400 over 1 7 days in March and April and temperature (■) recorded in a shallow pit below a flat stone over the same period. How does soil surface temperature compare beneath two different types of debris? The soil surface temperatures were recorded under two recycled 600 x 900 mm plastic ‘feed’ sacks filled with compost, a 50 mm thick concrete paving stone and at the soil surface over five days from March 12th to 16th. One sack regularly had specimens of A. triangulatus beneath it while the other had none. The sacks were 5 m apart. During a 24 h period temperature under the two compost-filled plastic sacks fluctuated in a similar manner from 5 to 9°C, a range of 4°C. The mean temperature for the sack with specimens of A. triangulatus was 7.1°C (SD = 2.6) and without specimens it was 7.3°C (SD = 1.0). Under the paving stone it fluctuated from 1 to 1 1 °C, a range of 10°C. Since the exposed soil surface temperature ranged from 1 to 14°C, a range of 13°C, compost- filled plastic sacks clearly buffered the changes in temperature over 24 hours. By contrast, the paving stone had a limited buffering effect (Fig. 3). Fig. 3. Temperature over the 12th to 16th March under a paving stone (•) (regression line, R" = 0.8767, P < 0.001) and a compost-filled plastic sack which had many specimens of Arthurdendyus triangulatus under it (■) (regression line, R~ = 0.2526, P < 0.001) (R' = 0.3055 & P < 0.001 for the sack, data points not shown, under which specimens were not found but the temperature had the same distribution). 12 ^ ^ 1 -1 4 9 14 Soil surface temperarure °C What is the cold tolerance of A. triangulatus? A plastic tray of 600 x 400 x 100 mm was filled with water to a depth of 30 mm and six glass specimen tubes of 75 x 25 mm were stood upright at one end. A 200 mm diameter cooling coil was placed at the other end of the tray which was then lagged. When the water froze a specimen of A. triangulatus was placed at the bottom of each tube which was then sealed with a cork stopper. As the apparatus cooled, the temperature in the tubes fell to -2°C. The specimens were prodded every five minutes over a period of 20 minutes and then every 10 minutes to determine whether they were alive. When a specimen became frosted it was removed from the tube to thaw at 10°C and then gently prodded to determine whether it was alive. Dead specimens disintegrated. The activity of other specimens in Petri dishes in crushed ice was also observed. Of 49 specimens with body weights of 0.05-1.00 g, 42 (86%) died within 60 minutes. There is a correlation between body weight and survival time (Pearson’s correlation r = 0.34, n = 49, P = 0.05). However, this relationship was skewed by seven specimens which survived the longest with one of 0.84 g living for 3 h. Five of these were at the heavier end of the range (mean 0.48 g, SD = 0.24). This compares with 14 specimens of similar weights which died in less than an hour. Some small specimens took refuge under the cork 30 stoppers and survived longer than they might otherwise have done. Observations on specimens in Petri dishes with crushed ice showed that they became active as the temperature fell and then sluggish before freezing. Their bodies often blistered. Specimens with partially frozen posterior ends survived when thawed and although the frozen ends disintegrated the wounds healed during the following two weeks. Specimens that froze completely disintegrated when thawed as observed earlier. What effect does temperature have on the vertical distribution of A. triangulatus in the soil column! A glass-fronted vermarium with internal dimensions of 600 x 440 x 10 mm was prepared. Five mercury thermometers had been stuck horizontally, with clear silicone-sealant, 100 mm apart on the inner face of the glass front with their bulbs lying along the vertical midline but without touching the glass. The first thermometer was 500 mm from the base and the fifth 100 mm from the base. The vermarium was stood vertically on one of the 440 mm sides and filled to the level of the first thermometer from the upper end with fresh allotment soil that had been sieved with a 3 mm mesh. A doweling rod of 8 mm diameter was pushing vertically into the soil at equal distances apart on the side opposite the thermometers to produce three artificial earthworm burrows. Five specimens of A. triangulatus, each of 0.5 g body weight, were placed on the soil surface and the vermarium was closed with a lid that was held in position by clear silicone-sealant. A temperature gradient was created using a flat 200 mm diameter cooling coil placed vertically 20 mm behind the vermarium with its centre level with the upper soil surface. The vermarium was kept in a darkened room at 10°C and illuminated with a dim red light. As a vertical temperature gradient developed from the soil surface downwards over 6 hours the soil temperatures and the positions of the specimens were recorded every 15 minutes. The experiment was repeated four times with fresh soil and different specimens, as well as being repeated without using the cooling coil to give a control. The temperature of the soil in the vermarium decreased progressively over 300 minutes to produce a gradient from -3.0°C at the soil surface to 1 .5°C at a depth of 300 mm (Fig. 4 upper graph). Some specimens froze near the top of the vermarium due to an initial rapid drop in temperature. However, the positions of specimens showed that as the temperature fell most moved down the soil column to stop at a mean depth of 390 mm (SD = 39.2, n = 4) (Fig. 4 lower graph) where the temperature was above 0°C (Fig. 4 upper graph). A Mann-Whitney U-test showed that specimens moved significantly further down the vermarium under the experimental conditions than under the control conditions (median depths: experimental 404 mm, control 20 mm; W = 591, P<0.001). In a similar experiment the temperature gradient was created using a red Philips 250 W heating lamp suspended 200 mm above the soil surface. Temperatures at the various depths and the positions of the five specimens were recorded after 24 h. The experiment was replicated three times with, and three times without, the lamp. Fig. 4. Upper Graph. Soil temperatures in the vermarium as cooling proceeded over 300 minutes at: the surface ♦, a depth of 100 mm •, 300 mm A, and 400 mm ■. The data points are mean values for the four experiments, error bars are omitted for clarity as are data for a depth of 200 mm. Lower Graph. The positions of four specimens within the vermarium that remained alive during the cooling period. Each worm, after a lag period at the soil surface (■ 105 min, ♦ 75 min, • 60 min, ▲ 30 min; mean = 67.5 min, SD = 31.2), moved down into the soil column where it remained until, due to the accumulation of cold air at the bottom of the apparatus (400 - 600 mm), it moved upwards to where the soil temperature was above freezing (see Fig. 4a). Minutes from leaving soil surface The temperature gradient within the soil column ranged from a mean of 23.3°C (SD = 1.2, n = 3) at the upper soil surface to a mean of 13.7°C (SD = 1.5, n = 3) at a depth of 400 mm. The specimens moved down the soil column during the experiment and after 24 h they were found in positions where the soil temperature was between 15 and 22°C (mean 18.9°C, SD = 2.4, n = 12). As in the 31 previous study, the specimens moved significantly further down the experimental column than the control column (Mann-Whitney U-test: experimental median depth =101 mm, control = 22 mm, n = 15, W = 304.5, P<0.003). Do low temperatures inhibit the predatory behaviour of A. triangulatus? Two plastic tubes, each 120 mm in diameter and 600 mm long, were positioned 50 mm apart and glued at one end to a sheet of plastic that formed a base. The apparatus was stood vertically and filled with 2.5 kg of fresh allotment soil that had been sieved using a 3 mm mesh. A 200 mm diameter cooling coil was placed half way between the tubes with its centre level with the soil surface. The apparatus was lagged. Ten large specimens of the earthworm Lumbricus terrestris L. were placed on the surface of the soil in each tube which was then sealed. The apparatus was left for five days during which the earthworms made burrows. Five 0.5 g specimens of A. triangulatus were then placed on the surface of the soil of each tube which was resealed and the apparatus cooled for a week. The contents of the tubes were then examined to determine the number of earthworms eaten. The experiment was repeated and a control was run without the cooling coil. This gave four experimental tubes and two control tubes. The soil in the experimental tubes cooled to 3°C while the soil in the control tubes remained at a temperature of 1 0°C. Of the 40 earthworms used in the experimental tubes 34, or about nine tenths, were recovered while of the 20 earthworms in the control tubes nine, or about half, were recovered. Therefore, significantly more earthworms were eaten at the higher temperature (P < 0.001, = 25, df= 1). DISCUSSION In Edinburgh temperatures can vary between -7°C and 25°C (annual record of the Royal Edinburgh Botanic Gardens for 1999). The numbers of A. triangulatus found under debris correlated positively with changes in environmental temperature, and in the laboratory, specimens were found to move down the soil column as the surface temperature rose or fell. Some of the items of debris buffered the variations in temperature at the soil surface. In spite of this buffering the densities of specimens cannot be accounted for by the nature of the debris (Gibson and Cosens, 2000a). For example, the numbers of A. triangulatus beneath the two compost-filled sacks were different even though they were subject to the same temperature variation. These findings suggest that other physical factors are involved in controlling A. triangulatus numbers and the most important of these may be the soil-water content (Gibson and Cosens, 2004). In the heated vermarium, for example, the soil surface became dry and as a result specimens may also have descended the soil column to the avoid desiccation. In the vermarium studies, A. triangulatus was found to be unable to survive freezing for any length of time and descended the soil column when the surface temperature fell below 0°C. A. triangulatus was found to be able to tolerate soil temperatures up to 23°C for at least a few hours but preferred soil at 19°C. These findings are consistent with those of Blackshaw and Stewart (1992) who found an upper lethal limit of 24°C and an ability to survive for up to two weeks at 20°C. The lower lethal limit must be just below freezing. The abundance of A. triangulatus in the soil is dependent on a number of factors. One is the predation pressure on earthworms which is indicated by how long A. triangulatus has been present at a particular location (Blackshaw, 1990; Lillico et al, 1996; Gibson and Cosens, 2000b). Blackshaw (1991, 1997) found that the numbers of earthworms eaten by A. triangulatus increased when soil temperature rose from 5°C to between 12°C and 16°C. Similarly, in this study three times as many earthworms were consumed at 10°C as at 3°C. That is, the biomass of earthworms eaten increases with temperature. This has implications for agriculture in Scotland should the mean annual temperature rise due to global warming. Were this to occur, the threat to earthworm numbers, first noted by Blackshaw (1990), may become a serious reality. Until now the numbers of earthworms and A. triangulatus in Scotland appear to have reached an equilibrium dependent on a predator/prey oscillation (Boag et al., 1995; Gibson and Cosens, 2000b) but this balance could alter were temperature to rise. Flatworm consumption of earthworms is unlikely to increase in England because global warming would also result in significant drying of soils which is likely to be a major influence in controlling the distribution of A. triangulatus (Gibson and Cosens, 2004). The conclusion based on this study and those of Blackshaw, is that temperature per se may not directly limit the distribution of A. triangulatus within Great Britain. A. triangulatus appears to tolerate temperatures between freezing and 24°C and has the ability to move sufficiently quickly to avoid extremes (Gibson and Cosens, 1998) by using surface shelters, earthworm burrows or crevices (since it cannot burrow) within the soil. The upper lethal temperature probably depends on thermal inactivation of enzymes (Schmidt-Nielsen, 1979) and is likely to be well above 24°C. The most serious threats to the survival of A. triangulatus are, however, likely to be the soil-water content and relative humidity (Gibson and Cosens, 2004) which are, of course, related to temperature. ACKNOWLEDGEMENTS We wish to thank Simon Lillico for his help in collecting the data used in this study. REFERENCES Blackshaw, R.P. (1990). Studies on Artioposthia triangulata (Dendy) (Tricladida: Terricola), a predator of earthworms. Annals of Applied Biology 1 16, 169-176. Blackshaw, R.P. (1991). Mortality of the earthworm Eisenia fetida (Savigny), presented to the terrestrial 32 planarian Artioposthia triangulata (Dendy) (Tricladida; Terricola). Annals of Applied Biology 118, 689-694. Blackshaw, R.P. (1997). The planarian Artioposthia triangulata (Dendy) feeding on earthworms in soil columns. Soil Biology and Biochemistry 29, 229-302. Blackshaw, R.P. and Stewart, V.l. (1992). Artioposthia triangulata (Dendy, 1894), a predatory terrestrial planarian and its potential impact on lumbricid earthworms. Agricultural Zoology Reviews 5, 102-219. Boag, B., Neilson R., Palmer, L.F. and Jones, H.D. (1994). A further record of the New Zealand flatworm Artioposthia triangulata (Dendy) in England. Plant Pathology 43, 220-222. Boag, B., Yeates, W.G., Johns, P.M., Neilson, R., Palmer, L.F. and Legg, R.K. (1995). Distribution of the New Zealand flatworm {Artioposthia triangulata) in New Zealand and Scotland. Acta Zoologica Fennica 196, 212-214. Cannon, R.J.C., Baker, R.H.A., Taylor, M.C. and Moore, J.P. (1999). A review of the status of the New Zealand flatworm in the UK. Annals of Applied Biology 135, 597- 614. Gibson, P.H. and Cosens, D.J. (1998). Locomotion in the terrestrial planarian Artioposthia triangulata (Dendy). Pedobiologia 42, 241-25 1 . Gibson, P.H. and Cosens, D.J. (2000a). Density of the New Zealand flatworm, a predator of earthworms, in allotments in Edinburgh between 1993 and 1995. Annals of Applied Biology 137, 347-352. Gibson, P.H. and Cosens, D.J. (2000b). Studies on the local dispersal of the New Zealand flatworm, a predator of earthworms, in allotments in Edinburgh between 1993 and 1995. Annals of Applied Biology 137, 353-359. Gibson, P.H. and Cosens, D.J. (2004). The effect of water content and compactness of soil on the survival of the New Zealand flatworm Arthurdendyus triangulatus. Glasgow Naturalist In Press Jones, H.D. and Gerard, B.A. (1999). A new genus and species of terrestrial planarian (Platyhelminthes; Tricladida; Terricola) from Scotland and an emendation of the genus Artioposthia. Journal of Natural History 33, 387-394. Lillico, S., Cosens, D. and Gibson, P. (1996). Studies on the behaviour of Artioposthia triangulata (Platyhelminthes; Tricladida), a predator of earthworms. Journal of Zoology, London 238, 513-520. Schmidt-Nielsen, K. (1979). Animal physiology: Adaptation and environment. Cambridge: Cambridge University Press, pp 20-213. Willis, R.J. and Edwards, A.R. (1977). The occurrence of the land planarian Artioposthia triangulata (Dendy) in Northern Ireland. Irish Naturalist Journal 19, 112-116. 33 34 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 35-52 STUDIES ON THE CONSERVATION BIOLOGY OF IRISH LADY’S-TRESSES ORCWID, SPIRANTHES ROMANZOFFIANA; 1) POPULATION SIZES, GRAZING, VEGETATION HEIGHT AND CAPSULE STATUS AT REFERENCE SITES Richard Gulliver ', Mavis Gulliver ^ Margaret Keirnen\ and Christopher Sydes^ ' and ^ to whom correspondence should be addressed, Carraig Mhor, Imeravale, Port Ellen, Isle of Islay, Argyll PA42 7AL, ^ Grianan, Dmmclach, Isle of Colonsay, Argyll, PA61 7YR, Scottish Natural Heritage, 2 Anderson Place, Edinburgh, EH6 5NP ABSTRACT Studies were undertaken on Irish lady’s-tresses orchid to determine the annual variation in recorded population size; and to examine a range of factors which were considered likely to affect population size, including impact of grazing, means of reproduction and features of the vegetation. Two special attributes of the species, a) the presence of a mycorrhiza and hence a source of organic carbon from the fungus, and b) its status as a Biodiversity Action Plan species; lend special biological and conservation importance to the investigations. At two sites subject to a grazing break in summer, the number of plants in bloom has risen each year between 1999 and 2002 with a fall in 2003. In every year 65% or more of plants in bloom at these two sites were newly recorded plants. At two sites with no grazing break 1-3 plants had flowering stems in 2002 and 2003, however moderate sized populations were in existence, indicating that the number of plants with flowering stems is not always a useful guide to population size. Data are provided on the impact of grazing on leaves by slugs and vertebrates; and on flowering stems by domestic stock and rabbits. The locations of the sites studied are referred to by code numbers to reduce the likelihood of visits by collectors of rare orchids. Spiranthes romanzoffiana plants occurred at a wide range of vegetational heights at six sites including a) two which had experienced grazing breaks since 1996 (in June, July and part or all of August) b) one traditionally managed site with a summer grazing break, c) two with summer grazing and d) one with summer grazing break 1996 to 2000 but not thereafter. At one site they had a tolerance of a regime of heavy grazing by domestic stock which produced low swards. Paired data indicated lower vegetational heights in 2003 cf 2002 at two sites with a summer grazing break. Measurements of the lowest grazed level created by domestic stock were used to indicate the intensity of grazing and associated activities e.g. trampling and defecation. At three sites in the same dune system, frequency of Juncus articulatus and Juncus acutiflorus x articulatus (pooled) ranged from 60% to 28% to 0% in 10x1 0cm quadrats centred on the positions of Spiranthes romanzoffiana plants. In order to appreciate fully the importance of a range of biotic and abiotic factors and management influences, detailed records are required. Such detailed information may not be easy to summarise and communicate. Capsules which had developed to full size, and then dried and dehisced, were observed in Spiranthes spiralis in England, but not in Spiranthes romanzoffiana in Scotland. To gain a more complete understanding of the species, it would be desirable to investigate aspects of sexual and vegetative reproduction under controlled conditions. A long-term objective might also be to experimentally simulate the effects of defoliation, trampling and organic matter input from dung The fact that Spiranthes romanzoffiana can occur in the underground form for up to six years makes long-temi study of both the species and its environment especially important, i.e. seven years must elapse after the initial observation before one can assume that a plant at a marked position has died. Key words and phrases: vegetation height, orchid, population, conservation, grazing, Juncus species INTRODUCTION Spiranthes romanzojfiana (Irish lady’s-tresses orchid: Figures Al, A2, A3, Bl) is a white flowered orchid which blooms in July and August. The species occurs in Devon in England and the north, south and west of Ireland. In Scotland it is present in the Hebrides from Islay to Benbecula; and on the west coast. There is also one 1970 record approximately 30km inland from the coastline; for distribution map see Preston, Pearman and Dines (2002). It has recently (August 2002) been found on Tiree (Bowler 2003). It was formerly a British Red Data Book (BRDB) species but botanical exploration and recording in the nineteen eighties and nineties raised the number of hectads (10x1 0km squares) in which the plant had been found to above the critical level of 15, and it no longer qualified for this status. It is currently a ‘Nationally Scarce’ species sensu Cheffmgs (2004), i.e. Scarce sensu Stewart, Pearman and Preston (1994); and a Biodiversity Action Plan Priority Species, see UK Biodiversity Action Group (1999). New lateral buds (normally one, sometimes two i.e. twins, very rarely three or four) usually emerge above the soil surface between July and October, so the plant has green tissue present throughout the winter months, see Gulliver et al. (2000) for illustrations of the lateral bud, the nature of the inflorescence in September, and for further illustrations of the plant in bloom. As with all members of the Orchidaceae, the species is myco-rrhizal, and so has a source of organic carbon in addition to that generated by photosynthesis. 35 In the British Isles the other extant member of the genus, Spiranthes spiralis (autumn lady’s-tresses: Figure A4) occurs in England, Wales and Ireland, but not Scotland. Spiranthes aestivalis (summer lady’s-tresses) formerly occurred in the New Forest in England, in Guernsey and in Jersey, but is now extinct in the British Isles, Preston, Pearman and Dines (2002). Spiranthes romanzoffiana can frequently be observed growing amongst the stems of Junciis acutiflorus x articulatus (Jimciis surrejanus). Figure Al; and ofJimcus articulatus which is stated to be moderately resistant to grazing by Grime, Hodgson and Hunt (1986). However the absence of stems of Jiincus species (mainly Juncus acutiflorus X articulatus) outside the exclosure at KA on Colonsay, shown in Figure A5, is due to the presenee of heavy grazing. The rhizome system of Juncus articulatus is described as subcaespitose (almost tufted) and of Juncus acutiflorus x articulatus as far creeping by Blackstoek and Roberts (1986). Species with far creeping rhizomes which produce daughter plants at intervals are grazing tolerant, as consumption by grazers can normally only involve part of the total set of ramets. Tufted taxa may suffer total consumption; however the close grouping of the stems may make the mass of vegetation less attractive for plant species of limited palatability. The emergence of stems of other species from underground organs may be partly or wholly impeded by the dense grouping of stems, stem bases and associated roots of tufted plants. The roots of Juncus articulatus are sometimes mycorrhizal. Grime, Hodgson and Hunt (1986), and the same may well be the ease for Juncus acutiflorus x articulatus. The status of Spiranthes romanzoffiana on Colonsay in 1995 and 1996 is given in Gulliver (1996) and (1997); the impact of grazing on plants on Colonsay in 1999 in Gulliver et al. (2000) and the impact of grazing in general by Gulliver et al. (2003); the results of long term monitoring of plants on Barra by Robarts (2000); the vegetation associated with the species at 16 locations on Coll by Henderson (2001); the genetic constitution of populations in Ireland and Scotland using molecular methods by Forrest (2001) and Forrest et al. (2004); and the results of research into the conservation biology of the species in Gulliver (2002). In this account, plant taxonomy follows Stace (1997): English plant names follow the journal house style. These investigations of small, medium and large sized populations form part of two Scottish Natural Heritage research projects with further funding from the Botanical Research Fund. Studies which involved the use of exclosures on the variable appearance over time of individuals in small populations commenced in 2003, Gulliver et al. (2004b, this volume). The aim of the research was firstly to record the annual variation in number of flowering stems (1999-2003) and for 1999-2001 to record the composition of the populations. Secondly it was to examine a range of factors which are likely to affect population size, including impact of grazing, means of reproduction and features of the vegetation. METHODS General Five sites (LF, KA, KB, KC and SF) on Colonsay were visited regularly throughout the year during the period 1999 to 2001; four less intensively so in 2002 and 2003. Some general recording had taken place previously, between 1991 and 1998. From 1999 onwards individual plants were marked by the use of an adjacent black plastic peg with a ‘mushroom shaped’ black top. Referencing systems were established to allow plants to be relocated. One site (BB subsite A) was studied on Barra. Due to a previous history of rare orchids having been periodically dug up in the British Isles, locations are not identified in detail, but are referred to by code letters. Any one plant may either be in the flowering (F), vegetative (V) or underground (U) phase in any one growing year. By 1999 the longest period a plant had been recorded underground was 4 years (Dr James Robarts pers comm., in Gulliver et al. 2000). Continued observations by Dr Robarts on Barra have shown that the longest period recorded underground for any single plant is 6 years (Dr James Robarts pers. comm.). Flowering and vegetative plants are differentiated by the presence of a flowering stem, which is sometimes truncated by grazing. Flowering, vegetative and bud-only plants were recorded during the period of multiple site visits, 1999-2001. Bud-only plants were recorded from July to October and were a) plants appearing above ground having spent the previous 12 months (at least) in the underground state or b) the newly generated second member of a pair of twin buds on a plant (flowering or vegetative) above ground in summer. They are included in the total as they represent part of the population present on site from a conservation perspective, but they are actually in the next growing year to the flowering and vegetative plants present. On multiple site visits plants were occasionally observed in mid to late August or early September with no stem whatsoever. On a previous visit a stem was present. On a system involving a single visit per year recording such plants would be classed as vegetative, on a multiple visit system as flowering. (Plant 6 at site KC on 14 August 2000 is an example of this phenomenon). The cumulative totals detected over all visits in any one year were always higher than the number of plants recorded on a single visit (Gulliver, 2002). For sites KC and LF the cumulative number detected from August 1999 to September 2001 is 36 referred to as the ‘monitored set’, n = 18 and 37 respectively. The ‘monitored set’ of positions was examined in 2002 (August) and 2003 (May, July and September) at KC and LF. For KA and KB plants in bloom were recorded in August in 2002 and July in 2003. In addition at KA and KB a subset of the population was investigated in July 2003. Shrivelled inflorescences were examined at KA and KB in detail in September 2001, 2002 and 2003. Site characteristics Sites KA and KB were protected from grazing by sheep and cattle from very late May/early June to late September/early October in 2001, 2002 and 2003; having been surrounded by fixed, gated exclosures in Spring 2001; Figure A5 shows part of the exclosure round site KA. These populations have diffuse margins. Of the 49 and 25 plants recorded in bloom on all visits to KA and KB respectively in 1999, one at KA (31a) and one at KB (21a) are now outside the exclosure. Outside these two exclosures grazing was more or less continuous in 2001, 2002 and 2003. The grazing history of the whole area which includes sites KA, KB and KC is complicated. Appendix A gives relevant details from 1991 onwards of a) presence or absence of a summer grazing break over time b) changes associated with the erection of a gated, moor/dune fence in 1996. From 1996 onwards grazing stock might be on either or on both sides of the fence. Prior to its erection they could move freely between moor and dunes. Site LF was grazed lightly, with considerable variation in the length of the grazing period. Site SF was more or less continuously grazed over the whole area, however it was very variable in nature often resulting in differing and varying intensities of grazing from position to position. Site TG was usually grazed more or less continuously. Site LF is site 1 in Gulliver et al. 2000, Site KA is site 2, site KB is site 3 site KC is site 4 and site SF is site 5. Details of site management and characteristics for the study sites, including adjudged intensity of grazing integrated over all times of year and all types of vegetation within the site are provided in Appendix A and B. Some parts of site KA were very marshy, periodically resulting in the sinkage of single or groups of marker pegs and an absence of detectable plants in highly disturbed ground. A degree of peg sinkage also occurred at LF, KB, and KC. Dates of visits are given in the relevant tables. At site BB subsite A on Barra the management is relatively constant every year, stock are absent in June, July and August. Frequently sheep are kept on the whole area of unenclosed croft land, which includes subsite A, for longer than cattle in the spring, and they are often brought back in the autumn in advance of the cattle. In general a greater degree of regularity of management occurs at sites which are managed by a grazing committee or by a group of crofters following an agreed procedure each year, compared with those that are not. Vegetation height and Jimcus acutijlorus x articulatus plus Juncus articulatus Vegetation height was recorded as an index of the intensity of use of the area by domestic stock, e.g. very low heights indicate heavy grazing, tall heights indicate light grazing, variable heights indicate patchy (selective) grazing on either a more or less uniform sward or on a mixture in the sward of plants of differing palatability. Three height bands have been used by MacDonald et al. (1998a and 1998b) to assess, together with other measurements, the intensity of grazing in four habitats in upland Scotland, (smooth grassland, flush, tall herb and tussock grassland). In general, high levels of grazing are associated with high levels of trampling and high levels of dung production, though some zones may be avoided and some heavily used for defecation. Vegetation height was also recorded to gain data on the range of heights of vegetation in which the plant could survive, i.e. to characterise the habitat of the species. In agricultural swards, height is recorded by taking measurements either of single leaves or of the whole sward i.e. the vegetation, (‘t Mannetje 1978). The swards encountered in the immediate vicinity of Spiranthes romanzoffiana plants, or their previously recorded positions, all exhibited considerable variation in height. Hence a system was devised to accommodate this feature. This involved firstly taking ten height measurements at a sampling station e.g. the sward around the position of a plant and then selecting the median value of this single sample (first order median). The process was then repeated for every sampling station. From this set of values a second order median was obtained which characterised the site as a whole. Different configurations of sampling stations were used, depending on the nature of the site. Further details of this topic and other aspects of the measurement of vegetation height are provided in Appendix C. Vegetation height was measured to the point where the density of the component stems and leaves of the sward diminished appreciably, as used in the former (pre 1973) Nature Conservancy’s Meadow Grassland Survey, (the late Mr Derek Wells, pers. comm.). This point is frequently at about 66% of the height of the longest leaf or stem. The height of plants or small stands of Juncus acutiflorus x articulatus and Juncus articulatus was measured not to the top of the tallest stem but to the point where stems and leaves became less dense. Vegetation height was recorded in 5 mm steps between 10 and 30mm and in 10mm steps thereafter. The height of plants and small stands of Juncus acutiflorus x articulatus and Juncus articulatus was measured in 20mm steps. 37 Where there were clearly different layers within the vegetation, these were measured separately. For example at one sampling station one might encounter a low grass and sedge rich, well-grazed i.e. first layer e.g. see Figure Al, a taller, lightly- grazed layer of Jiincus acutiflorus x articulatus {Juncus surrejaniis) and an ungrazed layer of Junciis effusiis. At site KA and BB subsite A vegetation height was measured at the position of 164 and 44 plants in 2002. Height at a sample of 30 and 38 respectively at the same positions was recorded in 2003, constituting paired data in time. The occurrence of the taxa in the Juncus acutiflorus X articulatus/Juncus articulatus group was investigated at four sites. Records were made of the presence of Juncus acutiflorus x articulatus and/or Juncus articulatus in a 10x10 cm square centred on the position of a Spiranthes romanzoffiana plant at site KA (sample of 120) and KB (sample of 101) on 3-7 October 2001. The number of 10x1 0cm subdivisions that were occupied by Juncus articulatus and/or Juncus acutiflorus x articulatus in a 50x50cm quadrat which was centred on the stem of Spiranthes romanzoffiana was recorded in a sample of 13 at the unmonitored site TG on Colonsay. Capsules The nature of the capsules of Spiranthes romanzoffiana was examined at the August, September and October visits. Five plants of Spiranthes spiralis in fruit were examined in Cumbria, England on 28 October 2002. Plants at the same site were photographed on 18 October 2002 by Mr David Benham. RESULTS Population sizes A standard method in plant surveying involves a count of inflorescences on a single visit per year. Single visit data are presented in Table 1. In Table 1 'Flowering plants' includes those with stems partly grazed or grazed to ground level but still discemable. The number of plants with stems on 8 August 2002 and on 27 - 29 July 2003 at site KC was 1 , 1 and at site LF was 3, 1 respectively. The number of plants in all three states (vegetative, flowering and bud- only) from the ‘monitored set’ for 2002 and 2003 for KC was 1 1 and 14 and for LF was 22 and 17. Furthermore at LF on 27 May 2003 26 plants (from the ‘monitored set’ of 37) were recorded. The number of buds on 23-24 September 2003 at sites KC and LF was 13 and 20 respectively. Thus the count of plants in bloom was small in two populations which actually contained a moderate number of plants. The presence of 1 1 plants in 2002 in the set of 1 8 monitored positions at KC and 14 in 2003 demonstrates the tolerance of the species to heavy grazing regimes producing low swards, (see section ‘habitat characterisation’). The number and percentage of flowering plants at new positions is shown in Table 1. In 1999 all the plants at site KB, KC, LF and SF were detected from the presence of flowering stems and this was the case for 49 of the 54 plants at KA. As this was the first year of recording all were ‘new’ plants i.e. at previously unknown positions. For 2000 and 2001 known positions include vegetative, flowering and bud-only plants. Active searching for and recording of vegetative and bud-only plants ceased in 200 1 . Thereafter only plants which flowered in a previous year were added to the set which was used to determine if a plant in bloom was a new plant or not. Hence the number of plants added each year to the cumulative total from 1999 will be less in the post 2001 system. This change is indicated by the superscript b in Table 1. The percentage of new flowering plants falls over the first three years at all sites except KC. This fall in the early years is a consequence of the methodology. Thereafter the percentage remains high at sites KA and KB with summer grazing breaks, and low at KC with no summer grazing breaks in 2001, 2002 and 2003. The percentage is also low from 2002 onwards at LF with light grazing levels. Using only values of flowering plants from a single annual visit, the cumulative number of records rises progressively from 1999 to 2003 at sites KA and KB, (Table 1). At least one pair of twin mature plants occurred at each of the five sites. During the period 1999 - 2001 the cumulative number recorded over all visits within one year, in all above-ground states (vegetative, flowering and bud-only), rose at all sites, (Table 1). An extrapolation of these trends would suggest that at all sites populations exist which are larger than the all-visits-within-year figure for 2001. This is highly likely to be the case for KA, with values of 122, and 37 new flowering plants for 2002 and 2003; and for KB with 41 and 20 (single visit data. Table 1). The fact that plants can spend up to six years in the underground state indicates that individuals can be quite long lived. The status of the lowest stem leaf Table 2 shows the incidence of slug and vertebrate grazing at the leaf at the base of the stem at three sites in August 2002. The high incidence of vertebrate truncated leaves at the two exclosures is due to grazing in the spring prior to the gates being closed. At this time the upper part of these leaf blades were above ground, but the stems and inflorescences (sheathed in bracts) were below ground. On Barra at site BB subsite A the level of winter and spring grazing was less and this is reflected in a low percentage of vertebrate grazed leaves (18%), conversely the level of slug grazing is higher (34%) than that at the other two sites. The maximal number of leaves per plant occurs when the stem is fully developed. However the upper stem leaves are only exposed to grazing when 38 the stem attains its foil length. The leaves which ultimately become the lower stem leaves appear above ground before the stem itself. The results suggest that the status of the lower stem leaves may be used as an index of spring grazing levels, especially at sites with a summer grazing break i.e. where the stem is not subject to heavy grazing by stock. Grazing on stems Sites/years with a summer grazing break Within the exclosure at site KA the number of flowering stems bearing inflorescences in a sample of 21 on 18 July 2001 declined progressively; 10 being present on 7 August, 4 on 14 August, 1 on 22 August and 0 on 2 September. These plants were located adjacent to a rabbit {Oryctolagus cuniculus) burrow and rabbit grazing is assumed to be the cause of the loss of stems. At site KC in 2000 stock were absent from 1 June to 4 August. One plant was in bloom on 3 July, the flowering stem had been grazed by 17 July, i.e. before the stock were re-introduced. Five forther plants were in bloom on 26 July. By 14 August the original plant had its stem grazed right down and one plant had no stem left. By 3 1 August two had truncated stems, one had the only two basal leaves visible and for three there was no trace of the above ground parts of the plant. Sites/years with no summer grazing break In 2001 site KC was subject to more or less continuous grazing. Appendix A. The balance of intact to partly or wholly grazed or trampled flowering stems changed from 7:2 on 18 July to 3:6 on 29 July and 0:9 on 7 August. The values include a diminutive flowering stem and proto-inflorescence only 14mm tall in total, present on 29 July but grazed by 7 August. On the 27 August there was no trace of the above ground parts of one of the plants. In 2002 and 2003 at KC the stem present had been grazed by the time of the late July/early August visit (Table 1). General The results from site KC emphasise the importance of regular recording and of the use of marker pegs without which the basal leaves and the remaining base of the stems (when present) might not have been observed. At many sites/subsites, on five Hebridean islands, individual stems have been located that have been cut through in the irregular manner associated with slugs (members of the order Stylommatophora); and slugs have sometimes been observed on the stems themselves. Habitat characterisation Spiranthes romanzoffiana plants in bloom were found at a wide range of vegetation heights. Table 3. Of special importance are the results from within the exclosures at sites KA (n=164) and KB (n=56) on Colonsay on 9 - 13 August 2002. The populations at these two sites were ungrazed for June, July and part or all of August from 1996 onwards (details in Appendix A): and hence members of the two populations had survived over a number of years in conditions in the summer months broadly similar to those present in 2002. The values refer to the grass and sedge rich i.e. first vegetational layer and exclude the Juncus articulatus and Juncus acutiflorus x articulatus layer. At site KC the sward height around the positions of 1 8 members of the monitored set on 8 August 2002 was low, median 25mm; mean value 24mm, and uniform. Coefficient of Variation 17%, (Table 3). No species of the genus Juncus were present, although the site is wet during the winter months, their absence apparently being due to the high level of grazing. KC is subject to the same grazing regime as the outside of the exclosures at KA and KB, Appendix A. Site BB subsite A on Barra has no summer grazing and light to moderate winter grazing. As at sites KA and KB (within the exclosures) Spiranthes romanzoffiana was found at a wide range of vegetation heights, the median and mean heights being 70mm and 84mm respectively, (Table 3). Site LF was lightly grazed in 2002; the mean and median vegetation heights in August 2002 are actually greater than those within the exclosures, ungrazed in June, July and August, (Table 3). Vegetation height and height of Juncus articulatus and J. acutiflorus \ articulatus in 2002 and 2003 at the positions of Spiranthes romanzoffiana plants at sites with a summer grazing break. For measurements paired in time, the vegetation height at the positions of Spiranthes romanzoffiana plants at BB subsite A and KA was very highly significantly lower in 2003 than in 2002, (Table 4). For the non paired measurements the vegetation height (of the grass and sedge rich layer) at the positions of Spiranthes romanzoffiana plants in the exclosures at KA and KB was not significantly lower in 2003 than in 2002, (Table 4). In these two cases the vegetation height at the positions of all detected Spiranthes romanzoffiana plants in bloom in the relevant year was recorded; (in 2002 n=164 for KA, n= 56 for KB; in 2003 n=57 for KA, n=24 for KB: exclosures ungrazed by stock). The height of Juncus articulatus and J. acutiflorus X articulatus plants up to the point where the stem and leaves become less dense was highly significantly less for KA in 2003 and very highly significantly less for KB in 2003, non paired data, (Table 4). The dry summer of 2003 appears to be linked to the lower vegetation height at BB subsite A and KA for paired data: and for heights of Juncus articulatus and J. acutiflorus x articulatus at KA and KB, Table 4. Conceivably it is partly responsible for the lower number of plants in bloom at these three sites in 2003 compared with 2002, (Tables 1 and 4). 39 J uncus articulatus and J uncus acutiflorus x articulatus There was a 60% frequency of Juncus articulatus and/or Juncus acutiflorus x articulatus in 10x1 0cm quadrats at KA; and a lower frequency (28%) at KB: the presence/absence ratios being very highly significantly associated with site, Chi = 23.21, 1 df. (Table 3). As indicated by number of plants in bloom the population of Spiranthes romanzoffiana at KA is larger than at KB, (e.g. 164 inflorescences at KA on 9-12 August 2002; 56 at KB, Tables 1, 3 and 4). These two Juncus taxa are absent from the nearby site KC. Both taxa were present at the whole site level at KA, KB, LF and also BB subsite A (Table 3). Both taxa occur at exclosure site ten Gulliver et al. (2004b, this volume) on the margins of the diffuse site SF. Further studies on the balance of this these two taxa on a quadrat basis at ten exclosure sites are presented in Gulliver et al. (2004b, this volume). At the unmonitored site TG on Colonsay the median number of 10x1 0cm subdivisions in a 50x50cm quadrat that were occupied by Juncus articulatus and/or Juncus acutiflorus x articulatus was 12, the range being 1 to 22. The quadrat was centred on the stems of 13 Spiranthes romanzoffiana plants on 19 September 200 1 . The cumulative number of plants that flowered at TG in 2001 was 33. Site TG is summer and winter grazed. There are no obvious indications that the observed plants of Spiranthes romanzoffiana are being adversely affected by the two Juncus taxa, but only long-term studies will detennine the matter absolutely. The samples taken at site KB on 7 October 2001 which had been ungrazed since 30 May included an example of Juncus articulatus (with capsules) that was 9cm tall and an example of Juncus acutiflorus X articulatus (with capsules) that was 10cm tall; these dwarf plants representing one end of the great range of variation in size that can occur in the two taxa. The current authors found Juncus acutiflorus x articulatus at the two Spiranthes romanzoffiana sites where they carefully examined the Juncus species on Coll. Henderson (2001) reported a 20% frequency of the taxon that she recorded as Juncus articulatus at 81 10x10 cm sampling stations centred on Spiranthes romanzoffiana plants on Coll. Pearman and Preston (2001) do not record Juncus acutiflorus x articulatus from Coll. Rich and Jenny (1998) describe Juncus acutiflorus x articulatus as ‘probably under-recorded’. Changes in vegetation height over time Table 5 shows the change in vegetation height over time for the five sites including the paired values on each side of the exclosure fence at sites KA and KB in 2001. In 2001 sheep and cattle were excluded from 30 May onwards at KA and KB, and hence no separate inside values for 16 May 2001 were recorded. 18-20 September 2002 values are also presented. The low May values indicate the intensity of winter and early spring grazing. For site KA wet and dry the ungrazed height values increase progressively from June to October. Maximum values are shown in bold in Table 5. The nature of the grazed wet sward at KA on 19 September 2002 is shown in Figure A5. The measured i.e. first vegetation layer on the ungrazed side is masked by Juncus spp. (mainly Juncus acutiflorus x articulatus) plants, which appear to be more dense on the photograph than is the case on the ground. For site KB dry ungrazed a peak is reached in mid-late August with declining values thereafter. For KB wet ungrazed there is a peak in early August with lower but somewhat erratic values thereafter. Further aspects of vegetation height and its ecological significance at sites KA and KB are presented in Appendix C. Site KC, was more or less continuously grazed in 2001 and 2002 (see Appendix A and B) by the same stock as the outside sward of KA and KB and the height values are similar to the dry outside values at KA and KB. On the visit to KC in July 2003 tillers of Molinia caerulea and Nardus stricta and parts of Calluna vulgaris plants were seen on the ground, apparently having been pulled up by stock. Dead tillers of Molinia caerulea had also been seen on the sward surface on previous visits. Site LF was lightly and intennittently grazed in 2001 and 2002 (see Appendix B) and its vegetation height values resemble the inside wet and dry swards of KA, but with a drop in September 2001 due to grazing which commenced on 29 August 2001. Site SF was grazed more or less continuously, but with varying intensities. The moderately high vegetation height indicates that the plant stations studied were grazed to an intermediate level of intensity. Vegetation height will vary with factors which change throughout the season. These include the rate of growth of the sward, changing palatability of plant species present, the intensity of grazing due to varying numbers of stock, and stock preferences for particular zones within the grazed area. Vegetational height will be affected by local differences in soil conditions within the total area, these differences become fiilly expressed in the absence of grazing. The vegetation height for the dry section of KB ungrazed is consistently lower than the wet section. However the dry' and wet sections of KA are similar. Differences within the wetness category are shown by comparisons of KA ungrazed with KB ungrazed. The BCA dry values are considerably higher than the KB dry values. Similarly the KA wet values are considerably higher than the KB wet ones. 40 Observations on Spiranthes wmanzoffiana and Spiranthes spiralis plants in August, September and October Fully-developed, dry capsules with longitudinal slits have not been observed by any of the authors in the study period, 1999 - 2003; e.g. during the survey on 3 to 5 October 2001 of the 39 plants with surviving stems at KA and the 19 at KB in 2001, Gulliver (2002). The observed course of development is always that the capsules develop part way and then become flaccid, turning from green to brown and then shrivel. The capsules of Spiranthes spiralis dry in the fully expanded state in Cumbria, see Figure A6 (photograph kindly supplied by Mr David Benham). The shrivelled capsules of Spiranthes romanzoffiana of plant A4 at site GF on Coll on 1 October 2002 are shown in Figure A3 (photograph kindly supplied by Ms Emma Grant); and in Figure 1C in Gulliver et al. (2000) for plant 2 from site LF on Colonsay on 15 September 99. It was therefore not possible to examine the performance of plants of Spiranthes romanzoffiana in swards and grazing regimes of different types by comparing aspects of sexual reproductive output at different locations. This technique was used by Legg, Cowie and Sydes (1997) who showed a 49% loss of scapes to sheep and rabbit grazing in Primula scotica in 1993 and a 42% loss in 1994 using caged plots. Our observations are confined to the state of the capsules. They do appear to lend support to the experiences of Robarts (2000) who states T have however yet to be convinced that Spiranthes romanzoffiana in Barra is in fact setting seed’. DISCUSSION Detection of populations The existence of a summer grazing break allows Spiranthes romanzoffiana plants to flower and hence their presence to be detected. The number of plants observed in bloom at sites KA and KB has risen from the values of 49 and 24 in 1999, (Table 1, single visit data) to 164 and 56 respectively in 2002. In 2003 the values were 57 and 24, (Table 1). At both sites new plants represented 65% or over of the plants in bloom, 1999 to 2003. At sites KC and LF, with no exclosures, the number of plants in bloom in 2002 were 1 and 3 and in 2003 1 and 1. Nevertheless the numbers present at the ‘monitored set’ of positions for 2002 were 1 1 at KC and 22 at LF; and for 2003 were 14 for KC and 17 for LF. At these two sites moderate-sized populations were in existence, despite the very low numbers of plants in bloom. Thus number of inflorescences is not always a useful guide to population size. Even with a summer grazing break, number of plants in bloom can vary greatly. Forty-four were in flower at BB subsite A in 2002, five in 2003, single visit data, (Table 4). Reproduction The non-production of expanded capsules on any of the plants examined at the reference sites emphasises the need for more investigation on the presence and extent of sexual and vegetative reproduction in the species, ideally under both natural and controlled conditions. Habitat characterisation, vegetation height, effects of stock presence The survival of plants at a range of vegetation heights, measured in August 2002, indicates that the individuals of Spiranthes romanzoffiana exhibit a fairly broad band of tolerance with regard to this habitat characteristic. The populations at sites KA (n=164) and KB (n=56) had been ungrazed for June, July and part or all of August from 1996 to 2002 (for details see Appendix A; from 2001 the exclosure plots were established). Thus there has been a seven-year period for competitive interactions between species to exert their influence. The results from site BB subsite A, subject to more or less the same grazing regime every year, show the same trend, (Table 3). Spiranthes romanzoffiana populations also exist at a range of vegetation heights at sites with no summer grazing break, e.g. site LF. The survival of the moderate-sized population at Site KC indicates the potential of the species to tolerate regimes of more or less continuous grazing producing low, uniform swards, at some sites at least. The number of buds on 23 September 2003 in the monitored set of positions at site KC was 13, suggesting a continuation of population of a similar size in 2004. Differences in vegetation height on either side of exclosures against summer grazing at heavily grazed sites reveal the impact of the grazing process and provide an index of the potential biomass in the absence of grazing. Continuous grazing is inevitably associated with continuous trampling and defecation activity. The possible positive, negative or neutral effects of these two features of stock presence have yet to be investigated. Defoliation results in loss of the photosynthetic tissues, but potential competitors may also be affected. Trampling will result in tissue damage but may also result in root fracture which could conceivably be a form of vegetative reproduction in the Orchidaceae (Rasmussen, 1995, p.l29); fiirthennore in marshy areas, patches with high levels of light penetration are created as small pieces of vegetated sod are pushed into the substrate. Dung deposition by cattle will result in the total loss of light to the plant for a period. However the organic carbon source from cattle and sheep dung may well ultimately be of importance to the fungal associate. It would be possible to study all these topics in the field; but more precise results would be obtained from controlled experimental manipulations in botanic gardens, if material were 41 available and the relevant conservation organisations approved such a course of action. Many Spiranthes romanzoffiana sites in Scotland are beside lochs, and the plant may obtain temporary relief from grazing during periods of high water following heavy summer rains. Grazing at such locations during dryer periods may ensure general habitat suitability for Spiranthes romanzoffiana. At the smaller subsite of LF, near the loch; three individuals were under water on 15 September 1999, but all produced vegetative plants in 2000. Several of the Irish lough-side sites are ungrazed or very lightly grazed F. Horsman, D. Lupton, N. Kingston; {pers. comm.). Hence there may be factors other than grazing at such sites which are advantageous to the species, e.g. the perturbation of the marginal zone by wave action and/or the deposition of silt and humus during periods of high water levels. J uncus articulatus and J uncus acutiflorus x articulatus Plants of Spiranthes romanzoffiana can occur growing with stems of Juncus articulatus and/or Juncus acutiflorus x articulatus, e.g. see Tables 3 and 4. At sites which are not heavily grazed they appear to gain a measure of protection from consumption by the presence of these less palatable (but not inedible) plants. The long-tenu fate of Spiranthes romanzoffiana individuals growing in stands of Juncus articulatus and/or Juncus acutiflorus x articulatus is yet to be investigated. Recording site conditions and the longevity of Spiranthes romanzoffiana For any given study site stock may be moved on and off at regular or irregular intervals or very rarely moved; and the number of livestock plus the balance of species of livestock may change over time. In addition perceived management objectives may alter; and sites may be subdivided by fencing or expanded by allowing access through gates, or by fences becoming functionless. The position of temporary features e.g. winter feeding stations may vary from week to week or year to year or be constant. Furthenuore sand blow, fire, local flooding and draining activities plus other site- specific biotic or abiotic factors may affect vegetation and patterns of grazing. All these changes may have conservation significance and it is important to record them. Only with these data can one interpret population changes. Appendix A is an example of the presentation, in table fonuat, of changes to some aspects of site conditions for a ten- year period. Future ecological analysis requires extensive records, however such detailed data may be difficult to condense and present in a meaningful summary fonn. The fact that Spiranthes romanzoffiana can occur in the underground fonn for up to six complete years (Dr James Robarts, pers. comm.) makes long-term study of both the species and its environment especially important, i.e. seven years must elapse after the initial observation before one can assume that a plant at a marked position has died. Monitoring populations Fully developed capsules have not yet been observed in Scotland. Six populations (B-G) of marked individual plants on Barra have been monitored by Dr James Robarts since 1993 and one (A) since 1992, e.g. see Gulliver (2002); and populations of unmarked plants have been studied since 1988 (Robarts 2000). One subsite of site WC on Coll has supported plants since 1978 (Ferreira 1978 record in Scottish Natural Heritage files and current authors’ visits of 2000 and 2001). It therefore seems likely that vegetative reproduction may be very important for the maintenance of populations of the species. Other rare species maintained wholly or largely on vegetative reproduction in Scotland are Carex chordorrhiza and Saxifraga hirculus (Legg, Cowie and Sydes 1997); together with Linnaea borealis, Cicerbita alpina and Saxifraga cernua (Wilcock 2002). The current studies reveal an absence of any annual, readily-quantifiable measure of sexual reproductive output, e.g. the number of mature capsules. (If present measurements of the total weight of fiilly developed seeds per capsule; or an estimate of total number of seeds based on samples from different parts of the ovary wall; would ideally additionally be recorded). Therefore other measures will have to be used to investigate yearly performance and importance or otherwise of intensity of grazing. These include recording the expansion or contraction of population size, once the majority of existing plants have been recognised. Such a technique requires a very long period of monitoring of pennanently marked and mapped plants. This approach has been developed to a high degree by Dr James Robarts on Barra (Robarts 2000; and the analyses of Dr Robarts’ data 1992-2001 in Gulliver (2002) for the seven populations (A-G) which have a summer grazing break and light winter grazing and the two which have some grazing suinmer and winter (H-I). The technique accommodates scenarios where the populations never produce seed; only produce seed occasionally; where fertility barriers exist which vary in their effectiveness from year to year; and where heavy grazing results in the high or total loss of inflorescences. It could therefore be very usefully applied to study the variation in population performance e.g. at sites subject to different grazing regimes. Ideally the sites should be ones where the full range of abiotic and biotic factors present are also being recorded 42 ACKNOWLEDGEMENTS Kind thanks are extended to following organisations, individuals and funding bodies. The owners, farmers and erofters of the sites for permission to make these observations; the staff of the Seottish Natural Heritage offices at Islay, and South Uist; and the Enquiry Service of English Nature and Publications Section of Scottish Natural Heritage. Dr James Robarts and Dr Frank Horsman for sharing their experience of the species. Mr David Benham, Dr Neil Cowie, Professor Michael Crawley, Dr Mike Dodd, Mr Alan Forrest, Ms Verena Forster, Dr Mark Gardener, Ms Emma Grant, Mr Nigel Grant, Dr Mark Hill, Dr Naomi Kingston, Dr Marilyn Light, Mr Alex Lockton, Mr Darach Lupton, Ms Rae McKenzie, Mr John McKinnell, Mrs Wendy Nelson, Mr Philip Oswald, Professor Clive Stace, Dr Robin Sutton and Dr Christopher Wilcock. Scottish Natural Heritage for funding the two research projects and the Botanical Research Fund. The administrators of the Professor Blodwen Lloyd Binns Bequest for a grant towards attendance at a British Ecological Society conference at which results of the investigations were displayed. 43 REFERENCES Bowler, J. (2003) Irish Lady’s-tresses (Spiranthes romanzoffiana) discovered on the Isle of Tiree, Argyll. BSBI News - the newsletter of the Botanical Society of the British Isles 92, 25-26. Blackstock, T.H. and Roberts, R.H. (1986) Observations on the morphology and fertility oiJuncus x surrejanus Druce ex Stace & Labinon in north-western Wales. Watsonia 16, 55-63. Cheffings, C. (2004) New plant status lists for Great Britain. BSBI News - the newsletter of the Botanical Society of the British Isles 95, 36-43. Diack, I.A., Burke, M., and Peel, S. (2000) Grassland management for nature conservation: towards a consistent approach to sward measurement and description. Pp. 155-156 in Rook, A. J. and Penning, P.D. (editors.) Grazing Management - British Grassland Society symposium No. 34. British Grassland Society, Reading, Berkshire:. Forrest, A. (2001) Genetic structure within and among populations o/Spiranthes romanzoffiana Cham, in Scotland. M.Sc. Thesis. Royal Botanic Garden Edinburgh and University of Edinburgh. Forrest, A.D., Hollingsworth, M.L., Hollingsworth, P.M, Sydes, C., and Bateman, R.M. (2004) Population genetic structure in European populations of Spiranthes romanzoffiana set in the context of other genetic studies on orchids. Heredity 92, 218-227. Grime, J.P., Hodgson, J.G. and Hunt R. (1986) Comparative plant ecology: a functional approach to common British species. Unwin Hymen, London Gulliver, R.L. (1996) The status of Spiranthes romanzoffiana Cham. (Orchidaceae) Irish Lady’s Tresses, on Colonsay, (v.c. 102) in 1995; with special reference to associated plant communities. Watsonia 1\ (2), 202-204. Gulliver, R.L. (1997) Irish Lady’s Tresses {Spiranthes romanzoffiana) on Colonsay, (VC 102). The Glasgow Naturalist 23 (2), 55-56. Plus brief note in The Glasgow Naturalist 23 (3), 77. Gulliver R.L. (2002). Dr Richard Gulliver action plan research into the ecology o/Spiranthes romanzoffiana in Scotland, viii & 124pp. Unpublished report to Scottish Natural Heritage. Gulliver, R., Gulliver, M., Keimen, M., and Sydes, C. (2004b). Studies on the conservation biology of Irish lady’s-tresses orchid, Spiranthes romanzoffiana 2) The establishment of 10 exclosures, dung counts and further studies on associated Juncus taxa (species and hybrid rushes). The Glasgow Naturalist. Current volume. Gulliver, R.L., Gulliver, M. and Sydes, C. (2003) The relationship of a Biodiversity Action Plan (BAP) orchid, Spiranthes romanzoffiana, to grazing in the West of Scotland. Aspects of Applied Biology 70: Crop quality: its role in sustainable livestock production. Published by the Association of Applied Biologists c/o Horticulture Research International, Wellesboume, Warwick CV35 9EF, UK. (This conference had a section on biodiversity) pp. 143-150. Gulliver, R., Keimen, M., Gulliver, M. and Sydes, C. (2000) Observations on Irish lady’s-tresses orchid {Spiranthes romanzoffiana) on Colonsay (vcl02). The Glasgow Naturalist 23 (5), 9-12. Henderson, S.A. (2001) The vegetation associated with Spiranthes romanzoffiana Cham. (Orchidaceae), Irish Lady’s-tresses, on the Isle of Coll, Inner Hebrides. Watsonia 23, 493-503. Legg, C., Cowie, N., and Sydes, C. (1997) The importance of regeneration studies to successful conservation management of Scottish rare plants. Botanical Journal of Scotland A9 , 425-432. MacDonald, A., Stevens, P., Armstrong, H., Immirzi, P., and Reynolds, P. (1998a)^ guide to upland habitats: Surveying land management impacts. Volume I . Background information and guidance for surveyors. Scottish Natural Heritage, Edinburgh. MacDonald, A., Stevens, P., Armstrong, H., Immirzi, P., and Reynolds, P. (1998b) A guide to upland habitats: Surveying land management impacts. Volume 2. The field guide. Scottish Natural Heritage, Edinburgh, f Mannetje, L. (1987) (editor) Measurement of grassland vegetation and animal production. Commonwealth Bureau of Pastures and Field Crops, Hurley, Berkshire. Pearman, D.A., and Preston, C.D. (2001) ,4 flora of Tiree, Gunna and Coll. Privately published by the authors, Dorchester. Preston, C.D., Pearman, D.A. and Dines, T.D. (2002) New Atlas of the British and Irish Flora. Oxford University Press, Oxford. Rich, T.C.G. and Jermy, A.C. (1998) Plant Crib 1998. Botanical Society of the British Isles, London. Robarts, J. (2000) A study of the orchid Spiranthes romanzoffiana on the island of Barra. Hebridean Naturalist 13, 30-35. Rasmussen, H.N. 1995. Terrestrial Orchids from Seed to Mycotrophic Plant. Cambridge University Press, Cambridge. Roberston H.J. and Jefferson R.G., (2000) Monitoring the condition of lowland grassland SSSIs. I English Natures ’s rapid assessment method. English Nature, Research Report No. 315, Peterborough. Stace, C.A. ( 1 997) New Flora of British Isles, Second Edition. Cambridge University Press, Cambridge. Stewart, A., Peannan, D.A. and Preston, C.D. (eds.) (1994) Scarce plants in Britain. Joint Nature Conservation Committee, London. Stewart, K.E.J., Bourn, N.A.D. and Thomas, J.A. (2001) An evaluation of three quick methods commonly used to assess sward height in ecology. Journal of Applied Ecology 28,, 1148-1154. UK Biodiversity Action Group (1999) Tranche 2 Action Plans Volume 3. Plants and fungi. English Nature, Peterborough. Wilcock, C.C. (2002) Maintenance and recovery of rare clonal plants: the case study of the twin flower {Linnaea borealis). Botanical Journal of Scotland 54, 121-130. 44 Table 1. Number of flowering plants (see general note) of Spiranthes romanzoffiana recorded on a single summer visit and the number and percentage which had not previously been recorded as flowering (F) and vegetative (V ) and bud-only (B) plants for 1999 to 2003; plus cumulative number of V, F and B plants recorded over all visits in any one year for 1999, 2000 and 2001; on Colonsay. Single yearly visit data - F plants Multiple visit data F, V and B plants Site Summer Date of single Number of Number Percentage 1999-2003 Within year Year for letter grazing visit - flowering F never F never cumulative cumulative cumulative code break sometimes ering plants previously previously number F value within year and see App- each subzone (F) record- recorded as recorded as based only flowering. visit totals num- pendix A visited ed. Max. F, V or B F, V or B on single vegetative. ber sequentially value in plants. plants. visit data & bud-only bold LF(1) N 21/08/99 8 8 100 8 8 1999 LF(1) N 13/08/00 14 13 93 21 27 2000 LF(1) N 04/09/01 8 4 50 25 32 [37“] 2001 LF(1) N 08/08/02 3 F 33" 26 LF(1) N 27/07/03 1 O'- 0" 26 KA (2) Y 14/08/99 49 49 100 49 54 1999 KA (2) Y 15/08/00 43 38 88 87 127 2000 KA (2) Y 30/08/-06/09/01 113 76 67 163 213 2001 KA (2) Y 9-12/08/02 164 122'’ 74" 285 KA (2) Y 28/07/03 57 3T 65" 322 KB (3) Y 14/08/99 24 24 100 24 25 1999 KB (3) Y 14/08/00 32 31 97 55 86 2000 KB (3) Y 28/08/01 67 53 79 108 142 2001 KB (3) Y 13/08/02 56 41" 73" 149 KB (3) Y 28/07/03 24 20" 83" 169 KC (4) Y 26/07/99 1 1 100 1 1 1999 KC (4) Y 14/08/00 5 4 80 5 7 2000 KC (4) N 26-29/07/01 9c 8 89 13 17118''] 2001 KC (4) N 08/08/02 1“ 0" 0" 13 KC(4) N 29/07/03 1'' 0" 0" 13 SF (5) N 04/08/99 3 3 100 3 3 1999 SF (5) N 25/07/00 2 1 50 4 4 2000 SF(5) N 25/07/01 0" 0 4 618“] 2001 SF(5) N Not surveyed SF(5) N Not surveyed' Notes General: 'Flowering plants' include those with stems partly grazed or grazed to ground level but still discemable. " For LF, KC and SF the values in square brackets are the totals of F, V and B plants pooled over all visits in all years, 1999 - 2001, i.e. for LF and KC the subsequently 'monitored set'. In 2002 and 2003 additions to the cumulative number recorded consist only of plants flowering in that year, recorded on a single visit, i.e. there is an alteration to the basis on which the number and percentage new plants in bloom is calculated. Includes one miniature plant with diminutive stem and proto-inflorescence, total height 14mm. ** Flowering stem grazed a time of late July/early August visit. ' One plant was in bloom on 4 September. ^ An area on the margin of the diffuse site SF with a 1996 plant record became plot ten, Gulliver etal. 2004b. 45 Table 2. The grazed or withered"' status of the lowest stem leaf in August 2002 at sites KA and KB exclosed since 1 June in 2002; and at Site BB area A; not subject to summer grazing. Percentages are rounded to the nearest whole number and hence in two cases appear to add up to 101. Sites and Dates Leaf Status KA 07-08/08/02 Number Percentage KB 07/08/02 Number Percentage BB subsite A 14/08/02 Number Percentage Ungrazed 35 22 19 34 10 23 Vertebrate truncated 100 63 30 54 8 18 Slug truncated and/or damaged 16 10 6 11 15 34 Withered^ 9 6 1 2 11 25 Total 160" 56 44 Notes ■' So badly withered that the end of the leaf could not clearly examined or was absent. Slightly withered leaves were allocated to one of the other three categories. 160 was the count during a general survey on 7-8 August. Between 9-12 August every plant was mapped, and the count was 164, see Table 1. 46 Table 3. Frequency of values of vegetation height measurements in the vicinity of Spimnthes romanzoffiana plants in August 2002 at one site subject to more or less continuous grazing, two exclosed sites, one site with no summer grazing; and one lightly grazed site: plus other sward details. Height values for the layer made up of Juncus species, present at some sampling stations at KA, KB and LF, are not included. Site KC KA KB BB sub-site A LF“ Grazed/ Exclosed Well Exclosed Exclosed No Summer Lightly Grazed Grazing Grazed^* Date 8/8/02 9-12/8/02 13/8/02 14/8/02 8/8/02 Vegetation height of the sward - 5nim steps between 20 and 30mm at KC: 10mm steps at other sites Median value shown bold 20 25 30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 180 190 5 12 13 0 0 14 0 4 37 5 14 31 13 2 30 17 3 26 13 4 4 3 2 10 4 1 3 0 5 2 1 2 2 0 1 0 0 2 2 0 1 1 0 0 2 Median height Mean height Coefficient of Variation No. of samples Number of plants with flowering stems on date of visit 25 60 24 67 11 32 18*’ 164 L 164 70 72 20 56 56 70 84 49 44 44 % of flowering stems which are newly recorded plants 0 74 73 36 0 1 5 4 10 4 2 1 0 0 1 0 80 80 23 28"^ 3“ (33f % occurrence on 3-7/10/01 at sites KA, KB and KC of Juncus artkulatus and J. acutiflorus x articulatus in a 10x1 0cm square centred on the position of a Spiranthes romanzoffiana plant Percentage 0 60 29 No. of sampling stations 18 120 101 Chi square presence/absence of Juncus species for sites KA/KB very highly significant Chi = 21.63, 1 df Presence on entire site (2001 and 2002) Juncus articulatus . + + + + Juncus acutiflorus x articulatus - + + + + Notes ^ Site LF was lightly and intennittently grazed in 2002. The height values refer to main population centred on the five plants first recorded in 1999, (as also in Table 5). In 2002, one plant in bloom in the main population, two in the sub-population. 33% new plants refers to whole site. ’’ For sites KC and LF these are positions of plants recorded 1999 to 2001, not necessarily above ground plants at the time of the height measurements. Stem was grazed. 47 Table 4. Vegetation height and height of Jimcus articiilatus and J. acutiflorus x articulatus at the positions of Spiranthes romanzoffiana plants in late July/early August 2002 and 2003 at sites with a summer grazing break. 2002 2003 Signific Mean Median Number Number Mean Median Number Number -ance of flower- of flower- of2002 to values ing values ing 2003 single single compar date date -isons (two tailed) Barra (site BB) and Colonsay (site KA) paired measurements; n is different from number flowering in 2002 Site BB subsite A 81 65 38" 44 63 50 38"*’ 5 VHS 14/8/02 & 8/8/03 z=7.37 Site KA 67 60 30" 164 52 50 30bc 57 VHS 9-12/8/02 & 28/7/03 z=3.76 Colonsay non paired measurements; n is both sample size and number flowering (single visit) Site KA 67 60 164‘‘ 164 67 60 57"* 57 NS 9-12/8/02 & 28/7/03 z=0.09 Site KB 72 70 56"* 56 65 60 24"* 24 NS 13/8/02 & 28/7/03 z=1.55 Jiinciis articulatus and J. acutiflorus x articulatus height (see text), Colonsay; non paired measurements At positions of Spiranthes romanzoffiana plants Mean Median Sampling Total Mean Median Sampling Total stations number stations number with sampl- with sampl- Juncus ing Jimcus ing spp. stations spp. stations SiteKA 291 300 [67]" [96]" 242 260 53 57 HS 9-12:8:02 & 28:7:03 z=3.27 Site KB 223 200 27 56 142 140 20 24 VHS 13:8:02 & 28:7:03 z=7.69 Notes HS Highly Significant, VHS Very Highly Significant, NS Not Significant a) 3 1 negative differences, 7 zero differences, 0 positive differences. i.e. there are no differences in values which mn opposite to the main trend. b) All were at positions of plants in bloom in 2002. c) 22 negative differences, 3 zero differences, 5 positive differences, i.e. there were 5 differences which nm opposite to the main trend. d) 100% sampling of plants in bloom in 2003. e) 96 positions sampled iox J uncus spp. within entire set of 164 plant stations. 48 Table 5. Overall median of vegetation heights in mms (maximum values shovi^n bold) between May and October 2001 and on 18-20 September 2002 at two sites with grazed and ungrazed sections and dry and wet parts plus one well-grazed, one lightly grazed and one variably grazed site. Height values for the layer made up of Jimcits species, present at some sampling stations at KA, KB, SF and LF, are not included. NR = Not Recorded For details of the configuration of the sampling stations see Appendix C Site KA dry KA dry grazed ungrazed 0 I (outside) (inside) n = 10 n=10 Differe- ence of medians KB dry KB dry grazed ungrazed 0 I (outside) (inside) n=10 n=10 Differe- ence of medians KC heavily grazed n = 3/18‘‘ LF^ Lightly grazed n= 10 SF Variable intensities of grazing n = 3 Values for KA dry & KB dry 2001 (KA & KB dry ungrazed 30 May - 7 October) May 15 see 0 value [0] 10 see 0 value 40] 15 mid June 15 25 10 15 25 10 late June 20 35 15 15 28 13 15 early July 50 50 mid July 15 48 33 15 38 23 40 73 45 early August 28 85 57 25 50 25 30 100 80 mid-late August 30 95 65 28 58 30 30 90 80 early September 25 110 85 23 50 27 25 100 55 late September 18 110 92 20 50 30 20 50 October 10 140 130 10 45 35 20^' 2002 (KA & KB dry ungrazed 1 June - 25 September) 18-20 September 20 120 100 20 55 35 20 105 NR Values for KA wet & KB wet Day of Month KA wet KA wet Differe- KB wet KB wet Differe- KA/KB LF SF grazed ungrazed ence of grazed ungrazed ence of /KC 0 I medians 0 I medians see notes (outside) (inside) (outside) (inside) c & d n= 10 n= 10 n= 10 n= 10 2001 (KA & KB wet ungrazed 30 May - 7 October) May 10 see 0 value [0] 10 see 0 value :[0] 1676'* - mid June 15 33 18 23 35 12 137-'* - late June 20 53 33 18 45 27 28 early July 5 5 mid July 30 55 25 20 45 25 12 19 18 early August 30 85 55 28 85 57 7 2 2 mid-late August 35 90 55 35 73 38 22 14 14 early September 38 95 57 35 63 28 6 4 4 late September 28 100 72 30 60 30 20 22 October 20 120 100 20 70 50 673'* - 2002 (KA & KB wet ungrazed 1 June - 25 September) 1 8-20 September 25 120 95 30 80 50 18'*/ 19'= 20 20 Notes “ Median of 18 sampling points on 3 October, otherwise median of 3 sampling points. ’’ Main subsite. For KA and KB. For KC. 49 Appendix A. Details of the different management phases at Sites KA®, KB^", KC*’ within a dune system (i.e. on the seaward side of the 1996 moor/dune fence) and TG‘^ (immediately on the 'moor' side of the 1996 moor/dune fence) on Colonsay from 1991 (when observations by RG and MG began) to 2003. See Appendix B for further site details Year or period Phase Sites Period of grazing by sheep and cattle Period of 1 signifies more or less affected summer grazing continuous grazing, break 2 signifies a summer grazing break 1991 - 1995 - No moor/dune fence, no summer grazing break 1991 - 1995 Phase 1 A KA, KB. more or less continuous** KC, TG 1996 - Moor/dune fence established, with gates 1996 - 1999 Phase 2A KA, KB, 15 August - 3 1 May (following year) KC 1996- 1999 Phase IB TG more or less continuous** Similar to 1 A but with the moor/dune fence and gates, hence stock either a) all on the moor or b) all on the dunes or c) in both areas 2000 Transition Phase KA, KB, up to 3 1 May 2000, more or less KC continuous after 4 August 2000 none 31 May - 15 August none 31 May -4 August 2000 Phase IB TG more or less continuous** none 2001 - Exclosures*^ at KA and KB established in the spring 2001, 2002 &Phase2B KA, KB 2003 similar to Phase 2A but with a longer summer grazing break 2001, 2002 & Phase IB KC 2003 up to late May/early June and From late after late September/early October May/early June to late September/ early October more or less continuous**; grazing none heaviest when all stock are on the dunes only (for conservation management for chough) i.e. when moor /dune gates are shut and stock are on the seaward side 2001, 2002 & Phase IB TG more or less continuous** but with no none 2003 grazing when all stock are on the dunes only (for conservation management for chough) Notes General: The exclosures have 6 sides at site KA and 5 at KB and are irregular in shape. “ Sites KA and KB are in zones within the dunes which are wet partly due to the impervious nature of the underlying rock, see also Appendix B. *’ Sites KC is on a slope, but the sandy substratum is periodically wet, especially in the winter. *" Site TG is in damp gully with a bum in the zone between moorland and dunes; the substrate includes windblown sand. ** With some breaks e.g. when sheep are dipped *' Some plants occur outside the fence at sites KA and KB. Of the 49 and 25 plants recorded in bloom in 1 999 at KA and KB (all visits), one is outside the 2001 exclosure at KA and one at KB. 50 Appendix B. Characteristics of the study sites. Slug activity has been recorded from all sites Site Site Site Grazing Summer Adjudged Slope. code number Unenclosed vertebrate. grazing intensity Different used in (extensive) Most break of zones may Gulliver =UE abundant grazing “ have et al. or Enclosed first different 2000 (a field) =E characteristics Colonsay LF 1 E*’ C & S, R, G N L F KA 2 UE S&C, R Y H F, GS KB 3 UE S&C,R Y H VGS KC 4 UE S&C,R Y/N^ H MS SF 5 UE S&C,R N V F (mainly) VGS (in part) TG UE S&C,R N M(V) BSG Barra BB/A UE c&s Y L F Wetness LF Adjacent to loch KA Wet in part, dry in part. Pegs and plants may be pushed deep down into the substrate in the marshy parts 1 stock KB Centre of the very shallow hollow is wet, very slight downslope water movement KC Some downslope water movement in winter SF Mainly next to a bum; part of site slopes gently to bum TG Gully bottom wet, bum present, sides dampish due to impeded drainage BB / A Surface water movement in wet periods Abbreviations: The code for the most abundant animal is always presented first. BSG = Bottom and sides of a gully, C = Cattle, E = Enclosed site (field), F = Flat, G = Geese, GS = Gentle Slope, H = Heavy, L = Light, M = Moderate, MS = Moderate slope, N = No, R = Rabbit, S = Sheep, UE = Unenclosed (extensive), V = Variable, VGS = Very gentle slope, Y = Yes Notes General; Further details in Gulliver (2002). “ And associated animal activity e.g. trampling and defecation. ’’ Whole field very approximately 3 ha in extent. ^ Summer grazing break in 1999 and 2000 not in 2001 and subsequently. 51 Appendix C. Vegetation height: further details. Aspect of the definition The definition of vegetation height utilised herein i.e. to the point where the density of the component stems and leaves of the sward diminished appreciably, is very similar to the one used by English Nature to assess the status (e.g. conditions favourable/unfavourable) of Sites of Scientific Interest (Robertson and Jefferson, 2000), in which height is ‘assessed as the average height rather than the extreme of flowering spikes of grasses and tall herbs’. Devices used to measure vegetation height Trials were conducted on Colonsay on the measurement of height using a 10cm sward disc, also known as a sward plate, (Diack, Burke and Peel, 2000). The presence of many ungrazed and grazed stems of Juncus spp. caused the plate to lie at an angle or raised it above the main sward surface, and so the technique was not used subsequently. Investigations on the relative merits of direct measurements of vegetational height (as used in this study), use of a sward disc of 30cm diameter (both 10 and 30 cm discs are in current use) and the Hill Farming Research Organisation’s sward stick, carried out on chalk grassland in southern England, are reported in Stewart et al. (2001). Each has certain advantages and certain demerits, the direct measurement technique was found to be less strongly affected by vegetation structure than the other two systems. Configuration of samples KA and KB 2001 and 2002, wet and dry; n = 10; arrangement - linear - details as text below KC 2001, n = 3 or 18; arrangement at the position of Spiranthes romanzoffiana plants: KC 2002, 30; arrangement - a 10 x 12m plot LF 2001, 10; arrangement - a rectangular frame: 2002, 30; arrangement - a 10 x 12m plot SF 2001, n = 3; arrangement - at the position oi Spiranthes romanzoffiana plants At sites KA and KB data on vegetation height were gathered in spatially paired measurements 1 m inside and 1 m outside of the exclosures from 13 June to 6 October 2001 and on 19 September 2002. At both sites ten pairs of values were taken along one fence line where the soil was wet, with standing water at or very near the surface during wet periods; and ten pairs along another fence line where dry soil was present. Figure A5 shows the whole fence line used for the wet comparisons and the first part of the fence line used for the dry comparisons on 6 October 2001. Further aspects of vegetation height at different parts of KA and KB in 2001 and 2002 For site KA wet and dry the difference between the inside and outside medians increases progressively throughout 2001 (Table 5). For site KB wet and dry the maximum vegetation height measurements inside the exclosure occur in August. For the wet set of values this produces the highest differences between inside and outside in August. For the dry set the outside sward was very low in October, and this resulted in the greatest difference between inside and out occurring in October. The 19 September 2002 values broadly resemble the 6 October 2001 values, except for the KA dry set values which have some affinities with 20 September 2001 and some with 6 October 2001. Allowing for some seasonal variation, these results suggest a fair degree of similarity for early autumn values between the two years. The wet and dry zones at sites KA and KB all had the same substrate (wind blown sand mixed with varying quantities of mildly acidic humus) and the separation between KA and KB was less than 400m. Nevertheless there were major differences in the vegetation height when comparing the same ungrazed types between the two sites (i.e. KA wet ungrazed with KB wet ungrazed, and KA dry ungrazed with KB dry ungrazed). Height differences between apparently comparable sites (e.g. KA and KB dry ungrazed) may well reflect differences in sward biomass and provide an index of the level of competition for light amongst the sward components. 52 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 53-68. STUDIES ON THE CONSERVATION BIOLOGY OF IRISH LADY’S-TRESSES ORCHID, SPIRANTHES ROMANZOFFIANA; 2) THE ESTABLISHMENT OF 10 ENCLOSURES, DUNG COUNTS AND FURTHER STUDIES ON ASSOCIATED JUNCUS TAXA (SPECIES AND HYBRID RUSHES) Richard Gulliver’, Mavis Gulliver^ Margaret Keirnen^ and Christopher Sydes^ ' and ^ to whom correspondence should be addressed, Carraig Mhor, Imeravale, Port Ellen, Isle of Islay, Argyll PA42 7AL, ^ Grianan, Drumclach, Isle of Colonsay, Argyll, PA61 7YR, Scottish Natural Heritage, 2 Anderson Place, Edinburgh, EH6 5NP ABSTRACT Exclosures were established at ten sites in the Hebrides (six on Colonsay, three on Barra and one on Vatersay) at sites in two groups. Group A: where plants of Spiranthes romanzoffiana (a Biodiversity Action Plan species) had been recorded in previous years, but were not located in 2002; to ascertain if individuals were still present. Group B: where 1 - 2 plants were observed in 2002, and where it was suspected that the cryptic population was larger. The locations of the exclosures are referred to by code numbers to reduce the likelihood of visits by collectors of rare orchids. 100 Ixlm quadrats were examined inside the exclosures for plants of Spiranthes romanzoffiana at three dates in 2003. On the same visits 100 Ixlm quadrats were examined outside the exclosures. Vegetation and site features were recorded inside and outside. At two of the 2003 exclosure sites (pooled), seven newly recorded plants were found outside the exclosure, one newly recorded plant was found inside and five newly recorded plants within 100m of the exclosures. At the remaining eight sites no new plants were discovered. The thirteen newly recorded plants were made up of eight vegetative plants and five flowering plants. Within each of two of the exclosures one plant, present in 2002, occurred above ground in 2003. At a third, highly-disturbed site a plant in 2003 outside the exclosure was in the approximate position of one recorded in 2002 and 1996. Combining new and known plants, inside and outside, four sites had Spiranthes romanzoffiana plants above ground in 2003. The total number of plants (pooled over all exclosure sites, inside and outside) which were last recorded in the period 1990-1996, but which were not detected in 2003, was eighteen. The number recorded in 2001 but not in 2003 was eight; and the number recorded in 2002 and not in 2003 was two. The two exclosure sites with the greatest number of detected plants in 2003 were in the middle of the range of values of combined counts of sheep and cattle dung, which can be used as an indicator of grazing activity. These two exclosure sites had the lowest percentage sheep dung as a percentage of sheep and cattle dung and the lowest percentage sheep hoof holes as a percentage of sheep and cattle hoof holes in late May/early June, inside and outside pooled. The frequency of Juncus articulatus and Juncus acutiflorus x articulatus combined in 1 x Im quadrats varied from 6 to 100. Juncus articulatus frequency as a percentage of the frequency of Juncus articulatus or Juncus acutiflorus x articulatus and Juncus articulatus varied from 5 to 100. Studies on the status of the small populations at and near the reference sites will continue in 2004. Monitoring of exclosure sites in general is facilitated if the vegetation is internally uniform and is comparable between locations. Key words or phrases: conservation, dung, exclosures, grazing, management, Juncus articulatus INTRODUCTION Spiranthes romanzoffiana (Irish lady’s-tresses orchid) is a white flowered orchid (Figure Bl) which blooms in July and August, (Gulliver et al. 2000). It can frequently be observed growing amongst the stems of Juncus articulatus and Juncus acutiflorus x articulatus {Juncus x surrejanus). It is currently a ‘Nationally Scarce’ species sensu Cheffmgs (2004), i.e. Scarce sensu Stewart, Pearman and Preston (1994) and a Biodiversity Action Plan Priority Species, see UK Biodiversity Action Group (1999). The number of plants recorded in large and medium sized study populations, the relationship to grazing, the height of vegetation at and near plant stations, and the results of observations on capsules are presented in Gulliver (2002), Gulliver et al. (2003) and Gulliver et al. (2004a, this volume). The species can spend up to six years in the underground state, (Dr James Robarts pers. comm.). The number of plants in bloom is not a good indication of population size Gulliver et al. (2004a, this volume). Flowering stems are grazed by stock, and the vegetative plant is hard to detect. It is therefore very difficult to ascertain the true status of a population occurring at a site subject to grazing by stock. Gated exclosures can be used to create a summer grazing break. This generates maximum visibility of the plants that are present; and allows grazing in the autumn, winter and spring to ensure sward consumption for much of the year. Ten were established in March 2003 as part of a Scottish Natural Heritage research project at sites with small populations in two groups. Group A: where Spiranthes romanzoffiana plants had been recorded in previous years, but were not located in 2002. Group B: where 1 - 2 plants were observed in 2002, and where it was suspected that the cryptic population was larger. Allocation to groups A and 53 B was based specifically on presence in 2002 and the plant(s) being present in the exact site of the future exclosure. Sites six and nine had plants present inside the position of the subsequent exclosure in 2001 but not 2002. At site five in 2002 no plant was present within the position of the ftiture exclosure; but one did occur in the equivalent study area outside the boundary of the future exclosure. The aim of the research was to determine if the populations with no 2002 records were still extant; and if larger populations existed where 1 - 2 plants were detected in 2002. The break from grazing by stock during the summer months was provided to improve the possibility of plant detection. In addition, factors which might affect Spiranthes romanzoffiana e.g. intensity of grazing and the attributes of the site, were investigated. METHODS Marking the position of individual plants of Spiranthes romanzoffiana Individual plants were marked by the use of an adjacent black plastic peg(s) with a ‘mushroom shaped’ black top. The marker peg(s) was/were 10cm from the plant. Either 1, 2 or 4 pegs were used. Referencing systems were established to permit plants to be relocated. Any one plant may either be in the flowering (F), vegetative (V) or underground (U) phase in any one year. The underground status can only be applied after a minimum of three years study, e.g. F in year 1, U in year 2, V in year 3. Members of the Orchidaceae are maintained by mycotrophy while in the underground state, Rasmussen (1995). At certain positions peg sinkage and loss occurred. These included locations where the ground was very soft, where it was very marshy and where there was a high level of disturbance to the substratum by stock, sometimes these features occurred in combination. The use of marker pegs commenced in 1999. Exclosures site details There has been a history of rare orchids being periodically dug up in the British Isles. Therefore the locations of exclosures are not identified in detail, but are referred to by code numbers. Three of the ten exclosures were on Barra (sites one - three), one on Vatersay (site four) and six on Colonsay (sites five - ten). The characteristics of the sites and the dates of the original records of Spiranthes romanzoffiana from the sites of the current exclosures, and from the associated sets of 100 quadrats outside the exclosures, are given in Appendix A. Nine of the exclosures were ten metre squares, inside which 100 1 x Im quadrats were studied in late May/early June 2003, late July/early August and late September/early October 2003. The dates of the visits are given in Appendix B. Due to topographic constraints at site five an 8 x 13m exclosure was used; data are presented for 100 of the 104 quadrats for comparative purposes. In addition the proposed positions of the exclosures were studied in 17-20 August 2002 on Barra and Vatersay and 20-24 September 2002 on Colonsay. Each exclosure had two 2.4m gates which faced each other. These gates were closed at the end of the late May/early June visit and opened at the end of the late September/early October visit. The facing gates allow through visibility, which will help to ensure stock are not discouraged from entering and grazing during the winter months. At each site 100 Im x Im quadrats were studied outside the exclosures. These were located first and foremost in positions where it appeared possible that Spiranthes romanzoffiana might exist. The local micro topography frequently resulted in strong variation in the degree of wetness and vegetation structure over very short distances inside the exclosures. The second criterion for determining the position of the outside set of quadrats was to replicate the variation inside as far as this was possible. The micro topographic features included shallow bums, rock outcrops, and ditches, Appendix A. In addition stands of e.g. Calluna vulgaris, and Juncus effusus added to the variety of the stmcture of the vegetation. Site five provides a particularly good example of this heterogeneity. It is bisected by a very shallow bum, hence there are some bum and bumside quadrats both inside and outside. Figure B2. A track used by stock (but no longer used by humans) passes through the outside set of quadrats at right angles to the bum, resulting in much trampling and disturbance, especially adjacent to the bum. However comparatively dry substrates are present at the quadrats which are most distant from the bum, inside and out. Thus the study plots do not resemble the more or less homogeneous stands that would be selected for the direct measurement of stand/grazing pressure interactions e.g. as utilised by Welch (1984), albeit at a larger scale, for dung studies. The outside quadrats were either in a single block; or in two, three or four blocks, (Appendix A). At site four the outside quadrats were in a block of 80 with a contiguous extension of 21 quadrats. At site five there were 104 quadrats outside the exclosure (as well as 104 inside the exclosure). In both cases 100 quadrats from the larger number was used for comparative purposes. Every quadrat was carefully examined for the presence of Spiranthes romanzoffiana leaves. The frequency of a range of plant species including Calluna vulgaris (heather) was recorded on the late July/early August and/or late September/early October 2003 visits. Assessment of grazing intensity For each exclosure site the adjudged overall grazing intensity was assessed on the following scale, H = Heavy, L = Light, M = Moderate, V = Variable, VH = Very Heavy, VL = Very Light. The assessment integrates a) changes throughout the year (2003) 54 and b) the effects of localised small areas of very light or very heavy grazing. Counts 'were made of the number of hoof holes per quadrat and of the number of items of sheep, and cattle dung per quadrat. Presence/absence of rabbit dung was noted. Vegetation height (Gulliver et al. 2004a, this volume) was measured by first taking 10 height measurements within the 1 x Im quadrat and then selecting the median value of this single sample (first order median). Vegetation height was recorded in 5mm steps between 10 and 30mm and in 10mm steps thereafter. Vegetation height was measured to the point where the density of the component stems and leaves of the sward diminished appreciably. The data gathered allowed comparisons to be made within different subsections of the same site, between sites, and over time. Data from each site are summarised in the form of medians and means. As the initial values are in 10mm steps, the medians routinely vary by values of 1 0mm. This coarse scale may mask differences that are apparent in the mean values, which are presented to 1mm values. When interpreting these means it is important to bear in mind that the original data were gathered in 10mm steps. Within the Ixlm quadrats more than one vegetation layer was often present. Data were gathered on the height and percentage cover, in 5% steps, of the one, two or three layers present as appropriate. The height band of each layer was broadly similar throughout, so in non grazed (or extremely lightly grazed) quadrats there would be no first layer. Juncus articulatus and Juncus acutiflorus x articulatus The relative balance of Juncus articulatus and Juncus acutiflorus x articulatus {Juncus x surrejanus) was recorded in terms of frequency, i.e. presence/absence per quadrat in 2002 or 2003. Juncus acutiflorus x articulatus was identified using the characters given in Blackstock and Roberts (1986), Stace (1997) and Rich and Jermy (1998), with particular attention being paid to the length of the ripe capsule in relation to the perianth segments and to the colour of the capsule. Sometimes the quadrat contained specimens where all the inflorescences were grazed, and taxonomic determination was not possible. Plants of Juncus acutiflorus x articulatus are partially fertile and produce limited quantities of viable seed, Blackstock and Roberts (1986); the taxon chiefly reproduees by far creeping rhizomes. By contrast Blackstock and Roberts (1986) state that the rhizome system in Juncus articulatus is subcaespitose or shortly creeping. Rhizome length will determine the openness of stands of the two taxa, indicating the desirability of recording the balance of the taxa present. Juncus articulatus and Juncus acutiflorus x articulatus varied in abundance in the quadrats from occasional plants in the grass and sedge rich sward to quite dense stands, especially inside the exclosures in ditches and at the edge of the lochan (at site six). The plant of Juncus articulatus or Juncus acutiflorus x articulatus with the height nearest the middle value in each quadrat was selected and the height, up to the point where stem and leaves thinned out markedly, measured in 20mm steps. The Ellenberg indicator values (British version), Hill et al. (1999) for Juncus articulatus for Reaction (preference for soil pH), Nitrogen and Moisture are 6, 3, 9: the corresponding values for 7. acutiflorus being 4, 2, 8; indicating that J. acutiflorus is more associated with acid, nutrient poor soils than is Juncus articulatus. Values for the hybrid are not provided. The values for Spiranthes romanzoffiana are 6, 4, 8. Capsules The nature of the capsules of Spiranthes romanzoffiana was examined at the late September/early October visits. RESULTS Number of plants within the 2003 exclosure sites and the associated set of 100 Ixlm quadrats A summary of the results is presented in Table 1. Plants found in 2003 are indicated in bold and their V/F status is stated. Details of the sites are given in Appendix A. The dates of the visits are provided in Appendix B. The newly located plants were all found at two sites, five and nine, (Table 1). At the remaining eight sites no new plants were diseovered. At these two sites pooled, seven newly located plants were found in the 100 1 x Im squares outside the exclosure: one plant was found within the exclosure and five plants within 100m of the exclosures, (one of which is shown in Figure Bl). The thirteen newly recorded plants were made up of eight vegetative plants and five flowering plants. In 2002 plants had been recorded from the subsequent positions of the exclosures in three instances. Within one exclosure (at site four) one of the two 2002 plants had above ground growth in 2003; within one exclosure (at site seven) the 2002 plant did not produce above ground growth in 2003, and within one exclosure (at site eight) the single 2002 plant had above ground growth in 2003, (Table 1). Site five has a highly disturbed zone (Figure B2) adjacent to the bum, where pegs sink and sections of sward are pushed do-wnward or sideward by cattle, located within the set of 100 quadrats outside the exclosure. The plant recorded in 2003 was in the same broad position as one in 2002 and 1996, (Table 1). In the outside set of 100 quadrats at site seven one plant had no above ground tissue in 2003. It had flowered in 2001. Combining new and known plants, inside and outside, four sites had Spiranthes romanzoffiana plants with above ground gro-wth in 2003, (Table 1). 55 The total number of plants (pooled over all exclosure sites; inside and outside) which were last recorded in the period 1990-1996, but which were not detected in 2003, was eighteen. The number recorded in 2001 but not in 2003 was eight; and the number recorded in 2002 and not in 2003 was two. Vegetation height in late July/early August at sites with Spiranthes romanzoffiana in 2003 To facilitate comparisons between sites the height of a single general level of vegetation was measured at sites in late July/early August 2003, (excluding the zone created by the Jimcus spp.). Spiranthes romanzoffiana plants occurred at sites five, eight and nine (pooled) at low e.g. 30mm and at relatively tall e.g. 130mm, vegetation heights, (Table 2). The mean values for the entire set of quadrats (n=100) were always very highly significantly greater inside than outside the exclosures. The great variability of height values at sites five and nine, caused by the range of micro topographic features and the variety of vegetation types, has had the consequence that the coefficient of variation was high at these two sites; in three cases it exceeds 40, (Table 2). Whereas the outside sets of quadrats were laid out over terrain that was approximately equivalent to the inside set, it is possible that an element of the inside/outside difference is due to physical and/or vegetational differences as well as an absence of grazing. The objective in selecting the positions of the outside quadrats was to maximise the possibility of locating Spiranthes romanzoffiana plants. At site four one plant with above ground growth occurred inside the exclosure. It was on a very small mound in the slow flowing bum; first vegetation height of quadrat 80mm, second vegetation height 1 10mm. There was no dung in the hundred quadrats outside the exclosure at site four in early June and none in early August 2003, indicating minimal grazing at this site. Dung and Hoof holes Dung values at sites one to ten in late May/early June - i.e. immediately before the exclosure gates were closed Table 3 shows the dung counts (sheep and cattle combined) totalled over all 100 1 x Im quadrats inside and outside the exclosures in late May/early June 2003. The mean of the two sets of values and the rank are also shown. The gates were closed at the end of the visit to each site, hence these values are linked to the same stock densities in the relevant area as a whole. At site eight the upper layers of the substrate were fluid and dung deposited was quickly amalgamated with the mixture of water, peaty humus and mineral material. Hence no reliable values were obtained for site eight. As a result the ranks run from 0 to 9. The ranking largely accords with general observation, site ten (rank 2 in Table 3) appeared to be the most heavily grazed overall and site four (rank 9 in Table 3) the most lightly grazed. The high count of dung at site six (rank 1 in Table 3) may be partly influenced by stock coming to the edge of the lochan to drink and/or graze the emergent vegetation. At sites one, three, six, seven and nine the dung counts were considerably higher outside the exclosure even though the gates were open, (Table 3). For sites three, six, seven and nine the first vegetation height measurements indicated more grazing activity outside. The mean height of the first level outside and inside being; 33 and 39mm, 41 and 56mm, 37 and 43 mm and 42 and 61mm respectively. (Means are quoted to 1mm: the data were gathered in 10mm steps: n ranges from 44 to 100 - not all quadrats had a first vegetation layer). At site ten the dung count was lower outside the exclosure; the first vegetation height measurements, 36 and 29mm respectively, indicated the same trend. At site five the dung count was 36 outside and 42 inside the exclosure; the height measurements here are fairly similar to each other at 34 and 37mm respectively. Greater animal activity will result in higher dung counts and lower vegetational heights. Local features may exert an influence on the difference between the inside and outside counts in Table 3. The outside part of site three was bisected by parallel sheep paths. At site one the high outside values occurred near to the fence. Although this had only been present for circa two months, it may have been favoured as a rubbing area, such a utilisation was very conspicuous at site six, later in the year. To accommodate localised high concentrations of dung, it would be possible to exclude the top quartile of the values when comparing sites. The two sites at which new plants were detected, sites five and nine, are midway in the range of values, both being rank 4.5, (Table 3). As well as being an indicator of grazing activity, dung counts may also show the extent of one of the sources of humus. It is conceivable that the supply of organic carbon reserves from the fungal associate of Spiranthes romanzoffiana plants is greater at sites with medium to high levels of dung input. Rasmussen (1995, p.98) states ‘Little is known about the distribution of orchid endophytes in nature, but it is assumed that they are widespread saprophytes, and some are possibly parasites or form other mycorrhizal associations as well.’ Further infonuation on the biology of the fungi associated with Spiranthes romanzoffiana, both in the free living and mycorrhizal states, would be desirable. Frequencies per 1 00 quadrats were calculated. The ranks of the means of the inside and outside frequeney values were exactly the same as the ranks of the total counts. As there is more scope for differences to be expressed using counts compared with frequency values (which in this case can only have one of 100 values; or 200 when inside and outside values are pooled), counts have been used in this account. 56 Hoof hole counts Conditions within the exclosiires and the set of 1 00 Ixlm quadrats varied from dry, peaty soils supporting Calluna vulgaris to wet, marshy zones; edges of small lochans; ditches; and small bums. A comparison of the hoof hole counts and dung counts showed that at one extreme there was firm ground which would not register hoof holes whilst dung counts were possible; ana the other there were moderately wet zones where dung persisted but hoof marks were absent and marks made in wet substrate would soon cease to exist (as the material resumed its original shape). At site eight the substrate was so fluid that dung was frequently amalgamated into it. Over all sites hoof hole counts per 100m' could not be used for assessing the intensity of grazing, but they were used to examine the balance of types of stock present. Balance of sheep and cattle present as indicated by dung and hoof holes The percentage dung due to sheep was similar for sites six (rank 4) and ten (rank 5); (Table 4). In addition the percentage hoof holes due to sheep was similar for sites six (rank 5), and ten (rank 4). Overall the rankings for percentage sheep in the two measures in Table 4 show good agreement but the effect of the three values from the Barra sites, all of 1 00%, should be noted. The two sites (five and nine) with the highest count of Spiranthes romanzoffiana plants had the greatest percentage of dung and hoof holes being due to cattle and the lowest percentage being due to sheep dung. Hence there is a suggestion that the type of grazing animal may be an important site factor. Lang (1989) reports the observation made by J. Raven that the greatest number of flowering spikes appear in areas where cattle have been fed in the previous winter, implying that disturbance of the soil may be beneficial to the species. Seasonal changes in dung counts Table 5 shows the total counts for the two sites at which new plants were detected, sites five and nine, in late May/early June (M/J), late July/early August (J/A) and late September/early October (S/0) 2003. Inside the exclosures no new dung is being added after the late May/early June count and values decline as the humus decays or is washed into the ground. At site five the values decline progressively outside the exclosures, though to a somewhat lesser degree compared with inside. This suggests that the grazing pressure has been strong during the late winter and spring; and decreases in the summer, possibly as a result of stock moving to alternative grazing areas in the summer. At site nine outside there is no change in the component of the dung value due to cattle from late May/early June to July/early August, but a small increase in value for sheep dung. Overall dung inputs and breakdown seem to be approximately in equilibrium for this period. The value falls in late September/early October, as at site five. Castle and MacDaid (1972) found that dung pats produced by dairy cows on two replicate plots of a low N application and two of a high N application disappeared in 114 days, there was no significant difference between the treatments. However dung ‘disintegrated’ (= disappeared) in 133, 131, 109 and 79 days (overall mean 113) when deposited in mid May, late May, June and July respectively, the differences being highly significant. They do not attribute a cause to the observed results. The average area of the dung pats was found to be 580cm': a voiding rate of 12.5 per day is provided in their discussion, but not in the results. Welch (1985) gives an average area of 61 1cm' and Marsh and Campling (1970) give a voiding rate of 12.0 per day. It is possible that there is a higher rate of decay of the organic matter in the summer when temperatures are somewhat higher. On the other hand higher winter rainfall levels may be important for carrying elements of the dung pat or dung pellet into the substrate. Climatological data for the period 1951 to 1980 for Colonsay is provided by Clarke and Clarke (1991). Juncus articulatus and Juncus acutijlorus x articulatus Abundance and frequency of the taxa Juncus articulatus and Juncus acutiflorus x articulatus pooled varied in frequency from six at site two (inside) to 100 at sites seven, eight and ten (inside and outside), (Table 6). Site ten had a very high adjudged grazing level and the second highest value for dung counts, (Table 6). Juncus articulatus was the only taxon in the Barra and Vatersay plots, but Juncus acutiflorus x articulatus did occur near to the outside quadrats at site four. Quadrats containing Juncus articulatus as a percentage of quadrats containing Juncus articulatus or Juncus acutiflorus x articulatus and Juncus articulatus in plots five to ten in late September 2002 (before the erection of the exclosures) varied from 5% to 100% (Table 6). Both taxa were present at all these six plots. All the plots on Barra and the Vatersay plot were in positions exposed to strong winds from the sea. The two plots on Colonsay with the highest percentage of Juncus articulatus were the two nearest to the shoreline; they also appeared to be the most exposed of the six. Height in late July/early August at sites with Spiranthes romanzoffiana in 2003 At the quadrats containing Spiranthes romanzoffiana at sites five, eight and nine (pooled) the height of Juncus articulatus and Juncus acutiflorus x articulatus (measured in 20mm steps to where stems and leaves thinned out markedly) ranged between 100 and 200mm, mean 156mm, n = 10. The mean heights for all quadrats inside at sites five, eight and nine were 176, 205 and 264mm, (n = 95, 100 and 97 respectively, see Table 6). The 57 height outside, for plants which had no obvious signs of having been grazed by stock, were 168, 149 and 222, n = 98, 100, 94 respectively. Juncus articulatm was not present at the quadrat position of Spiranthes romanzoffiana at site four; the overall mean height inside was 202mm, n = 13 (August 2003). Grazing Grazing by rabbits was observed at the distal ends of Juncus articulatus and Juncus acutiflorus x articulatus stems in some of the exclosures in late July on Colonsay, and grazing by slugs at other exclosures. Sometimes both types of grazing occurred. At heavily grazed sites Juncus articulatus and Juncus acutiflorus x articulatus can have grazed stems of approximately the same height as the rest of the grazed vegetation. At parts of very lightly grazed sites they may be ungrazed. Frequently a range of heights of both grazed and ungrazed plants was present in each quadrat. The two taxa (combined) might be used for an assessment of the intensity of recent grazing, utilising the lowest grazed stem height present in each quadrat. Observation indicates a rapid regrowth of stems following grazing, assuming no repeat grazing. Jumping plant louse Livia juncorum galls The jumping plant louse Livia juncorum modifies the inflorescences of Juncus species to form a tassel gall (Redfem and Shirley, 2002). This gall can occur at ground level as well as higher up the stem. At plots one, two and three the frequency was three, zero and nine respectively in August 2002. No data were collected at plot four. On Colonsay in September 2002 site nine had two quadrats with the gall, all other sites had no quadrats with the gall, though the gall is not uncommon on the island. The Colonsay sites were generally more heavily grazed than the Ban'a and Vatersay sites. Floristic indications of variation between sites The frequency of Juncus effusus, Calluna vulgaris and of Sphagnum species (pooled) is shown in Table 7. Juncus effusus is rank position 1 and 2 (mean of outside and inside) at sites three and ten. It is considerably less frequent at the other sites. Calluna vulgaris is most frequent at site nine followed by sites four and five, it has frequency values over 40% at all three of these sites. It shows a positive association with Erica tetralix, the Speamian rank correlation coefficient is 0.87. Sites one to three on Barra, site four on Vatersay and site nine on Colonsay have mean frequencies of 60% or above for Sphagnum species. Sites six and seven, on wind blown sand, have low or zero frequencies of Calluna vulgaris and Sphagnum spp. The first and second rank positions for these three study species, shown bold in Table 7, occur at six different sites: this is one indication of the variability between sites. Capsules All the examined capsules of Spiranthes romanzoffiana were slirivelled. None had expanded. Full capsule expansion and natural drying of the capsule wall was observed in Spiranthes spiralis in England, in October 2002, Gulliver et al. (2004a, this volume), but has not been known to occur in Spiranthes romanzoffiana in Scotland. DISCUSSION Reproduction and distribution of plants The non-production of expanded capsules on any of the plants at the reference sites Gulliver et al. (2004a, this volume), and also at the exclosure sites, emphasises the need for more investigation on the presence and extent of sexual and vegetative reproduction in the species. The presence of the five plants (at two stations pooled, sites five and nine) within 100m of the exclosures suggests that at some sites the species may be very thinly distributed. Detection of plants In the set of quadrats inside or outside the exclosures 73% of plants (both newly located and previously known) were vegetative. (The plants within 100m of the exclosures have not been included in this data set as they were only located because either they were in bloom or were immediately adjacent to plants in bloom). These findings emphasise the need for careful examination of the sward in order to assess the presence of Spiranthes romanzoffiana plants. A detailed search strategy of this kind is implemented at the exclosure sites. There is a marked contrast between the number of plants detected in the unfenced set of quadrats (n = 8, seven newly located plants) and in the exclosures (n = 3, one newly located plant). The exclosures and therefore the inside quadrats were located around the position of previously recorded plants. The outside quadrats were adjacent, but positive efforts were made to select positions for the one to four blocks of quadrats where it was felt likely that Spiranthes romanzoffiana plants might occur. Surveys in 2004 will help to establish whether this difference in spatial distribution is genuine, i.e. whether or not there are approximately equal numbers within and outwith the exclosures. Juncus articulatus and Juncus acutiflorus x articulatus Juncus articulatus and Juncus acutiflorus x articulatus can tolerate heavy grazing e.g. at sites six and ten, as can Spiranthes romanzoffiana (Gulliver et al. 2003, Gulliver et al. 2004a, this volume). The evidence from the exclosures and from the reference sites (Gulliver et al. 2004a, this volume) indicates that at the present grazing regimes, both species appear to be co-existing, but long term studies are needed to verify this interpretation of the situation. The reduction of the biomass of the two Juncus taxa due to grazing may result in lower levels of competition with adjacent species for e.g. light and nutrients. In addition 58 grazing of Jimciis articiilatiis tissue may limit the development of dense tufts, groups of which may form a barrier for the emergence of underground plant parts of other species. Hoof hole counts and dung deposition The ranking of combined sheep and dung counts shows a positive relationship with adjudged grazing intensities. Dung data are more rapid to gather than measurements of vegetation height, especially where more than one vegetation layer exists, each layer itself commonly requiring a cover estimate as it frequently does not occupy the full area of the quadrat. The results indicate that dung counts appear to be informative as a measure of grazing intensity. However local features such as drinking areas, comparatively sheltered areas, fence posts and fence lines (and the presence of barbed wire), and sheep paths may provide high counts. For four of the sites, variations which existed between values from the inside of the exclosure (before the gates were closed) and values from the outside, were reflected in differences in vegetation height of the first vegetation layer. The percentage of hoof holes that were due to sheep was strongly linked to the percentage dung produced by sheep. For general studies of grazing activity at the sites which had a good registration of hoof holes of sheep and cattle, it may well be possible, after conducting pilot studies, to generate a regression line of dung counts on hoof hole counts. This would then allow the impact of either of the two factors on the sward to be estimated from one set of values. Impact of grazing animals Four of the ways in which the presence of grazing animals may affect Spiranthes romanzoffiana positively are a) suppression of potental competitors, b) dung deposition adding to the stock of humus in the soil and favouring relatively high rates of organic matter breakdown generally; which may result in enhanced heterotrophic activity by the fungal associate of Spiranthes romanzoffiana plants, c) trampling being associated with root breakage and possible movement of root fragments by the hooves of cattle and sheep, d) the creation of open areas by stock movement and trampling, these having high levels of light availability. Grazing will have a negative effect by removing leaves, stems and inflorescences. Trampling will damage tissue of Spiranthes romanzoffiana and dung deposition will blanket out the light for a period. Site monitoring, including dung counts, and studies on the interaction between a variety of biotic and abiotic factors will continue in 2004. This monitoring will generate further infomiation on the conservation biology of Spiranthes romanzoffiana. ACKNOWLEDGEMENTS a) Groups and organisations; the owners, farmers and crofters of the sites; Scottish Natural Heritage; the Botanical Research Fund; the Professor Blodwen Lloyd Binns Bequest Fund of the Glasgow Natural History Society: Scottish Executive Rural Affairs Department. b) Individuals, Dr Neil Cowie; Mr Alan Forrest; Ms Verena Forster, Dr Mark Gardener; Ms Emma Grant; Mr Jonathan Grant, Mr Nigel Grant; Dr Mark Hill; Dr Frank Horsman; Dr Marilyn Light; Mr Alex Lockton; Mr Darach Lupton, Ms Rae McKenzie, Mr Philip Oswald; Dr James Robarts; Professor Clive Stace; Dr Robin Sutton. REFERENCES Blackstock, T.H. and Roberts, R.H. (1986) Observations on the morphology and fertility of Jimciis x siirrejanus Dmce ex Stace & Labinon in north-western Wales. Watsonia 16, 55-63. Castle M.E, and MacDaid, E. (1972) The decomposition of cattle dung and its affect on pasture. Journai of the British Grassland Society 21, 133-137. Clarke, P.M. and Clarke, J. (1991) The flowering plants of Colonsay and Oransay. Printed by the Harlequin Press, Oban, Argyll: privately published by the authors. Cheffmgs, C. (2004) New plant status lists for Great Britain. BSBI News - the newsletter of the Botanical Society of the British Isles 95, 36-43. Gulliver R.L. (2002). Dr Richard Gulliver action plan research into the ecology of Spiranthes romanzoffiana in Scotland, viii & 124pp. Unpublished report to Scottish Natural Heritage. Gulliver, R., Gulliver, M., Keimen, M., and Sydes, C. (2004a). Studies on the conservation biology of Irish lady’s-tresses orchid, Spiranthes romanzoffiana 1) Population sizes, grazing, vegetation height and capsule status at reference sites. The Glasgow Naturalist. Current volume. Gulliver, R.L., Gulliver, M. and Sydes, C. (2003) The relationship of a Biodiversity Action Plan (BAP) orchid, Spiranthes romanzoffiana, to grazing in the West of Scotland. Aspects of Applied Biology 70; Crop quality^: its role in sustainable livestock production. Published by the Association of Applied Biologists do Horticulture Research International, Wellesboume, Warwick CV35 9EF, UK. (This conference had a section on biodiversity) pp. 143-150. Gulliver, R., Keimen, M., Gulliver, M. and Sydes, C. (2000) Observations on Irish lady’s-tresses orchid {Spiranthes romanzoffiana) on Colonsay (vcl02). The Glasgow Naturalist 23 (5), 9-12. Hill, M.O., Mountford, J.O., Roy, D.B., and Bunce, R.G.H. (1999) Ellenberg's Indicator values for British Plants. ECOFACT Volume 2 Technical Annex. Institute for Terrestrial Ecology, Huntingdon, Cambridgeshire. Lang, D. (1989) A guide to the wild orchids of Great Britain and Ireland. Oxford Universiy Press, Oxford. Marsh, R and Campling, R.C. (1970) Fouling of pastures by dung. Herbage Abstracts 40, 123-130. Redfem, M. and Shirley P. ( 2002) British plant galls; identification of galls on plants and fungi. Field Studies 10, 207-531. Rich, T.C.G. and Jermy, A.C. (1998) Plant Crib 1998. Botanical Society of the British Isles, London. Rasmussen, H.N. 1995. Terrestrial Orchids from Seed to Mycotrophic Plant. Cambridge University Press, Cambridge. 59 Stace, C.A. (1997) New Flora of British Isles, Second Edition. Cambridge University Press, Cambridge. Stewart, A., Pearman, D.A. and Preston, C.D. (eds.) (1994) Scarce plants in Britain. Joint Nature Conservation Committee, London. UK Biodiversity Action Group (1999) Tranche 2 Action Plans Volume 3. Plants and fungi. English Nature, Peterborough. Welch, D. (1984) Studies in the grazing of heather moorland in north-east Scotland 1. Site descriptions and patterns of utilization. Journal of Applied Ecology 21, 179-195. Welch, D. (1985) Studies in the grazing of heather moorland in north-east Scotland IV. Seed dispersal and plant establishment in dung. Journal of Applied Ecology 22,461-472. Table 1 . Number of flowering plants (F) or vegetative plants (V) of Spiranthes romanzoffiana detected in 2003, shown bold, and details from previous years at 10 locations with exclosures. To economise on space, categories which currently hold no records are not included in the table. Exclosure Unfenced set of 100 Ixlm quadrats outside the exclosure Within 100m of exclosure Site and Non Non appear- Non Presence Newly Non Presence Newly Newly Group appear- ance of appear-ance of 2002 located appear- of2002 located located (A or B) ance of 2001 of2002 plants plants ance of plants plants plants see Intro- 1990- plants. plants. above 2001 above duction 1996 Marker Marker ground plants ground plants. pegs used, pegs used. Marker some lost none lost pegs not through through used sinkage sinkage 1(A) 4 2(A) 4 3(A) 7 Vatersay 4(B) = Colonsay 5(A^) 1 6(A) 1 6 7(B) - 8(B) - 9(A) - 1 10(A) 1 IF IF^ 1F,4V 2F,2V 2V IF Total 18 7 2 2 7 7 5 Notes “ Group A for the area inside the exclosure. One 2002 plant (see note b) occurred outside the exclosure. This may the same plant as one recorded in 2002 and 1 996. However there has been a sinkage of pegs from 2002 to 2003 (pegs were not used in 1 996, a sketch plan was produced). Present in 2002, first found 2001, see Appendix A. 61 Table 2. Heights of the single vegetation level e.g. excluding Juncus spp., at sites five, eight and nine, measured in July 2003 outside and inside the exclosures^ plus values for individual quadrats containing Spiranthes romanzoffiana plants. Site five Site eight Site nine Outside Inside Outside Inside Outside Inside Mean height 54 83 65 123 72 101 Coeff of Var. *’ 43 26 17 17 48 45 Median height 50 80 60 120 60 90 Range 30-120 30-140 40-100 70-180 30-220 50-300 No. of values 100 100 100 100 100 100 Value ofz(n=100) 9.40 24.10 5.11 Significance VHS VHS VHS Single vegetation level heights at individual quadrats containing Spiranthes romanzoffiana plants 30,40,50 90 130 30,50 50,50, 50 Notes VHS Veiy Highly Significant " In early August 2003 one plant occun'ed at site four inside the exclosure, on a very small mound in the slow flowing bum; lower vegetation height of quadrat 80mm. upper vegetation height 1 1 0mm. There was no dung in the hundred quadrats outside the exclosure. *’ Coefficient of variation, values of 40 and above are shown bold. Table 3. Total count in 100 1 x 1 m quadrats of sheep and/or cattle dung at exclosure sites 1 - 10 in late May/early June 2003. Site Number 1 2 3 4 5 6 7 8 9 10 Outside exclosure (0) 29 14 0 36 90“^ 2T ND 4r 64^ Inside exclosure (I) 8 15 24 0 42 67 15 ND 17 73 Mean Outside + Inside 18.5 14.5 46 0 39 78.5 21 29 68.5 Rank 7 8 3 9 4 1 6 ND 5 2 Adjudged intensity L L M(V) VL M H(V) M M M VH(V) of grazing and rank 7.5 7.5 4.5 9 4.5 2 4.5 not inc.'* 4.5 1 Notes H = Heavy, I = Inside exclosure, L = Light, M = Moderate, ND Not Determinable: dung sinks rapidly into the fluid upper layers of the substrate O = Inside exclosure, V = Variable, VH = Very Heavy, VL = Very Light The vegetation height data also indicated heavier grazing outside the exclosure cf inside for 3, 6, 7 and 9. Dung counts were not determinable at 5 quadrats due to 100% standing water. ‘'The vegetation height data indicated lighter gi'azing outside the exclosure cf inside at 10. Not included in the ranking to allow comparability with dung counts. 62 Table 4. Percentage of total count of dung and hoof holes that was due to sheep at exclosure sites 1-10. Data gathered late May/early June 2003 immediately before gate closure, inside and outside values pooled 9 10 41 93 7 5 19 57 7 4 Notes ^ No dung present. ND Not Determinable: dung sinks rapidly into the fluid upper layers of the substrate. ‘'No hoof holes present. Site Number 1 2 3 4 5 6 7 8 Dung count % sheep 100 100 100 No dung^ 13 95 86 ND^ Rank 1-8 2 2 2 8 4 6 Hoof hole count % sheep 100 100 100 No hoof 1 56 43 ND'' holes Rank 1-8 2 2 2 8 5 6 Table 5. Total count in 100 1 x 1 m quadrats of sheep and/or cattle dung at exclosure sites five & nine in late May/early June (M/J), late July /early August (J/A) and late September/early October (S/0) 2003. Site Number 5 5 5 5 M/J J/A S/0 M/J - S/0 9 M/J 9 J/A Outside exclosure 42 39 21 Inside exclosure 36 13 8 21 17 20 10 28 41 8 1 9 7 40 63 Table 6. Jiincus articulatus and Juncus acutiflorns xarticulatus at sites 1 - 10. Site Number 23456789 10 Frequency of both Juncus articulatus and Juncus acutiflorus x articulatus, pooled, in 2003 late Sep./early Oct. 2003 late Jul./early Aug. 2003“ (for comparison with (relates to height measurements in text) taxonomic determinations) Outside Inside 20 7 46 17 35 6 85 21 98 75 100 100 94 100 95 99 100 100 97 100 Juncus articulatus frequency as a percentage of the frequency oi Juncus articulatus or J. acutiflorus x articulatus and Juncus articulatus for quadrats with one or both taxa*’ late Sep./early Oct. 2003 (28 Sep. -6 Oct.) (plants ungrazed) 100'-' 100'-' 100^' lOO*-'' late Sep. (20-24) 2002 (plants grazed: records from 100 Ixlm quadrats before exclosure erected) 92 100 65 43 10 Notes “ Full frequency count and height measurements were in late July /early Aug 2003 at sites 5-10. In Sep. /early Oct. 2003 a sample was taken. Plants not determinable due to grazing of floral parts excluded throughout from this data set. " All outside i.e. grazed plants (recorded in 2003) were Juncus articulatus at sites 1-4. '^Juncus acutiflorus x articulatus occurs nearby. Table 7. Frequency of Juncus effusus, Calluna vulgaris and Sphagnum species in 2003, to indicate aspects of the wetness and variability of the sites (see also Appendix A). Rank positions 1 and 2 are shown bold Site Number 2 3 4 5 7 8 9 10 Juncus effusus Late September/early October Outside 1 0 26 Inside 3 7 64 Mean Outside + Inside 2 3.5 45 Rank 10 8.5 1 Calluna vulgaris “ Late September/early October Outside 8 18 3 Inside 6 17 1 Mean Outside + Inside 7 17.5 2 Rank 546 Sphagnum spp. Late July/early August Outside 96 100 59 Inside 100 97 62 Mean Outside -h Inside 98 98.5 60.5 Rank 2 1 5 Notes Erica cinerea had a frequency of 2 Outside at site five; and 2 Outside, 1 Inside at site nine. 11.5 4 79 70 2 93 92 92.5 3 61 43.5 3 31 19 0 1.5 7 0 9.5 0 12 47 15 8 34 7.5 10 40.5 6 5 2 2 70 0 0 73 0 1 71.5 0 8 1 9.5 13 75 4 18 89 0 15.5 82 2 7 4 8 The Speannan rank correlation coefficient for the association of Calluna vulgaris and Erica tetralix is 0.87, using mean of Outside and Inside values. The association is Very Highly Significant. 64 Appendix A Appendix A Site characteristics of exclosure plots plus original record of plants in the general locations of the 2003 exclosures in two categories; I) inside = within the exclosures when subsequently erected O) outside = in an unfenced set of 100 Ixlm quadrats adjacent to or around the exclosure. Code Grazing “ Study site attributes, I and 0 unless indicated Date No. of plants Site & Grazing Adjud Slope: Presence Light (D) Presence Number Date of Inside In subs- Group verte- ged flat(F), of bum or heavy (P)of of original subsequ- equent (A or B) brate. overall gentle (BU), (DD) dry blocks record ent 2003 unfenced see Most inten- slope ditch disturb- zones. of exclosure set of 100 Intro- abundant sity of (GS) (DI), ance of other quadrats plot Ixlm duction first graz- slopin lochan margins than the outside quadrats ing' g(S), shore ofBU, DI margins the very (LS), or LS: or of exclos- gentle marshy of marshy protrud- ure slope subsect- or flat wet ing (VGS) ion (MS) or flat wet area (FW). areas Rocks Barra 1(A) S L VGS 4 1990 4 2(A) S L F FW 3 1990 4 3(A) S M(V) S DI D P 1 1990 7 Vatersay 4(B) S,C^ VL F BU D 1 15/08/02 2 Colonsay 5(A) S,C M F BU DD P 1 18/08/96 1 1 6(A) S,C H(V) F LS DD P 2 27/7/95^ 1/6' 7(B) S,C M GS MS-0 D 2 30/07/01 1 1 8(B) S,C M F FW DD pd 1 14/08/01 1 9(A) S,C M F DI DD 3 14/08/01 1 10(A) S,C VH(V) S P 1 17/08/96 1 Notes C = Cattle, F = Flat, G = Geese, GS = Gentle Slope, H = Heavy, L = Light, MS = Moderate slope, R = Rabbit, S = Sheep, V = Variable, VGS = Veiy gentle slope, VH = Very Heavy, VL = Very Light “ And other activities related to stock presence e.g. trampling. Grazed by cattle in the winter months. 27/7/95 for I plant: 16/5-14/08/01 for 6 plants. ‘‘Asa low bank c.30cm wide on one side of both I and O areas. 65 Appendix B Appendix B Dates of visits to exclosure sites in 2002 & 2003 Site 2002 2003 2003 2003 Barra 1 20 August 1-2&4 June 3 August 28-29 September 2 21 August 2-3 June 4 August 29-30 September 3 Vatersay 1 9 August 4 June 5-6 August 30 September to 2 October 4 1 7 August 5 June 7-8 August 3-6 October Colonsay 5 23 September 21-22 May 26-3 1 July 20-21 September 6 20 September 26 May 28-29 July 19 September 7 24 September 25 May 29-30 July 1 8 September 8 23-24 September 23 May 25 July 17-18 September 9 22-24 September 28-30 May 23&31 July 24 September 10 22 September 24 May 25-26 July 21-22 September LEGENDS TO COLOUR FIGURES FOR PAPER 1 AND PAPER 2 BY Richard Gulliver, Mavis Gulliver, Margaret Keirnen, and Christopher Sydes. Figure Al. Irish lady’s-tresses orchid (Spiranthes rotnanzojfiana) with 13 fully open flowers on 1 1/8/99 at site KA. The stems and leaves of Jiincus aciitiflorus x articulatiis fonn an upper layer at 300mm. The first vegetation layer, around the base of the plant, is 100mm; with a zone of stoloniferous, dicotyledonous plants at 20mm. Figure Bl. A newly discovered plant of Spiranthes romanzoffiana on 2^11103 within 100m of the exclosure at site five. The two bottom stem leaves and one surviving basal leaf have been truncated by vertebrate grazing. Twelve of the flowers were open, seven unopen. The plant was surrounded by Jimcus aciitiflorus x articulatiis stems, leaves and flowers (some present in the photograph but somewhat out of focus); Juncus articulatiis occurred 80cm from the plant. Figure A2. Spiranthes romanzoffiana in bloom on Colonsay on 9/8/95. Careful examination of the illustration shows that the bract of the left hand flower has been truncated. In addition the two upper stem leaves appear to have been truncated and the partly withered. These features suggest that grazing has occurred at the time when the whole inflorescence was emerging from the ground. Such an event might have removed the upper flowers; these are the last to open, when the inflorescence is fully developed. This would explain the uniformity in stage of development of the six flowers that are present. Figure A3. Spiranthes romanzoffiana, plant A4 at site GF on Coll on 1/10/02, showing withered, unexpanded capsules. Photograph Ms. Emma Grant. Figure A4. Autumn lady’s-tresses orchid {Spiranthes spiralis) in bloom on 13/08/90 on the Gower in South Wales Figure A5. The exclosure at site KA on 6/10/01. The fence line immediately behind the comer post divides the grazed wet sampling stations (right) from the ungrazed wet sampling stations (within the exclosure). To the left of the comer post is the first of the dry sampling stations. Inside the exclosure Juncus spp. (mainly Juncus acutiflorus x articiilatus) are present in the second vegetation layer (which appears more dense on the photograph than is the case on the ground). Outside the fenceline they have been grazed down. Figure A6. Spiranthes spiralis in Cumbria on 18/10/02. Photograph Mr David Benham. The lower capsules have become fully expanded (‘inflated’), dried naturally and then developed slits which allow the seeds to be shed. This is best shown in the lowest capsule on the right hand side. Figure B2. Part of row A (extreme left) and of row B, outside the exclosure, at site five on 27/7/03, showing the small bum with trampled margins. The track which passes through the outside quadrats at the site is more clearly visible in the background. The tape on the right of the picture shows the outer boundary of the outside set of quadrats. 66 Figures A1 upper left, B1 upper right, A2 lower left, A3 lower right. Figures A4 upper left, A5 upper right, A6 lower left, B2 lower right. Glasgow Naturalist 2004. Volume 24. Part 2. Pages 69-70. REDISCOVERING THE FIRTH OF FORTH BELUGA Andrew C. Kitchener and Jerry S. Herman Dept, of Geology and Zoology, National Museums of Scotland, Chambers Street, Edinburgh EHl 1 JF E-mail:a.kitchener@nms.ac.uk, i.herman@,nms. ac.uk The beluga, Delphinapterus leucas (Pallas), is an Arctic cetacean, which has been recorded rarely in Britain from sightings and standings (Barclay & Neill, 1821; Fraser, 1934, 1974; Evans, 1991, Flower, 1880; Millais, 1906; Turner, 1912). Records of strandings from Scotland comprise two young males that were stranded in August 1793 east of Thurso in the Pentland Firth, a specimen at Auskerry, Orkney in October 1845, a specimen near Dunrobin, Sutherland in June 1879, another at Wick in 1884 and finally one near Stirling in the Firth of Forth in October 1932. There is an earlier record of a beluga in the Firth of Forth, where an animal was frequently observed for about three months in 1815 in between Alloa and Kincardine (Barclay & Neill, 1821). It swam upstream on the incoming tide and downstream on the ebb. Many attempts were made to kill it before it succumbed eventually to the firearms and spears of salmon-fishers on 6"’ June 1 8 1 5 at the Abbey of Cambuskenneth, Stirling while in the pursuit of salmon. The animal was purchased by Robert Bald, who sent it to Professor Jameson in Edinburgh, having saved it from going to Glasgow (!) or being “cut to mince-meat for a soap-work”. This beluga was described by Neill and dissected by Barclay (Barclay & Neill, 1821). The animal was a male and measured 13 feet 4 inches (4 metres) in a straight line. This specimen was supposedly the source of a skin that was mounted for display in the Natural History Museum of the University of Edinburgh (Evans, 1892). Unfortunately, this stuffed beluga appears to have been disposed of by the Royal Scottish Museum in the early 1950s. However, close examination of Barclay & Neill (1821) reveals that it was highly unlikely that this specimen’s skin could have been prepared for display. Neill described the soft parts as “quickly passing into a state of putrefactive fermentation” and Barclay regretted that his dissection was very incomplete because of the “putrid state of the body, and the shortness of time which 1 had to examine it”. Plate XVIII (Barclay & Neill, 1821) shows that extensive areas of skin had apparently been cut away to allow Lizars to sketch the viscera in situ, although this could have been artistic licence. Barclay also described putting pieces of the skin into spirit, which hardly seems consistent with saving its skin for a museum display. Curiously, no mounted specimen of beluga is recorded in the registers of the National Museums of Scotland (NMS) and its predecessors. Therefore, it is unclear from where the stuffed beluga originated. It is possible that Evans (1892) inadvertently linked the story of the Firth of Forth beluga with the stuffed specimen in the Edinburgh Museum of Science and Art (now NMS). All that appears to have been preserved from the Firth of Forth beluga was the skull, vertebral column and ribs of the skeleton (Barclay and Neill, 1821, p. 388). However, the whereabouts of this skeleton is unknown. It could have been in the collection of the Royal College of Surgeons in Edinburgh, where Barclay was based. However, much of Barclay’s collection was destroyed in the 1960s and no beluga skeletons survive there today. Today NMS’s collections contain three beluga skulls (Herman, 1992). Two of these are of Arctic origin, but one (register no. NMSZ 1956.36.54) lacks data and originated from the Turner Collection in the Anatomy Department of Edinburgh University (Turner, 1912; register no. C. Dpt. 1.1). Turner’s (1912) only comment about this skull was that it was from the Monro Collection. Herman (1992) erroneously recorded this skull as being from a female, but it is unsexed. We compared the published measurements of the skull above (Turner, 1912) with those of the Firth of Forth beluga (Barclay & Neill, 1821). Although the skull widths matched (i.e. 1 1 inches [=279 mm]), the length of the Tumer/Monro specimen was one inch shorter (i.e. 20 inches compared with 21 inches [=508 mm compared with 533 mm]) . However, we re-measured the Tumer/Monro skull and found that it was exactly the same length as the Firth of Forth specimen (Barclay & Neill, 1821). This suggested that the Tumer/Monro specimen could be the lost Firth of Forth beluga. Other evidence to support this view comes from damage caused to the skull, presumably during the dissection. For example, Barclay noted that “1 once observed the front teeth in the lower jaw, but before we proceeded to the dissection, some person had secretly extracted them”. The teeth are absent from the mandible of the Tumer/Monro skull and the alveoli are covered in as much soot from gas lamps as the rest of the mandible, suggesting that they have always been missing since the specimen was prepared. The cranium has been crudely split in two, with saw marks still evident on the right hand side. Barclay did attempt to look at the brain, but found “The brain was putrid”. Taking the skull measurements, the missing teeth and damaged cranium together, it is highly suggestive that the Tumer/Monro beluga skull is that of the missing Firth of Forth beluga. Other specimens from the Turner collection are known to have originated from the Royal College of Surgeons, so the transfer of the skull to the Anatomy Department is not unusual. However, a mystery remains - the location of the rest of the skeleton. 69 References Barclay & Neill (1821). Account of a beluga or white whale, killed in the Frith of Forth. Memoirs of the Wernerian Natural History Society 3, 371-395. Evans, W. ( 1 892). The mammalian fauna of Edinburgh. M’Farlane and Erskine, Edinburgh. Fraser, F.C. (1934). Report on Cetacea stranded on the British coasts from 1927 to 1932. British Museum, London. Fraser, F.C. (1974). Report on Cetacea stranded on the British coasts from 1948to 79(56.British Museum (Natural History), London. Flower, W.H. (1880). Exhibition of, and remarks upon, the skull of a beluga or white whale {Delphinapterus leucas). Proceedings of the Zoological Society of London 1879, 667-669. Herman, J.S. (1992). Cetacean specimens in the National Museums of Scotland. National Museums of Scotland, Edinburgh. Millais, J.G. (1906). The mammals of Great Britain and Ireland, vol.3. Longman, Green & Co, London. Turner, W. (1912). The marine mammals of the Anatomical Museum of the University of Edinburgh. Macmillan, London. 70 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 71-78. CLIMATE CHANGE AND ITS EFFECTS ON CATCHMENT HYDROLOGY OF THE RIVER SPEY, SCOTLAND, THE RIVER NECKAR, GERMANY, AND THE RIVER ALPENRHINE, SWITZERLAND Gerrit Klemm Department of Civil Engineering, University of Glasgow, G12 8QQ, ABSTRACT Climate change is controlled by many different factors. These include solar radiation, atmospheric gases and volcanism. The prediction for a climate change in the near future is a temperature increase in the range of 0.5°C and 4.5°C by the year 2030, but varies for different regions. The responses of hydrological regimes to climate change are reviewed, and the role of General Circulation Models, Hydrological Models, and their coupling, are described in the context of climate change. Potential changes in sediment discharge carried by rivers is also described. The change in discharge of rivers in response to climate change is difficult to predict because the locality of different rivers has very specific characteristics. The level of uncertainty of a prediction of the response of hydrological regimes is quite high due to difficulties in modelling climate relationships. Recently published results, based on GCMs, predict a general increase in precipitation and in some regions (e.g. mountain areas, like the Alpenrhine catchment) also a change in the seasonality of rainfall. It is also difficult to assess the changes in river discharge, because there are too many factors of the prevailing catchment to be considered for a general statement to be made. The effects of climate change on sediment transport have rarely been the topic of climate change research so far. Three rivers are considered in detail, the River Spey in Scotland, the River Neckar in Geiinany, and the River Alpenrhine in Switzerland. In the River Spey catchment, the present-day rainfall pattern shows weak seasonality with a slight tendency towards more rain in the autumn and winter period. The consequence is that floods are approximately evenly distributed throughout the year, but in the case of large basins, such as the Spey catchment, more pronounced winter- dominated flooding is to be seen. For continuous climate warming, the future prediction is that there will be an increase in runoff due to a significant increase in winter rainfall. In other words there will be more pronounced seasonality for the winter season and there may also be more concentrated periods of rain. The influence of snowmelt is almost insignificant. In the River Neckar catchment, the present pattern of rainfall and runoff shows that there are two main seasons, winter/spring and summer/autumn. The winter precipitation is partly stored as snow but this is not as high as for the River Alpenrhine. Floods usually occur in late autumn and at the end of winter/beginning of spring. In the autumn the floods are just rain-fed, while melted snow will additionally feed the spring runoff. All in all, the winter and spring runoffs are higher than the summer and autumn ones. Increased temperature would lead to an increased runoff in winter, and a decreased runoff in spring and summer, of which the summer would be the driest season. The River Alpenrhine catchment shows clear seasonality for rainfall and for floods. At the present all four seasons are almost independent. In winter the precipitation is stored as snow and the temperatures are so low that the mnoff is minimal. In late spring the runoff caused by precipitation is additionally increased by the volume of melt-water from the snowmelt. The summer and autumn differ in the amount of precipitation and runoff Here, the changes caused by increased temperature will be very significant. The time during which the winter precipitation is stored as snow will decrease considerably, and therefore the immediate runoff will increase drastically (Kwadijk and Rotmans, 1995). As a consequence the volume of melt-water will decrease and will not be available anymore for the spring and partly also for the summer. Additionally, the precipitation will decrease in the summer and at the beginning of autumn. Basically, this means that the River Alpenrhine would change from a snowmelt fed river to a rain-fed river. INTRODUCTION Throughout the Earth’s history its climate has changed by cooling and warming, with other aspects of climate such as rainfall also fluctuating as a result. Global patterns of climate also change differently in different regions. The timescales of periods of global wanning and cooling include decades, centuries, millennia and millions of years. Most interesting for the purposes of environmental biologists and water engineers are changes that occur on a relatively short time scale of decades or centuries. On such a short time scale, the last period of global cooling ended in the mid-1960s, which 71 Since then, the Earth has been in a period of global wanning and the prediction is that this period will continue. However it is almost impossible to say for how long this will go on (Dawson, 1992). In this article the author will review the factors which influence global climate, and then consider the effects of global wanning on three representative rivers, one in Scotland, one in Germany, and one in Switzerland. The research presented is based on knowledge and facts as they were available in 1999. CLIMATE CHANGE Solar Radiation The most fundamental factor exercising the greatest influence over the climate is the Sun. Solar radiation is not constant. It fluctuates in a cyclical manner mainly due to the Earth’s orbital mechanisms. An explanation for the onset of ice ages is that they are triggered by changes in the tilt of the Earth around the Sun and to changes in the tilt of the Earth’s axis, which change the distance between the Earth and the Sun and so cause a climate change (Dawson, 1992). Smaller climate changes over short periods (a few decades) are also set off by this mechanism. Atmospheric Gases The Earth’s atmosphere is another major factor. In general tenns, the response of the atmosphere to solar radiation is threefold: absorption, scattering and reflection. Changes in the concentration of atmospheric gases will induce global changes in temperature. The absorption of outgoing long-wave radiation by carbon dioxide (CO2), methane (CH4) and water vapour has come to be known as the “greenhouse effect” owing to the retention of heat by these gases (Dawson, 1992). The greenhouse effect is caused by the blocking of outgoing radiation by a number of gases in the troposphere, which is the lowest layer of the atmosphere, in which weather processes take place. Principal among these are carbon dioxide and methane, both of which absorb, scatter and reflect this radiation. As a result temperatures in the troposphere, are much higher than they would otherwise be. The large increase in the amount of carbon dioxide, as the major greenhouse gas in the atmosphere, is mainly due to the burning of coal and oil and the removal of large areas of forest cover. The role of other absorbing gases (nitrous oxide, ozone, methane, and chlorofluorocarbons (CFC’s)) is claimed to be already as important as that of carbon dioxide (WMO, 1986). Should the present trends be maintained, it is estimated that by about 2030 AD the combined effects on the terrestrial radiation balance of all those gaseous constituents could be nearly the same as the effect that would be induced by doubling of the pre- industrial level of atmospheric carbon dioxide (WMO, 1986). Volcanism Volcanic activity is another very important factor inducing climate change. Scientists have long known that major explosive volcanic eruptions often result in global cooling. Such eruptions are mostly associated with the injection of large quantities of volcanic ash into the stratosphere where the ash can persist for several years. This ash absorbs incoming solar radiation and is heated (Dawson, 1992). Predicted Future Climate Despite the enormous amount of information on past climates now available, the task of accurate climate prediction is still as elusive as ever (Dawson, 1992). Nevertheless, most predictions of global wanning in the near future assume doubling of the carbon dioxide concentration in atmosphere. It would entail large perturbations in the present climate, notably an increase in temperature and absolute humidity within the near surface air layer (Kellogg, 1979; Watts, 1980; Berger, 1981; Clark, 1982; Manabe, 1983; Flohn and Fantechi, 1984), a decrease in net terrestrial and global solar radiation (Chou et al., 1982; Ramanathan, 1981) and modifications in the annual rainfall regime and in the nebulosity (Manabe et al., 1981; Washington and Mehl, 1983, 1984; Mitchell, 1983, 1986). Predicted increases in temperature lie between 0.5°C and 4.5°C by the year 2030 and differ for various regions (van Dam, 1999). Hence, predictions of climate change show many uncertainties and their use for secondary predictions such as the response of hydrological regimes have to be handled with care. Nevertheless, this prediction was acknowledged by the Intergovernmental Panel on Climate Change (IPCC, 1985) as the standard of the recent research in the field of climate changes. RESPONSE OF HYDROLOGICAL REGIMES TO CLIMATE CHANGE Changes in Rainfall and Runoff All predictions of future climate scenarios are based on General Circulation Models, while the hydrological aspects are based on hydrological models. In order to obtain results for the response of hydrological regimes to climate change, two of these different types of model are usually coupled. General Circulation Models (GCMs) GCMs are used to obtain descriptions of current atmospheric processes. At present there are only a few of these models, because they require a considerable effort to design, and they can only be implemented and run on very large-capacity computers. The areas modelled by GCMs are subdivided into grid cells with horizontal dimensions of the order of 300x300 km“ to 1000x1000 km". In the vertical direction, the Earth’s atmosphere is subdivided into six, ten, or more layers, each several hundred metres thick. The temporal resolution is in the order of 30 minutes to 1 hour. 72 The atmospheric processes modelled by GCMs are based on mathematically formulated physical laws. The various GCMs differ not only in grid sizes and the number of layers, but also in the number of processes and relevant parameter values that can be included (van Dam, 1999). The weak points of GCMs are their inability to model the physics of clouds properly, and do not take account of the presence and effects of aerosols and volcanism, which can have a great influence on climate changes. Therefore the results obtained from various models are often quite diverse. Another reason for the differing results is that the important elements in GCMs are the interactions between land and water surfaces, and the fluxes of energy, water and CO2. The descriptions of these processes are based on small-scale field data or models which may not be valid for such large grid sizes. Recently, regional circulation models (RCMs) have been developed. The grid cells of RCMs are in the order of 50x50 km‘. This makes RCMs more useful for application to relatively small regions. They use inputs from GCMs, but the resolution of an RCM for a particular region is better than that of a GCM. Hydrological Models Hydrological models are usually built for small- scale problems and for a particular purpose, for example for modelling of runoff or groundwater in a given locality, and hence can therefore vary enormously. They can be physically based models, conceptual models, or “black-box” models, and can either be deterministic or partly stochastic. As inputs, hydrological models use prescribed data, spatial analogue data, temporal analogue data or output data from GCMs (van Dam, 1999). Prescribed data may consist of for example present day precipitation, multiplied by a particular factor (multiplicative method) or the same data with an addition (additive method) (Brandsma, 1995). Spatial analogue data consists of data recorded at other locations, where the present climate could be the future climate of the location under consideration (Brandsma, 1995). Temporal analogue data is data recorded at a particular location over a period in the past in which the climate was wetter or warmer, and which is similar to that which might be expected during or after climate change in the catchment area under construction. Coupling of GCMs and Hydrological Models The main problem here is one of scale. As van Dam points out (van Dam, 1999), there is a discontinuity between the size of the GCM grid cells and the basin area for which the interactions between the atmosphere and the hydrological cycle are modelled by using a water balance approach. This means that there is a problem of regionalization or upscaling of the fluxes obtained by hydrological basin studies to be used as an input for GCMs, and/or of downscaling the results of GCMs to individual basins (Feddes et ah, 1989; Feddes, 1995). A better understanding of the interactive processes that result in fluxes is therefore essential for more effective modelling. Changes in Precipitation and Runoff Throughout the relevant literature the published results for different regions vary considerably and they are rarely compatible, but general trends can be identified. The latest results in recent research are listed in the publication of the Intergovernmental Panel on Climate Change (van Dam, 1999). However estimates of the possible change in precipitation made with the aid of either GCMs or instrumental scenarios are extremely uncertain. In general, the changes in precipitation and runoff differ from region to region. For instance in arid and semi-arid regions (e.g. parts of the USA, Russia, Australia, Africa and South America), a change in temperature of 1°C to 2°C and a decrease in precipitation by 10% could reduce annual river runoff by up to 40% or 70%. Compared to that, the changes range from a possible decrease of 6% to 1 2% to an increase of up to 20% in humid tropical regions. Changes in River Discharge The changes of discharge for particular rivers due to climate change are not easy to assess. There are many factors of the prevailing catchment characteristics to consider, e.g. potential evapotranspiration, soil moisture deficit (SMD), urban land use, and vegetation. Each factor contributes to uncertainty of a future prediction and the combination of many unclear factors will increase the uncertainty again. That is why most of the models, apart from the RHINEFLOW model by Kwadijk and Rotmans, only deal with precipitation and runoff. Discharge data estimated from precipitation and runoff results, obtained from GCMs, are based on many assumptions. Therefore results generated from discharge data, which were Changes in Sediment Transport Very little research has been done on alterations in sediment transport caused by climate change. Most of the research carried out on sediment transport has dealt with changes due to alterations in land use (e.g. Ferguson, 1991), and some research has been done regarding the consequences of climatic change (e.g. Collins, 1991). These studies focused on the comparison of historical data and data gained in recent fieldwork. For instance Collins (1991), in his work on climatic and glaciological influences on suspended sediment transport from an alpine glacier, states that climate directly influences runoff. The process of how variations of flow during the ablation season are translated into suspended sediment transport depends on the areal interaction between the elements of the evolving basal drainage system with the glacier subsole. However, no work has been published yet on the direct effects of global warming on either bed load sediment transport or suspended load sediment transport. This may be due to the high degree of 73 uncertainty which would accompany any research on this topic. Precipitation Pattern and Seasonality In order to assess ehanges of sediment transport due to climate change, it is necessary to know the seasonal pattern of precipitation and consequently the pattern of runoff Both are predicted to vary due to climate change. Therefore it is useful to know what these patterns look like at the present and how they might possibly evolve in the future. Present patterns of precipitation are well known and can be easily assessed by using meteorological and hydrological records for any particular catchment. Usually the runoff pattern shows a seasonal character, but this might be different in the case of floods, as they vary considerably from region to region. The view of this pattern in this paper is limited to the three considered eatchments. One also has to bear in mind that ehanges in land use, which may have a considerable influence on seasonality, are not eonsidered for any of the three catchments. COMPARISON OF THE EFFECTS OE CLIMATE CHANGE ON RIVERS IN SCOTLAND, GERMANY AND SWITZERLAND With this background, the authors aim is to compare the potential effeets of climate change on three river catchments of approximately equal size, differing in the geographieal region they are situated in, in the climate zone and the character of flow. The tliree selected rivers are the River Spey in Scotland, the River Neekar in Genuany and the River Alpenrhine in Switzerland. THE RIVER SPEY IN SCOTLAND The catchment of the River Spey is situated in the northeast of Scotland and has an area of 3,010 km^ with a stream network length of 36,399 km, of which the main river comprises 120 km. The river flows in a predominantly northeasterly direction. from its source in Loch Spey, above Laggan to its entry into the Moray Firth at Tugnet, for approximately 157 km. The River Spey is the seventh in terms of length, the eighth largest in terms of mean annual flow and the ninth in terms of catehment area, in the British Isles. From the west there is only one major tributary, the River Dulnain. This drains the Monadliath Mountains. However, from the east and south of the main river there are a number of large tributaries, mainly draining the Caimgonn Massif, of which the River Avon is the largest, with a subcatchment of 1,390 km^. A feature unique to the River Spey in comparison to other major rivers in the northeast of Scotland is its rejuvenated character. That means that when land is uplifted, or the base level of a river is lowered for any reason, the river's erosive power is inereased. The gradient do-wn which it flows is increased and its powers of vertical erosion a re also increased. The river therefore becomes more active and is literally rejuvenated. The upper catchment to Newtonmore is relatively steep (1:225) as is the lower river below Grantown-on-Spey (1:380). The central part of the catchment is characterised by a broad meandering channel, wide flood plain, and 74 passage through Loch Insh, where the gradient (just 1 : 1200) is more typical of a lowland river. Geology Most of the Spey catchment is underlain by metamorphic rocks which are intmded in a number of places by granite plutons and are overlain at the northern end of the catchment by Devonian sandstone. For most of its length, the River Spey flows through a wide alluvial plain composed of silts, sands and water-borne pebbles. In the upland areas there are extensive beds of peat, some many metres thick. Effects of climate change on the River Spey catchment In the case of climate change leading to wanning between 1°C and 2°C the estimated changes in precipitation over Scotland differ considerably. A comparison of the results of five different GCMs for the entire United Kingdom (Hulmes and Jones, 1989) concluded that the annual precipitation could increase between 20 mm and 200 mm, but that the seasonal distribution of this change was unclear. A rise in annual rainfall in this range could lead to an increase in mnoff between 12% and 30% in the wetter regions of the United Kingdom, which include the south and the west of Scotland. Although an estimate of an increase of 5% to 15% in annual rainfall is a likely scenario, there is a high degree of uncertainty in this estimate, and the bulk of the increase is likely to be concentrated in winter (Rowntree, 1990). A greater increase can be expected in the west and north than in the east and south. The estimate also depends heavily on the model used to link temperature and potential evapotranspiration, and it is certainly true that none of the scenarios with their results can be regarded as more realistic than any other (Amell et al., 1990). Figure 3: River Spey Catchment Gradients (North-East River Purification Board (1995)) In the north-east of Scotland where the Spey catchment is situated, the present-day rainfall pattern shows weak seasonality with a slight tendency towards more rain in the autumn and winter period (Black and Werritty, 1997). The consequence is that floods are approximately evenly distributed throughout the year, but in the case of large basins, such as the Spey catchment, more pronounced winter-dominated flooding is to be seen. For continuous warming, the future prediction (Amell et al., 1990; Mansell, 1997) is that there will be an increase in mnoff due to a significant increase in winter rainfall. In other words there will be more pronounced seasonality for the winter season and there may also be more concentrated periods of rain. The influence of snowmelt is almost insignificant. THE RIVER NECKAR IN GERMANY The catchment of the River Neckar is situated in the south-west of Germany, drains an area of 13,958 km^, has a length of 367 km and flows almost without major diversions from south to north. The 75 River Neckar is one of the major tributaries to the River Rhine in its upstream part and mainly drains the mountains of the Black Forrest. The main tributary from the west is the River Enz and from the east there are the Rivers Fils, Rems, Murr, Kocher and Jagst. In order to obtain a catchment comparable with that of the River Spey, a part of the entire catchment was used. This was the part upstream of the gauging station at Plochingen with a catchment size of 3995 km". From its source in the middle of the moor Schwenniger Moos to the gauging station at Plochingen the River Neckar has a length of approximately 120 km. In the upstream part there is Just one major tributary, the River Eyach from the south-east. The River Neckar has a main gradient of approximately 1:580 for its entire length. In its upstream part it is slightly steeper and at the confluence with the River Rhine it is slightly flatter. Geology In the upstream part of the catchment, the River Neckar is underlain by Jurassic limestone and shell limestone. Further downstream, between Tuebingen and Plochingen, this changes into Keuper, a regional type of marl, before returning again to shell limestone. Most of the published studies on Gemian hydrological regimes deal with the river Rhine. Because of the Neckar’s geographical position close to the Rhine and the fact that it is one of the tributaries of the Rhine, the data of precipitation and runoff changes can be used. In the scenario study by Kwadijk and Rotmans (1995), using a climate assessment called ESCAPE coupled with a water balance model called RHINEFLOW, two main scenarios can be considered, HIGH (increase in precipitation) and LOW (decrease in precipitation). According to the LOW scenario, precipitation decreases from May to November which results in lower runoff in all seasons, because the increase in winter precipitation cannot compensate for the large decreases during the rest of the year. According to the HIGH scenario, the summer precipitation will also increase by 16% to 25% which will result in an increase in runoff throughout the entire year. These are two extreme scenarios for the changes in precipitation and runoff caused by a temperature rise of 1°C to 4°C (used in both scenarios). However, in the model used this leads to a more general result of an increase in discharge by 10% to 20% in winter and a change of discharge between -15% and +10% in summer. The present pattern of rainfall and runoff shows that there are two main seasons, winter/spring and summer/autumn. The winter precipitation is partly stored as snow but the significance of this is not as high as for the Alpenrhine. Floods usually occur in late autumn and at the end of winter/beginning of spring. In the autumn the floods are just rain-fed, while melted snow will additionally feed the spring runoff. All in all, the winter and spring runoffs are higher than the summer and autumn ones. Increased temperature would lead to an increased runoff in winter, and a decreased runoff in spring and summer, of which the summer would be the driest season (Kwadijk and Rotmans, 1995). THE RIVER ALPENRHINE IN SWITZERLAND The River Alpenrhine is situated in the east of Switzerland, where it runs from south to north and drains an area of 6,122 km" on a length of 164 km, before its entry into Lake Constance. It runs predominantly in a northern direction and has its source in the Alps near the St. Gotthard Massif, where the river is formed by the confluence of the Vorderrhine and Hinterrhine. Most of the major tributaries come from the south and east and a few from the north and west. The gradient of the entire river is very steep. Geology The geology of the Alpenrhine is dominated by alternating deposits of gravel, sand, clay and peat as the non-unifonned filling of former lake basins. Further downstream it changes to young valley soils with fields of fluvial coarse gravel. The entire Alpenrhine is rich in sediments and thus in sediment transport. The trends for the River Rhine are most pronounced for the River Alpenrhine (Kwadijk and Rotmans, 1995). According to all scenarios, an increase in temperature will dramatically increase the volume of melt water. Also, winter precipitation, presently stored as snow, will be directly available for stream flow. These two facts when combined lead to an increase of 30% to 60% in winter discharge. The other effect is that this melt water volume will not be available in spring and 76 summer, thus decreasing stream flows by 10% to 1 5% during these periods. In comparison with the catchments considered in Scotland and Gennany, the Alpenrhine catchment shows clear seasonality for rainfall and for floods. At the present all four seasons are almost independent. In winter the precipitation is stored as snow and the temperatures are so low that the runoff is minimal. In late spring the runoff caused by precipitation is additionally increased by the volume of melt-water from the snow-melt. The summer and autumn differ in the amount of precipitation and runoff. Here, the changes caused by increased temperature will be very significant. The time during which the winter precipitation is stored as snow will decrease considerably, and therefore the immediate runoff will increase drastically (Kwadijk and Rotmans, 1995). As a consequence the volume of melt-water will decrease and will not be available anymore for the spring and partly also for the summer. Additionally, the precipitation will decrease in the summer and at the beginning of autumn. Basically, this means that the River Alpenrhine would change from a snow-melt fed river to a rain-fed river. SUGGESTIONS FOR FURTHER RESEARCH As the research is based on the knowledge available in 1999 it is first and foremost necessary to assess the latest set of predictions on climate change and therefore increase in temperature regarding the described problem. Secondly, more research is undoubtedly required to assess the influences of all the uncertain factors brought about by catchment characteristics. These characteristics include sediment supply, land use, vegetation, the final amount of runoff, and especially the discharge caused by precipitation. It is useful to test all these factors on their own to assess the real influence they may have on the change in sediment transport due to climate change. The results could then be tested to find the best solution to include the factors with a significant impact into the method, either as single factors or as possible combinations of factors. Another point is to compare the predicted results of changes in sediment transport with results from historic periods. For such comparison data records of sediment loads over a long period of time are required. A solution to the first two points mentioned above is the coding of a specific computer environment. This enviromnent should enable more powerful and specific-oriented computation of the method. ACKNOWLEDGEMENTS The research for this article was carried out in the Department of Civil Engineering at the University of Glasgow and the Geography Department at the University of Glasgow, under the supervision of Prof Trevor B. Hoey, whose help and encouragement are gratefully acknowledged. I also wish to express my sincere thanks to Professor Alan Ervine, Department of Civil Engineering, University of Glasgow, for giving me the opportunity to attend the MSc Water Resource Engineering Management course. 1 am also indebted to the Bundesanstalt fiir Gewasserkunde (BAfG), Koblenz, Germany, namely Dr Disse, and to the Swiss National Hydrological and Geological Survey (SNHGS), Bern, Switzerland, namely C. Koch, for providing the author with sediment and discharge data for the River Neckar and the River Alpenrhine. REFERENCES Amell, N.W., Brown, R.P.C. and Reynard, N.S. (1990) Impact of climatic variability anchange on river flow regimes in the UK. Institute of Hydrology Report No. 107. Wallingford: Institute of Hydrology, UK. Berger, A. (Editor) (1981) Climatic Variations and Variability: Facts and Theories. NATO Advanced Study Institutes Series, Series C: Mathematical and Physical Sciences, Reidel, Dordrecht, 795 pp. Black, A.R. and Werritty, A. 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(Editor) (1999) Impacts of Climate Change and Climate Variability on Hydrological Regimes. International Hydrology Series, Cambridge: Cambridge University Press. 77 Kellogg, W.W. (1979) Influences of mankind on climate. Annu. Rev. Earth Planet. Set., 7: 63-92. Kwadijk, J. and Rotmans, J. (1995) The Impact of Climate Change on the River Rliine: a Scenario Study. Climatic Change, 30: 397-425. Manabe, S. (1983) Carbon dioxide and climatic change. Adv. Geophys., 25: 39-82. Manabe, S., Wetherald, R.T. and Stouffer, R.J. (1981) Summer dryness due to an increase of atmospheric CO2 concentration. Climate Change, 3: 347-385. Mansell, M.G. (1997) The effect of climate change on rainfall trends and flooding risk in the West of Scotland. Nordic Hydrology, 28 (1): 37-50. Mitchell, J.F.B. (1983) The seasonal response of a general circulation model to changes in the CO2 and sea temperatures. Q. J. Meteorol. Soc., 109: 113-152. Mitchell, J.F.B. (1986) On modelling the effects of CO2 on climate. 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Part 2: Results. Weather, 45 (2): 38- 42. Van Dam, J.C. (Editor) (1999) Impacts of Climate Change and Climate Variability on Hydrological Regimes. International Hydrology Series, Cambridge: Cambridge University Press. Washington, W.M. and Meehl, G.A. (1983) General circulation model experiments on the climatic effects due to a doubling and quadrupling of carbon dioxide 78 Glasgow Naturalist 2004. Volume 24. Part 2. Pages 79-85. GROWTH RATE OF AILSA CRAIG SLOW-WORMS ANGUIS FRAGILIS: PREY PREFERENCE AND TEMPERATURE EFFECTS Cara Lavery, J.R. Downie' and Suzanne R. Livingstone Institute of Biomedical and Life Sciences, Division of Environmental and Evolutionary Biology, Graham Kerr Building, University of Glasgow, Glasgow G12 8QQ. 'j.R. Downie@bio.gla.ac.uk (author for correspondence) ABSTRACT Slow-worms are abundant at lower altitudes on the island of Ailsa Craig (Firth of Clyde). They attain remarkable lengths: one male at 51.8 cm is the longest recorded in the UK. Most are found under metal and wooden sheet refuges, where congregations of about 40 individuals are common. Continuous temperature records in these refuges show significantly higher than ambient day-time temperatures. Prey choice experiments showed no size preference for slugs over a range of about 0.5-2.5g. Slow-worms kept at 27-28°C ate more and grew faster than slow- worms kept at 1 8°C. Keywords: Ailsa Craig, slow worm, Anguis fragilis, growth and temperature, prey preference INTRODUCTION The slow-worm Anguis fragilis is probably one of the UK’s commonest reptiles. There are no reliable population estimates for UK reptiles, but recorded sightings of slow-worms show a widespread distribution with most records in England being in the south, especially the south east, but in Scotland mainly in the south- west (Beebee and Griffiths, 2000). The slow- worm’s fossorial habits contribute to our relative lack of knowledge both of this reptile’s distribution and its ecology and habits. To understand this animal better, we need a good study site. One such is the island of Ailsa Craig which lies in the Firth of Clyde, 16 km west of Girvan. Ailsa Craig was designated a site of Special Scientific Interest in 1984, then a Specially Protected Area in 1990 and has now (2004) become a Royal Society for the Protection of Birds Reserve by agreement with the owner, the Marquis of Ailsa. The island’s main interest is the vast populations of sea- birds which nest on its cliffs, but since a rat eradication programme in the early 1990s, it has become apparent that the island supports a considerable variety of fauna and flora (Zonfrillo, 1994). The first report of slow- worms on the island dates from the 1800’s (Lawson, 1888); they were severely affected by the rat population, but, since the eradication, Zonfrillo (2000; personal communication) has noted not only that slow- worms are abundant but also that some individuals have attained remarkable sizes. The aims of the work reported here were: • To assess the sex ratio and sizes of slow-worms found in refuges on Ailsa Craig. • To measure the temperature in slow- worm refuge sites • To test prey size preference in slow- worms • To measure slow-worm growth rate at different temperatures MATERIALS AND METHODS Visits to study site Ailsa Craig was visited on 17“’ June, 22"“ July, 28“’ August and ll* October 2003. Each visit required a boat trip of 1.5h there and back, with 2-3h field-work on the island. Round much of Ailsa Craig, sheer cliffs face directly on to the sea, but on the east of the island, there is a relatively flat raised beach, rocky and shingly close to the sea, but well vegetated further on shore, rising steeply to a higher level. It is on the vegetated area that the light-house, disused quarry works and mainly ruined village are located (Fig. 1). Around the only habitable cottage. Dr Bernard Zonfrillo has placed a number of corrugated iron and plywood sheets to act as slow-worm refuges. Each visit, these sheets were lifted to sample slow-worms. Siow-worm sampling An original aim of this study was to estimate population size, using the technique of mark and recapture. Since there is no effective marking technique for slow-worms, we hoped to use photographs of the unique head parietal pattenr to identify individuals (Riddell, 1997): unfortunately, our camera had insufficient resolution and this aim had to be abandoned. Since the number of slow-worms under each sheet could be around 30-40, some fully exposed, some half-burrowed, it was not practicable to count total numbers in each refuge: they lie closely entwined and can burrow remarkably quickly. Our method therefore was to lift a sheet quickly and pick up as many slow-worms as we could and transfer them to a bucket. We took them for measuring, then returned and collected a second sample from the same refuge. All slow-worms were replaced under their original refuges after measuring, except those taken to the laboratory for feeding experiments (see later). In addition to sampling regularly under 79 Fig. 1 Study site map and location (re-drawn from Zonffillo, 1994). A: sketch map of Ailsa Craig, showing location of study site on flat area to south of jetty; B: location of Ailsa Craig. the metal and wooden refuges, we used a line transect technique to assess the presence of slow-worms further from the cottage. Slow-warm measuring and sexing On Ailsa Craig, slow-worms were weighed in bags using a Pesola spring balance accurate to O.lg; in the laboratory they were weighed in closed polythene tubs using a digital balance to 0.0 Ig. To measure length, slow-worms were straightened and held straight by one observer (this is not easy!) while the other assessed length to 0.1 cm using a meter stick. Individuals were sexed by the criteria of Beebee and Griffiths (2000): females have a dark dorsal stripe and are generally dark in colour; males lack the dorsal stripe and have a lighter uniform colour. Juveniles are not easily distinguished and even some larger individuals were unclear. Field temperatures Gemini data loggers (‘Tinytalks’) were used to measure field temperature from 17*'' June to if*’ October. These loggers are accurate to 0.1 °C and were set to measure temperature every 2h i.e. 12 times a day. Loggers were sealed in polythene bags to protect them from excessive moisture. One was located under a corrugated iron reftige; another under a plywood refuge; the third was placed in a sheltered but not shaded site on the ground surface near the cottage. The information recorded was later downloaded onto a computer and analysed using Gemini software. Laboratory experiments Samples of slow- worms from Ailsa Craig were taken to our laboratory at the University of Glasgow as follows; 17’'’ June - 13 slow-worms, varying sizes, both sexes (returned 28'*' August); 22"‘* July - 4 slow-worms (returned ll“’ October); 28'’’ August - 10 slow-worms (returned ll”’ October). Each slow-worm was maintained individually in a plastic tank 34x18x17 cm with a tight- fitting lid containing nan'ow air slits. Tanks were filled to 8-10 cm deep with moist peat. A water bowl was placed on each tank, the top flush with the peat surface. The peat was re- wetted weekly. To feed the slow-worms, slugs were collected regularly from allotments, gardens and damp walls round the west end of Glasgow. We did not attempt to discriminate between species of slugs. Runham and Hunter (1970) note the difficulties of distinguishing the species of slugs on external characteristics: from their descriptions, Agriolimax reticulatus is likely to have been the commonest species in our collections. Slow-worm tanks were kept in two rooms a) an unheated laboratory with natural lighting; temperature about 19°C. b) a temperature- controlled room at 27.5-28.5°C, with a 12h light:dark cycle. Prey size preference experiment The 13 slow- worms collected on 17"’ June were fed twice a week for 5.5 weeks with one large (>2g), three medium (l-2g) and three small (