A QUARTERLY JOURNAL OF NATURAL HISTORY FOR THE NORTH OF ENGLAND The Wasps and Bees (Hymenoptera: Aculeata) of the Upland Sites of Brimham Rocks, Caydale, Gundale and Seckar Moor with Woolley Edge Quarry in Watsonian Yorkshire — Michael E. Archer Recorder’s Seventh Report of the Aculate Hymenoptera in Watsonian Yorkshire — Michael E. Archer Seasonal Variations in Nutrient Levels in Bemersyde Moss, Borders Region, Scotland — Kathy Velander and Marina Mocogni Botanical Report for 1999 - Flowering Plants and Ferns — D. R. Grant Yorkshire Naturalists’ Union Excursions in 1999 — A. Henderson Published by the Yorkshire Naturalists’ Union Editor M. R. D. Seaward MSc, PhD, DSc, FLS, The University, Bradford BD7 1DP Notice to Contributors to ‘The Naturalist’ Manuscripts (two copies if possible), typed double-spaced on one side of the paper only with margins at top and left-hand at least 2.5cm wide, should be submitted. Latin names of genera and species, but nothing else, should be underlined. 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The University, Bradford BD7 1DP Volume 126 2001 Published by the Yorkshire Naturalists’ Union 3 THE WASPS AND BEES (HYMENOPTERA: ACULEATA) OF THE UPLAND SITES OF BRIMHAM ROCKS, CAYDALE, GUNDALE AND SECKAR MOOR WITH WOOLLEY EDGE QUARRY IN WATSONIAN YORKSHIRE MICHAEL E. ARCHER 17 Elmfield Terrace, Malton Road, York Y031 1EH Characteristically, the aculeate wasps and bees are active during warm sunny days and, except for the social species, stop foraging activities if the temperature drops too low. Thus with increasing latitude (Archer, 1996a) and altitude it might be expected that the number of species (or species diversity) would decrease. The effect of altitude on species diversity will be investigated for four upland sites in Watsonian Yorkshire. Brimham Rocks (SE2064) is situated in the natural area of the Yorkshire Dales at an altitude of about 260 m. It has an underlying geology of Millstone Grit, from which some spectacular rock formations have developed. Brimham Rocks is a National Trust property with an area of 157 ha. The moorland is mainly covered by heather and bilberry with scattered oaks and silver birches, and willows in boggy areas. The surrounding slopes are covered by deciduous woodland including oak, birch and rowan. The flowers of the heather, bilberry and willow provide important food sources for the aculeate species. Importantly nesting areas for the subterranean nesting species occur where the sandy soil is exposed, especially heavily used footpaths and at the bases of the rock formations. During May 1982, for example, 105 freshly dug burrows of the mining bee, Andrena cineraria, were counted in the sloping ground just below the wall in the front of the house. Further during June 1983 about 120 burrows of several species were counted in the path behind a house. Caydale (SE5386) is situated in the natural area of North York Moors and Hills at an altitude of 210-250 m. The valley cuts into the Coralline Oolite plateau with good exposures of the underlying Calcareous Grits in old quarries near the top of the valley sides (Atherden, 1985). The dry friable soils of these old quarries provide nesting sites for the subterranean nesters. The valley sides have some dry and damp limestone grassland which is being invaded by hawthorn and ash. The valley bottom around the stream is largely marsh. Some areas of old woodland and isolated trees are present, some of which are dead and now providing nesting sites for the aerial nesters. The flowers of hawthorn, raspberry, dandelion, white clover, rock rose, figwort and even heather are important food resources. The area for study is 49 ha centred around the public footpath, but similar habitats extend over a larger area of private property. Gundale (SE8089) is situated in the natural area of North York Moors and Hills at an altitude of 130-150 m. The study area of 59 ha consists of a northern section of long narrow exposures of extremely soft sandstone with surrounding herb-rich acid grassland and a southern section of herb-rich limestone grassland openings in the deciduous woodland. The tops of the sandstone exposures are dominated by heather and the bases of the slopes by flower-rich ruderal communities. Shrub hawthorn has also invaded the open areas. Bare soil areas of the limestone grassland and the sandstone exposures provide nest sites for the subterranean nesters. Seckar Moor (SE3214) at an altitude of 90 m and Woolley Edge Quarry (SE3013) at an altitude of 160 m are both situated in the natural area of the Coal Measures. The underlying geology is Millstone Grit of the Upper Carboniferous. Seckar Moor consists of heath, bog and birch woodland with occasional oak, alder and sallow. The study area of 20 ha is mainly in the open heath habitat which contains much heather and bilberry with some bramble, but the herb flora is poor. The nests of the subterranean nesters are associated with the well-used footpaths, particularly where a small slope or bank has appeared. Woolley Edge Quarry has been abandoned and consists of gritstone crags with sandy Naturalist 126 (2001) 4 Wasps and Bees (Hymenoptera: Aculeata) of Upland Sites in Watsonian Yorkshire pockets of soil in which subterranean nesters nest. The quarry has some heather and bilberry surrounded by deciduous woodland. Methods Between 1974 and 1998, 20 visits were made to Brimham Rocks, distributed throughout the year as follows: April (1 visit), May (3), June (8), July (4), August (3), September (1). Between 1984 and 1998, 13 visits were made to Caydale as follows: May (3), June (4), July (3), August (3). Between 1986 and 1994, 18 visits were made to Gundale as follows: April (1), May (3), June (5), July (4), August (3), September (2). Other people also visited Gundale: W. D. Hincks on 30 June 1956, J. H. Flint on 10 June 1979 and 3 May 1980, R. S. Key on 6 September 1985 and J. D. Coldwell on 19 June 1994. The specimens collected on these visits except Priocnemis perturbator (J. D. Coldwell) and Hylaeus communis (W. D. Hincks) have been seen. The remaining specimens collected by W. D. Hincks were found at the University Museum, Manchester. Between 1983 and 1999, 17 visits were made to Seckar Moor and Woolley Edge Quarry as follows: April ( 1 ), May (4), June (3), July (5), August (3), September (1). In addition W. D. Hincks visited Seckar Moor on 3 July 1943 and his specimens were found at the University Museum, Manchester except for Symmorphus bifasciatus. Unless specifically noted in future Seckar Moor will cover both Seckar Moor and Woolley Edge Quarry. During my visits which lasted for approximately two to three hours, all species of aculeate wasps and bees were recorded (Archer sample) and usually collected with a hand net for identification, except possibly the visits to Brimham Rocks during the 1970s. In the following account, the nomenclature can be related to that of Kloet and Hincks (1978). Species Present and Seasonal Progression of Species A full list of recorded species is given in the Appendix. At the family level, Tables 1 and 2 show the taxonomic distribution of species and records. A record represents a specimen differing in one of the following three variables: name, sex and day of visit. The solitary wasp family, Sphecidae, and the solitary bee families, Andrenidae and Halictidae, are the dominant families in terms of both the number of species and records. Table 3 shows the number of species and when species were first recorded for each month. The most productive month for the species of solitary wasps at the four sites was July except for Caydale when June and July were particularly productive. Most species of solitary wasps were first recorded either during June (Caydale), or July (Brimham Rocks, Seckar Moor) or both June and July (Gundale). Since June and July are summer months, most species of the solitary wasps would seem to have to wait until the warmer summer months before the adults can become active. During the spring months of April and May very few species of solitary wasps were found and at Brimham Rocks and Seckar Moor no species were found. The most numerous species of solitary wasp recorded from Brimham Rocks was Crossocerus dimidiatus which was usually found when hunting for its prey of small flies; from Caydale were Chrysis impressa and Trichrysis cyanea found on fence posts probably on a mating circuit; from Gundale were Crossocerus tarsatus and C. ovalis usually at their subterranean nesting sites, and from Seckar Moor, where very few solitary wasps were found, Lindenius albilabris was the most numerous species, usually found at its subterranean nesting sites in the very small banks at the side of the path. The most productive months for the species of solitary bees was June except for Caydale when June and July and Gundale when May and June were particularly productive. Most species of solitary bees were first recorded either during June (Brimham Rocks, Caydale), or May and June (Seckar Moor) or April, May and June (Gundale). Unlike the solitary wasps, solitary bees were present in numbers during both the spring and summer months. The most numerous species of solitary bee recorded from Brimham Rocks were Andrena fucata, A. cineraria and Lasioglossum calceatum ; from Caydale Andrena haemorrhoa and Halictus tumuloruni | from Gundale Andrena haemorrhoa, Lasioglossum fratellum and 5 Wasps and Bees (Hymenoptera: Aculeata) of Upland Sites in Watsonian Yorkshire TABLE 1 The number of species of aculeate wasps and bees recorded from Brimham Rocks, Caydale, Gundale and Seckar Moor with Woolley Edge Quarry. Brimham Caydale Gundale Seckar Solitary wasps Chrysididae 0 4 3 0 Mutillidae 0 0 1 0 Sapygidae 0 1 1 0 Pompilidae 5 5 7 1 Eumenidae 0 3 2 3 Sphecidae 9 13 14 6 Total solitary wasps 14 26 28 10 Solitary bees Colletidae 1 3 2 2 Andrenidae 10 9 13 13 Halictidae 10 11 13 12 Megachilidae 0 2 1 0 Anthophoridae 5 3 7 9 Total solitary bees 26 28 36 36 Total solitary species 40 54 64 46 Social wasps and bees Vespidae 4 5 5 3 Apidae 13 11 11 9 Total social species 17 16 16 12 Total aculeate wasps and bees 57 70 80 58 TABLE 2 The number of records of aculeate wasps and bees in the Archer sample from Brimham Rocks, Caydale, Gundale and Seckar Moor with Woolley Edge Quarry. Brimham Caydale Gundale Seckar Solitary wasps Chrysididae 0 13 4 0 Sapygidae 0 1 0 0 Pompilidae 6 8 16 1 Eumenidae 0 3 3 1 Sphecidae 26 20 42 8 Total solitary wasps 32 45 65 10 Solitary bees Colletidae 4 7 0 6 Andrenidae 30 23 54 48 Halictidae 32 27 70 39 Megachilidae 0 5 1 0 Anthophoridae 8 6 20 31 Total solitary bees 74 68 145 124 Total solitary species 106 113 210 134 6 Wasps and Bees (Hymenoptera: Aculeata) of Upland Sites in Watsonian Yorkshire TABLE 3 The number of species and when species were first recorded per month of solitary wasps and bees at Brimham Rocks, Caydale, Gundale and Seckar Moor with Woolley Edge Quarry. April May June July August September No. species Wasps Brimham 0 0 5 10 5 1 Caydale - 1 15 14 5 - Gundale 1 4 12 19 8 5 Seckar 0 0 0 8 2 - Bees Brimham 2 5 17 9 8 3 Caydale - 8 18 17 6 - Gundale 10 19 26 12 7 9 Seckar No. species 1 5 first recorded 16 25 13 6 Wasps Brimham 0 0 5 8 1 0 Caydale - 1 15 8 2 - Gundale 1 4 10 9 3 1 Seckar 0 0 0 8 2 - Bees Brimham 2 4 13 2 5 0 Caydale - 8 12 5 3 - Gundale 10 10 11 2 0 3 Seckar 5 12 12 3 4 - L. rufitarse, and from Seckar Moor Colletes succinctus , Andrena lapponica, A. cineraria and A. tarsata. All these species are subterranean nesters and are either found at their nesting sites, e.g. C. succinctus and A. cineraria in nesting aggregations, or at foraging sites, e.g. A. haemorrhoa on hawthorn flowers and C. succinctus on heather flowers. Some of the above findings at Caydale, and particularly at Brimham Rocks, must be regarded as provisional - it will be shown later that many further species may be present at these two sites. Species-area relationship and potential number of solitary species The species-area relationship can be found by plotting the number of solitary species recorded at a site against the area of the site, with both the number and area expressed as natural logarithms (In). Archer (1999) plotted a species-area figure for 18 sites from the north and north midlands of England and found that the points fell on a line indicating a highly statistical significant positive linear relationship. The regression equation for this linear relationship is: in number of species = 3.84 + 0.11 x In area (ha). From Archer (1998a, 2000) the sample can be increased to 20 sites, with no change to the regression equation. Of the four sites under investigation only the Caydale species - area dot falls within the range of the 20 sites, with the dots for Brimham Rocks, Caydale and Seckar Moor (excluding Woolley Edge Quarry) being below the range of the 20 sites. If a site falls below the range of sites with which it is being compared, this could indicate either more 7 Wasps and Bees (Hymenoptera: Aculeata) of Upland Sites in Watsonian Yorkshire recording is needed to find additional species, or the site is less favourable for solitary aculeates than the sites with which it is being compared. To test whether more recording is needed, the potential number of species can be estimated using non-parametric statistical procedures. The presence/absence Chao (in Colwell and Coddington, 1994) and jackknife (Heltshe and Forrester, 1983) estimators have been used, using the software of Pisces Conservation Ltd. The Chao estimate depends on the number of species found and the number of species found only on one (unique species) and two (2-occasion species) visits or samples. The jackknife estimate also depends on the number of species found and the number of unique species. The statistical procedures are run a number of times equal to the number of samples or visits. In practice the software selects 1, 2, etc. samples at random each time calculating a mean estimate of species richness. With a small number of samples the estimates are erratic, but as larger sample sizes are selected the estimates may stabilise giving confidence in the estimates. The estimates with increasing sample size are shown in Figs 1-4 and the estimates from the largest number of samples in Table 4. TABLE 4 Non-parametric estimates of species richness of solitary wasps and bees at Brimham Rocks, Caydale, Gundale and Seckar Moor based on the Archer samples. Chao estimate Jackknife estimate Brimham Rocks No species - recorded 40 40 - estimated 70 58 95% confidence limits 40-100 48-68 % of estimated species found 57 69 Caydale No species - recorded 54 54 - estimated 86 81 95% confidence limits 59-114 65-96 % of estimated species found 63 67 Gundale No species - recorded 59 59 - estimated 75 79 95% confidence limits 59-90 68-90 % of estimated species found 79 75 Seckar Moor No species - recorded 35 35 - estimated 46 48 95% confidence limits 32-60 41-55 % of estimated species found 76 73 For Brimham Rocks, the Chao estimates do not stabilise (Fig. 1), the two estimates disagree with each other and fewer than 70% of the potential number of species have been found (Table 4). These calculations indicate that more solitary species could be found at Brimham Rocks and further visits should be made. Since it is probably not possible to record all the species from a site, provided the estimates stabilise and the two estimates agree with each other, Archer (in press) arbitrarily decided that sampling could stop when 70% or more of the potential number of species has been found. For Caydale, the Chao estimates stabilise (Fig. 2), the two estimates are similar to each Species Richness Estimates Species Richness Estimates 8 Wasps and Bees (Hymenoptera: Aculeata) of Upland Sites in Watsonian Yorkshire FIGURE 1 The Chao presence/absence estimates of species for Brimham Rocks. No. Samples FIGURE 2 The Chao presence/absence estimates of species for Caydale. Wasps and Bees (Hymenoptera: Aculeata) of Upland Sites in Watsonian Yorkshire 9 other although just less than 70% of the potential number of species have been found (Table 4). Further visits could therefore be made. However the estimates probably refer to a larger area of Caydale than those covered by my visits and, as such, further visits should be made to the private larger area. It is unlikely access will be possible. If the larger area is considered as a truer measure of the area of the site then the dot would fall below the 20 comparison sites on the species-area figure. For Gundale and Seckar Moor, the Chao estimates stabilise (Figs 3, 4), the two estimates are similar to each other and more than 70% of the potential species have been found at each site. w100 0) 15 E 80 -4— > 1500 ft. Painted Lady, Cynthia cardui (L.) As such offspring of the Painted Lady as are produced by immigrants in Britain have no overwintering stage its credentials depend entirely on incomers. Like the Red Admiral it has been present, but more intermittently, throughout the period covered and, again like the former, frequents a variety of habitats. Dunning’s record from Storthes Hall in 1846 (Rimington & Beaumont 1996) is the earliest traced. Small Tortoiseshell, Aglais urticae (L.) The Small Tortoiseshell (earliest record 1846: Dunning) was described as very common by Hobkirk (1859, 1868), and according to Mosley (1883) continued to be “abundant throughout the district” until the early 1860s, yet 20 years later “only occasional stragglers” were to be seen. The interest of this decline of a common species is that it coincided with that of other species, and the demise of some, in the area. Porritt (1900a) noted in passing that the Small Tortoiseshell was present in his garden in 1899, “it being . . . now comparatively seldom seen here”, though the Annual Report of the Huddersfield NS had recorded that it “swarmed throughout the district” in 1895, which suggests that large, short- term, fluctuations in abundance were involved. A note that it was “more plentiful than usual” at Halifax in 1899 (Halifax SS) confirms Porritt’s comment for that year and shows that the decline noted by Mosley was temporary. In 1901 it “simply swarmed” at Skelmanthorpe (Morley 1902), but in manuscript notes, where he specifically records the presence there of single individuals in 1915 and 1917 he says that its appearance is “uncertain”, which indicates continuing scarcity. Wattam (1936) reported a definite increase “especially since 1933” - a welcome note of precision - and, with fluctuations in abundance (it was scarce in 2000), it has remained well represented and widely distributed in the area though, by virtue of its regular presence, it tends to go unreported unless included in a survey. In Garland’s (1981) survey of Sheffield it was the most frequently recorded species. Unlike colonial, largely sedentary, species, the Small Tortoiseshell is mobile, often spends much time in direct flights across diverse types of terrain, and frequently traverses several kilometres in a day. In Germany marked individuals have been recovered 150 km. from the point of release. In Britain there is a tendency to move more or less NNW until about mid- August and SSW later in the year (see Emmet & Heath 1989, for summary). Such movements add to the suspicion that occasional individuals arrive in Britain from the Continent. They also demonstrate that it is not only local factors that determine its presence and abundance in a particular area. Its scarcity in the period in the 19th century during which several species suffered extinction in the Huddersfield area may therefore have been 74 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire coincidental. While it has been a component of the local fauna throughout the period for which information exists, little can be said about what determines its abundance. Much remains to be learned about the biology of this common species. Large Tortoiseshell, Nymphalis polychloros (L.) Hobkirk (1859) recorded this species from South Crosland and (1868) reported a brood at Birkby in 1859. Mosley (1883) referred to “occasional specimens” from Farnley and Birkby (Porritt) and Penny Spring Wood (Varley). As Porritt was not born until 1848 his Birkby record mentioned by Mosley may be additional to Hobkirk’s for the same place. As well as Huddersfield (on his own authority) which would embrace all localities specified by Mosley, Porritt (1883) lists others not far distant, of which Horbury and Halifax, fall within the area considered here. It was also recorded from Clayton West by J. Firth, as reported by Brady (1884), who gives no date. It is significant that most of the Yorkshire records of this species, which Porritt describes as “not uncommon but not often taken in numbers”, are from the south-west of the county. Particularly noteworthy is that Wattam (1936) reported it as “still only seen occasionally” as it was by himself at Newsome in 1934 and 1935, and from where he had earlier reported seeing “a fine specimen of this locally uncommon butterfly” (Wattam 1921). Furthermore, he subsequently reported two from Newsome on August 4, 1949, on which day he saw another at nearby Lower Castle Hill, and saw three more at Newsome on October 15 in the same year - a late sighting (Dearing 1950). (The late 1940s were the last time that this species was comparatively common in Britain.) Unless the indication for V.C.63 in the period 1940-49 shown on the map of Emmet & Heath (1989) refers to Wattam’s 1949 sightings, all these records (some of the many treasures buried in the pages of The Naturalist) have been overlooked. Indeed, Sutton & Beaumont (1989) say that, apart from three individuals seen at Allerthorpe in 1948, the only other record for Yorkshire this century was from Keighley in 1910. Only the Keighley record is mentioned in YNU Lepidoptera Committee (1967). It seems that the Large Tortoiseshell, one of whose strongholds was in S.W. Yorkshire, and which is now thought by some to be extinct in Britain except as an intermittent immigrant (or even always to have depended on immigrants, see Emmet & Heath 1989) may have persisted in the Huddersfield area longer than suspected by most entomologists. Unfortunately adults of this species were irresponsibly released at Warley in 1984, and while there were no indications in the following four years that these survived or bred, there is bound to be a faint suspicion that an individual at Norton Tower, near Halifax, in 1989 (Cain 1990) was derived from this introduction. This and another seen at Sowerby Bridge in 1991 were deemed to be migrants by Cain (1992). Two more were seen in 1992, one at Cromwell Bottom on June 11 and one at Well Head on July 26 (Cain & Baggaley 1994). These were also thought to be migrants, as are most of the rare individuals now seen in Britain. While this may be correct, it may be more than coincidental that four individuals of this now very rare species should be seen in the very area that has yielded a series of records over many years. (Apart from Yorkshire the Large Tortoiseshell was not seen in northern England between 1940 and 1988 - Emmet & Heath 1989.) Is it conceivable that it still survives somewhere in the area? Camberwell Beauty, Nymphalis antiopa (L.) Very intermittent in its occurrence, and a rare migrant, the Camberwell Beauty has a long history of appearing in the area at lengthy intervals. It was recorded in Huddersfield in 1858 and at Newsome in 1859 (Hobkirk 1859, 1868); several were seen in 1872, and others at other times (Mosley 1883). There were subsequent occurrences at Huddersfield in 1899 (Porritt 1900b) and Longley Hall in 1924 (Morley 1926), but Wattam neither encountered nor listed it as he doubtless did not regard it as a true component of the local fauna. More recently it was seen at Southowram in 1983 (per B. D. Cain), and there were several sightings in 1995 - Upper Denby (D.Elliot), Almondbury (B. & M.J.L), Slaithwaite (B. Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 75 Brassington), and Lower Shibden (Cain 1997). While of intrinsic interest, such occurrences of this splendid insect are of little significance in the history of the local fauna. Peacock, Inachis io (L.) Listed by Dunning in 1846, the Peacock was described as common by Hobkirk (1859, 1868), but by 1883 Mosley, who confirmed that it was previously common, reported that it had become rare, a state of affairs that, with occasional short-lived local increases, persisted for more than a century. In this instance the decline was not merely local but was related to a general contraction of range in northern Britain that apparently began about 1860. It is interesting that, evidently against the general trend, in the Barnsley area, where it was “universally distributed, but nowhere abundant”, “it appeared somewhat plentifully” in 1 877 (Brady 1884). It clearly became very scarce, and does not appear at all in Halliday’s (1896) list for Halifax, though it had in fact been seen at Ovenden in 1894 (Halifax SS), a record evidently unknown to Halliday and others. This was the first report there since 1876, also from Ovenden. When Robertshaw (1900) caught one in 1900 near Luddenden Foot he noted that it had not been captured “for over a dozen years in the district”, and reporting another from Elland in the same year, Halliday (1901) also said that it had not been seen for many years. Fielding (1902) reported one at Wadsworth a year later. Butterfield (1911) said it used to be common in the Bradford area “but is fast dying out”. Morley’s manuscript list for the Skelmanthorpe area (1896-1908) says “only one or two records of its capture” over the 13- year period. When reporting it at Crosland Hall in 1899, Porritt (1900a) noted that “it is now rarely seen in the Huddersfield district” and the same occurrence elicited the comment “now a great rarity in our district” in the Annual Report of the HNS for 1899. Its long-continued rarity is indicated by the few references to it in Annual Reports of this society - Almondbury 1896, Newsome 1900 (one only) and 1912, and Meltham 1919, a state of affairs that was to prevail for another 50 years or so. It was reported from Mytholmroyd in 1911 (Halifax SS). Notwithstanding its general scarcity, the Peacock occasionally increased markedly for short periods in restricted areas. This was so near Skelmanthorpe in 1917 where it became common in many places (Morley 1918a, 1918b). Its reappearance here caused him to say that this was noteworthy as it had been very scarce for many years. For the same year Porritt (1917) noted that “a few V. urticae and V. io, both usually of rare occurrence here, were about”. At Elland in 1927 it was said to occur “fairly regularly” but to be variable in numbers (Halifax SS, probably on the authority of H. Spencer). Wattam (1936) reported it as “still rare”, sufficiently so to prompt him to refer specifically to having seen one at Newsome in 1934 and three there in 1935, and to add that he had never found larvae in the area. There was a trickle of records in the later 1930s, 1940s and 1950s. Wattam (1937) saw four at Newsome in 1936. In 1939 several were seen at Elland (Halifax SS) and it was reported from Edgerton, Huddersfield, and again from Newsome (Dearing 1940). Several were seen at Elland in 1941, where it was reported as uncommon in 1943, but in that year many were seen at Ovenden (Halifax SS), a parallel to the local increase at Skelmanthorpe in 1917, and evidently as ephemeral. It was seen at Elland in 1944, Halifax 1957 and Triangle 1959 (Halifax SS). It was absent from, or very rare in, the Colne Valley in the 1940s and early 1950s (G.F.). Collinson (1969) recorded that it was rarely recorded around Halifax and sometimes not seen in the parish for a dozen years. The area shared in a Yorkshire-wide revival in the late 1960s and early 1970s, that apparently occurred earlier in the Doncaster area in the south east, where the recovery began in the 1930s and 1940s and continued during the 1950s (Rimington 1992). For the years of expansion Jackson’s annual reports in The Naturalist refer to it as having had an “exceptionally good year”, being “very plentiful” and the like, while Sutton & Beaumont (1989) describe it as rare until the last two years of the 1960s, after which it became common. To this generalisation the Doncaster area was an exception. Over this period it became common and widespread in the Huddersfield area and, apart from a possible but ill- documented decline in the late 1980s, remained so to the bi-millennium. Like the Small Tortoiseshell it is a mobile species that, although with a preference for 76 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire woodlands, traverses a variety of habitats, and is often seen in gardens. It may move for considerable distances each day - in Germany marked individuals have been found up to 94 km from where they were marked. Like the Small Tortoiseshell it tends to move NNW until about mid August, then, in contrast to that species which moves SSW, it often moves SSE. In the Huddersfield area it has been seen at an altitude of c. 1050 ft. The history of the Peacock in the Huddersfield area raises interesting questions. Particularly striking were the localised, short-term increases, as around Skelmanthorpe, and evidently elsewhere, in 1917 and at Ovenden in 1943, in what were otherwise periods of general scarcity. Of such upsurges there may have been another in the Barnsley area in 1877. There were also years when it was not scarce at Elland, and a note by E. Dearing in the YNU Record Book in 1939 says “some years not uncommon”. Some (all?) of these upsurges were clearly the result of local breeding - at Skelmanthorpe the period of abundance was in August and September, after the emergence of locally- produced adults. About 450 to 500 eggs make up a typical batch, and as larvae live gregariously, protected by a web, large broods may reach maturity, and certainly do so at times. Having lived at low densities for a long period, populations would be unlikely to suffer greatly from parasitism. A few broods may therefore produce several thousand pupae that are exposed to predation for only about two weeks, so the sudden appearance of large numbers of adults in a restricted area is not difficult to explain. What is difficult to explain is why such increases sometimes appear to have been localised within the Peacock’s extensive range in the area. Such localisation hints at enormous levels of predation, parasitism, or disease, or adverse environmental conditions, from which these isolated populations were, apparently for a single season, free. Adult mobility would ensure dispersal, either before, or probably more effectively after, hibernation, explaining the ephemeral nature of the events. Mobility raises another issue. Rimington (1992) shows that the Peacock began to recover in the Doncaster area in the 1930s and maintained its improved status in the following decades, as it did in the adjoining Derbyshire, Nottinghamshire and Lincolnshire. Its mobility is such that individuals must have dispersed north and west from south Yorkshire. That its recovery took place later in the Huddersfield area, and indeed in much of Yorkshire, is therefore difficult to explain. If there are climatic limitations they must be very subtle, and suitable conditions must have prevailed intermittently in the Huddersfield area in the years of localised abundance. Genetically different lineages may have been involved. Much remains to be learned about the biology of the Peacock. One even wonders whether it was less rare in the Huddersfield area in the lean years than in much of Yorkshire. Although records are few they show persistent occurrence. Comparative quantitative information is, however, lacking. Comma, Polygonia c-album (L.) (Figure 3, p. 77) The Comma was recorded at Storthes Hall sometime prior to 1859, this being the only locality mentioned by Hobkirk (1859), who designated it as “rare”. Newman (1870-1871) includes “Huddersfield, rarely and singly” as a locality on the authority of Porritt who may have reported the old Storthes Hall record to him. Porritt (1883) does not include any Huddersfield record in his list, which suggests that he deemed the record too ancient to merit citing. Newman also includes Halifax on the authority of Birchall, which Porritt does include but cites the wrong page. This, cited from Newman, is the only 19th century record in the Halifax SS archives (under the erroneous date, 1883, evidently from Porrittt). It was unknown to, and not mentioned by, Mosley (1883) and Wattam (1936) as a component of the Huddersfield fauna. Indeed it had declined throughout Britain by the 1830s, and, following a temporary recovery in the north in the 1850s and 1860s, (to which period the Storthes Hall record may refer) again declined. By the early 20th century it had retreated to the Welsh borderlands and adjacent areas, from which it began to spread in 1914 (summary in Emmet & Heath 1989). It was not, however reported in Yorkshire until 1937 (YNU Lepidoptera Committee 1967) - at Scarborough - but in the 1940s it was seen in several places in the county, especially in the south west, including Elland 1946 (Halifax SS), Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 77 FIGURE 3. Three butterflies with contrasting histories in the Huddersfield area. The Orange Tip (top) persisted, sometimes tenuously, for more, perhaps much more, than 150 years. This historic illustration of a male, drawn and engraved by James Bolton, probably, but not certainly from a locally obtained individual, appeared in his Harmonia Ruralis vol. 2, 1796. The orange wing tips were coloured by hand in each copy. The Comma (lower left) has had a complex history in the area where it became extinct three times, once being lost for about 90 years, but is common at the beginning of the present century. The White-letter Hairstreak (lower right) was unknown until, at a seemingly unpropitious time in the 1990s, when its larval food plant - Elms Ulmus spp. - had recently suffered considerable losses to Dutch Elm disease, it made its appearance in at least six places. The lower figures, from Sutton & Beaumont (1989), were drawn by David Green, whose artistry it is a pleasure to share. 78 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire Cawthorne and Elland 1948 (Dearing 1949), as well as in places just outside the Huddersfield area. Following this invasion it declined and retreated. Perhaps the penultimate sighting was by B. D. Cain at Woodside, near Halifax in 1955, the last record for Yorkshire at that time being in 1957. Seago (YNU excursion circular 1953) caused some confusion when he said that Polygonia c-album was present in Deffer Wood. Although he used the present tense, it is not clear whether he implied that it continued to exist there in 1952 or 1953. However, in a bulletin circulated among YNU lepidopterists, in which he noted that the Comma reappeared in S. Yorkshire “around 1945”, he said that he saw one or more every year since that date “chiefly at Deffer Woods, Cawthorne, until 1950” after which he saw no more in Yorkshire. It is not clear that all these were seen in the area covered, or in which years he saw it in Deffer Wood, but he did not see it there in 195 1 or 1952 as his note in the excursion circular seemed to suggest. As it was recorded from Cawthorne in 1948 (Dearing 1949), at best Seago added one or more sightings in that area in the 5-year period concerned. In the 1970s it again recolonised Yorkshire, but was very rare. For this decade there are few published records, all in the south west, the earliest, Potteric Carr, 1973, being outside the Huddersfield area (Sutton & Beaumont 1989). To this can be added an unpublished record for Cowlersley, in the Colne Valley, where it was seen in September of that year by D. Elliot. It was seen in Halifax in 1974, (A. Bannister) and of two records for 1976 (Jackson 1983) one was for Oakenshaw. Jackson also records it from Beauchief, Sheffield, not listed by Sutton and Beaumont. Thus three of the five known occurrences in Yorkshire in that period were in the Huddersfield area. It was not seen again in Yorkshire until 1980. It then spread quickly throughout the county but was seemingly slow to establish itself in the Huddersfield area, though it was seen at Triangle, near Halifax, as early as 1983 by A. Dale (Sutton & Beaumont 1989). Being very distinctive, and at that time unusual in the area, one assumes it would be recorded if seen, but no further reports have been traced until 1990, at Rastrick and Triangle (Cain 1991), after which it increased, became widely distributed, and was reported in every year until the bi-millennium. Its spread was remarkable - embracing Bretton, Skelmanthorpe and Denby in the east and south-east, Fangsett, Ingbirchworth and Hepworth in the south, Slaithwaite, Scammonden, Ryburn Reservoir, Ripponden and Fuddenden in the west, and Wyke and Fow Moor in the north. It was recorded at many places within these bounds, such as Emley, Almondbury, New Mill, Honley, Holmfirth, Blackmoorfoot, Findley, Birchencliffe, West Vale, Elland, Cromwell Bottom, Cleckheaton, Brighouse and several others, and the increase continues. An unexpected sighting was near Holme at an altitude of over 1200ft - perhaps a wandering individual. Small Pearl-bordered Fritillary Boloria selene (Denis and Schiffermiiller) In a period of more than 150 years the Small Pearl-bordered Fritillary has been reported only once, at Triangle, near Halifax, in early July 1983, by F. Murgatroyd. This occurrence, almost 40 miles (64 km) from the next nearest known locality at which it existed at that time, remains enigmatic. If it represents a wandering individual it is informative of the way in which colonisation of an area is possible. Pearl-bordered Fritillary Boloria euphrosyne (F.) Always rare in the area and so designated by Hobkirk (1859, 1868), the Pearl-bordered Fritillary was known in the mid- 19th century from Storthes Wood. On the authority of J. Varley, Mosley (1883) added “behind Castle Hill”, this presumably being the same as the “Huddersfield, very rare” of Porritt (1883) who cited the same naturalist. There are no further records. It probably became extinct before 1870. Dark Green Fritillary Argynnis aglaja (F.) Evidence for the occurrence of the Dark Green Fritillary in the Huddersfield area is tantalisingly vague and unsatisfactory. There are no 19th century records and Wattam (1936) Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 79 makes no mention of it. It was present at Seckar Wood, near Wakefield, about two miles outside the area, in 1942. In comments on the Lepidoptera of the Cawthorne - Bretton area, Seago (YNU excursion circular 1953) stated that there were two colonies in the Cawthorne district. No other reference to such has been found, but Seago was a knowledgeable lepidopterist. Other reports are also unsatisfactory. Collinson (1969) refers to two large fritillaries watched for half an hour at Copley near Halifax in 1969 by H. Archer and a companion but, while the Dark Green seems the most likely candidate, left their identity unresolved. Sutton and Beaumont (1989), evidently referring to the same incident though quoting a different year, give “Copley, two seen 1967 (per B.D.C.)”. The original report made no positive claim to identity. From nearby Beacon Hill there was an unconfirmed sighting of two individuals in 1985, again no more than an unproven possibility. Silver-washed Fritillary Argynnis paphia (L.) This species was recorded at Storthes Hall in 1847 by J. W. Dunning (Rimington & Beaumont 1996) and listed for Honley by Hobkirk (1859, 1868), who added “rare” in the 1868 list. Mosley (1883), on the authority of Inchbald, said only “Formerly at Storthes Hall but very rare”. Porritt (1883), on the same authority, simply said “Huddersfield, very rare”, obviously no more than a less precise reference to the same locality. It is not clear whether Inchbald saw this species at Storthes Hall subsequent to 1847 but it would be coincidental if Dunning, who arrived there in 1846, should have seen it on its last appearance. It is worth noting that Dunning made the comment on his manuscript, “The most common Argynidd”, which, although apparently not referring to Storthes Hall, is relevant to the status of the Silver-washed Fritillary at that time. On the basis of reports from his correspondents, Newman (1870-1871) recorded it as not uncommon in Northumberland, Durham and Cumberland (i.e. in the 1860s and probably earlier) and it was known, but rarely, in Scotland. For the first two of these counties, however, Dunn & Parrack (1986) cite only a few records, either undated or from the 1850s. At the end of the 19th century it was widely distributed, but rare, in Yorkshire which was then near the northern limits of its range in Britain. It may, however, have been more frequent in at least one part of the Huddersfield area than generally assumed. As Collinson (1969) reports, Herbert Spencer (1884-1949) always insisted that in his boyhood the Silver-washed Fritillary “could be regularly (our italics) seen flying in the Elland woods near his home”. Apparently this was never accepted by the YNU, much to Spencer’s indignation. Herbert Spencer was well known to, and was one of the finest lepidopterists ever met by, G.F. Extremely knowledgeable about their habits, the study of these insects was his enduring passion. He had built up a magnificent collection, and also had an encyclopaedic knowledge of Elland Park Wood alongside which he lived. He could hardly have misidentified such a conspicuous and distinctive insect, which is readily recognisable on the wing, and would never confuse it with the Dark Green or High Brown Fritillaries, neither of which has with certainty been seen in the Elland- Halifax area. The High Brown Fritillary is virtually unknown in Yorkshire except for possible strays. Furthermore the Silver-washed Fritillary is much more a woodland insect than the Dark Green Fritillary which favours wild, open areas rather than woodland. There is no valid reason for doubting the report of this excellent entomologist, which is here posthumously rehabilitated. A tantalising rider to this matter is a note in the records of the Halifax SS for 1927 “Fritillary seen in garden at Elland probably this species”. If Spencer was the observer, unless he saw this insect only at a distance, he would easily have made a positive identification. Its probable identity may have been based on a description by another observer. While unacceptable one suspects that this record may indeed be valid. Subsequent events lend credence to the validity of the Elland records. From the turn of the century the Silver-washed Fritillary retreated southwards, but there is more positive information on its persistence in the Huddersfield area than published summaries suggest (and more records for Yorkshire in the 20th century than generally supposed). The earliest 80 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire documented 20th century record for the Huddersfield area is for Skelmanthorpe in 1918. Although well known in a general sense there has been much confusion over the details, and about the exact site. Indeed published information even suggests that two sites were involved and three woods have been named. The correct facts are as follows. Morley (1919) recorded that in August 1918 “I saw Argynnis paphia in a wood near Skelmanthorpe, and Mr J. Hooper confirmed the record by taking one in the same place the following day”. This was noted as “new to the local list”, that is to the area around Skelmanthorpe. No name was given to the wood, clearly deliberately, but in his notebooks Morley is more explicit, noting that “I saw a fine male in the old garden of the ruined keeper’s house in Bank Wood on Aug 10th. 1918. Mr J. Hooper caught a female on the same ground the following day”. This is clearly the exact locality. However, in a report of the YNU Entomological section (Corbett 1919) B. Morley is recorded as reporting that J. H. Hooper took a specimen of A. paphia in Deffer Wood! To add to the confusion the YNU record book says simply “One, Aug 11 1918 Bankhall Wood, J. Hooper”, and the S.W. Yorks. Ent. Soc. record book says likewise! Bank Wood and Deffer Wood are almost 3 miles apart and Bank Wood is nearly 3 miles from Skelmanthorpe. The suspicion is that the finders of this collectable species deliberately gave the wrong locality. Bankhall Wood may also be a deliberately misleading name. No such wood is known by those from whom enquiries have been made, and no wood of this name appears on maps of the area dated 1850 and 1893, nor on recent maps. The Silver- washed Fritillary was recorded again in the Huddersfield area in 1929. Wattam (1936), who described it as “very rare”, saw three at Broken Cross, Almondbury in July 1929. This is more significant than would have been the sighting of a single individual. This important record escaped notice by all subsequent recorders. There was another find, at Emley Park in July 1941, inconspicuously recorded (Anon. 1942) but now well known. Although the finder appears never to have been named it was W. Buckley, who lived at Skelmanthorpe ( SWYES record book). There is also an ambiguous reference to this species by Seago who, in a YNU excursion circular (1953), said that it “occurs very rarely at Bretton”. There is nothing to indicate whether this refers to earlier finds - West Bretton is not much more than a mile from Bank Wood and less than three miles from Emley Park - or to a more recent occurrence. As no additional finds have been substantiated in the area, we take this to be a vague reference to earlier finds made there. The last authentic report of this species in that area is therefore for 1941 and this may have been only a vagrant. Nevertheless the Huddersfield area was one of its last outposts in Yorkshire in the 20th century and we believe that there is good evidence that it bred around Elland in the early years of the century, and possibly did so elsewhere. Speckled Wood Pararge aegeria (L.) Said by Hobkirk (1859, 1868), who gave no date, to have occurred at Honley, the Speckled Wood persisted in the area for some time thereafter and was known at Castle Hill Side, Almondbury, by James Varley, whose main period of activity was between about 1855 and 1870 (Mosley 1883). There is no reference to this site in the second edition of Hobkirk (1868) in which Varley is thanked for additional information on the Lepidoptera, but it is unwise to draw inferences from this. Hobkirk may simply have deemed it unnecessary to update his entry, or Varley could have recorded it after he did so. Before Mosley prepared his catalogue it had “entirely disappeared”. The two 19th century localities are in well- wooded parts of the area: many western and south-western parts were probably never suitable for the Speckled Wood during the period of recording. In spite of its disjunct distribution in the second half of the 20th century - very roughly south Yorkshire southwards, and western and north eastern Scotland - which inspired baseless speculations about its arrival in Britain before the end of the Pleistocene glaciations, the Speckled Wood was more widely distributed in the 19th century. Dennis (1977), like Newman (1870-1871) before him, concluded that before about 1850 it had a continuous distribution throughout much of Britain save for the north of Scotland. High moorland, such as much of the southern Pennines, was of course unsuitable. For Yorkshire, Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 81 Porritt (1883) simply listed 10 localities without commenting on the pattern. Most are in the south and include Huddersfield, “formerly” on the authority of Varley, this being the most westerly locality. Several others were less than 30 miles to the east and south of Huddersfield the nearest being “Wakefield”. Brady (1884) added Hemsworth 1882, and Rimington (1992) gives additional 19th century sites from the Doncaster area. The range of the Speckled Wood contracted in the late 19th century. Its disappearance from the Huddersfield area may either have been part of this national decline or the result of some local cause. That it evidently became extinct here before it declined, but not always to extinction, elsewhere in Yorkshire, hints at a local cause. It began to increase again nationally in the 1940s. To the south-east and not far outside the area, one was seen near Barnsley in 1944 by E. G. Bayford, the first he had seen there (Dearing 1945). Expansion continued, particularly in the 1970s. However, in Yorkshire, while it shared in the recovery in the 1940s and early 1950s, it remained essentially confined to the south. It then again decreased, and even disappeared from its long held stronghold at Wentbridge in 1970. (YNU Lepidoptera Committee 1967, Jackson 1980, Sutton & Beaumont 1989, Rimington 1992). By the late 1970s it was reduced to very few known sites. Jackson (1980) commented on the strangeness of this decline at a time when in most parts of England it was holding its own or increasing its range. Shortly thereafter it again increased. It did so around Doncaster in the late 1970s and 1980s, recolonising Wentbridge in 1987, and was described by Rimington (1992) as then being probably as common in that area as at any time since recording began. Indeed more colonies were known in the Magnesian limestone belt of south Yorkshire than in the past. Further north it was known for some years south east of the Harrogate area, then in 1 992 it penetrated, suddenly and deeply, in several places (Barnham et al. 1993). It continued to spread in the south of Yorkshire throughout the 1990s, showing a small advance westwards (Blakeley 1997-1999). It therefore for long occurred, and increased, just outside the Huddersfield area which, however, it was slow to penetrate. Its national extension of range in the 1990s is mapped by Hill et al. (1999). They suggest that in the past 100 years or so changes in its distribution can be attributed to changes in climate, but this fails to explain the decline in Yorkshire in the 1970s when it was increasing elsewhere. Not surprisingly their model was unable to explain distribution on the basis of climate alone, “whereas a combination of climate suitability and woodland availability was more successful”(! ) Equally unsurprising was their conclusion that “distribution of habitat, in addition to climate, may have been important in determining distribution”. One does not expect to find the Speckled Wood on moorland or in open fields. Subsequent to its disappearance in 1870 or earlier the Speckled Wood was not certainly seen again in the Huddersfield area for more than a century. There was an unconfirmed report of its occurrence in the Upper Shibden Valley in the mid-1970s (when it was at a low ebb in Yorkshire) (Cain & Baggaley 1994) and a definite record in the early 1980s when one was photographed at Birds Royd, Brighouse by M. Murgatroyd (Sutton & Beaumont 1989, Cain & Baggaley 1994). It was not recorded again until 1992 when a fresh female was photographed, and another was seen, in North Dean Wood, near Clay House. In May 1996 it was reported in Bank Wood, between Emley and West Bretton, by R. M. Sunter (Blakeley 1998 gives it as 1997), this being the most westerly of several sites in which it has been found since 1995 by Wakefield Naturalists. (At the bi-millennium it is widespread around Wakefield - R. Bedford.) In 1998 it was found at Greenwood, Denby Dale by D. Hemingway, and in 1999 at Almondbury by Stephanie Coughlan. In 2000 it was found at Cromwell Bottom (per B. D. Cain), Langsett (D. Manchester, who also received a report of another at High Hoyland) and, as late as October 23, B. and M.J.L. had one in their garden at Fixby. A significant spread to the west is indicated by finds at Blackmoorfoot, Colne Valley, by M. L. Denton and in the Ryburn Valley by M. Earnshaw. Suitable habitats are fewer than they are for the Large and Small Skippers which colonised the area more rapidly, and until recent tree planting began to rectify this situation, were virtually absent from the Colne Valley. Colonisation, however, is now underway. 82 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire In 2000 several were seen for the first time in the Greenfield area, west of the Pennines, by L. N. Kidd, just outside our adopted boundary. North of the area, single individuals were seen in 1996, 1997 and 1998 in the Heaton area of Bradford (Blakeley 1997-1999). Wall Lasiommata megera (L.) Of the Wall, Hobkirk (1859) was able to say only “has occurred at Honley”, while Mosley (1883) noted, on the authority of Inchbald, that it formerly occurred on Castle Hill Side (near Almondbury), but that by then it had “entirely disappeared”. One was reported from Hipperholme in 1868 (Halifax SS). It may never have been common in the area in that period but information is inconclusive. Formerly common in north-east England, it suffered a decline in the early 1860s and few were recorded there, sometimes at long intervals, until the early 1970s when it rapidly increased in numbers and spread (Dunn & Parrack 1986). Its disappearance around Huddersfield may have been part of this decline, whose limits are, however, uncertain, or independent of it. It evidently persisted, but became scarce, towards the end of the 19th century, in parts of S. Yorkshire, N. Nottinghamshire and Derbyshire, and apparently never became extinct around Doncaster (Rimington 1992), where it increased in the 1930s 1940s and 1950s. In Huddersfield it became extinct. Wattam (1936) had no knowledge of it there during the previous 42 years, and it remained unknown for some time thereafter. After an absence of more than 70 years H. Spencer provided “the first local record at Elland” in 1944 (Dearing 1945), and by 1948 it was said to be “now established at Elland” (Dearing 1949). It seems not, however, to have spread widely in the early stages of recolonisation, but may have bred unnoticed for some time a little further down the Calder Valley (up which it clearly advanced from the east) for B. and M. J. L. found it to be very common at Cooper Bridge in the late 1950s, and in 1959 or 60 it was found in the Ryburn Valley, a steep-sided upstream tributary of the Calder (YNU record cards). Further east it had evidently established itself in the Bretton and Cawthorne area by 1952 or earlier as it was noted as occurring there by J. H. Seago (YNU excursion circular 1953). In 1957 one was recorded at Denby Dale by R. Crossley, by which year it was said by J. Hooper to be “now fairly common” at Wakefield, just east of the area covered (Hewson 1958), a picture typical for the lower-lying parts of south Yorkshire in the late 1950s where it was said by Seago (1959) to be widely distributed and “definitely increasing” in many localities. In 1958 R. Crossley saw one near Kirkburton and received a report of another near Fenay Bridge. In 1960 he found it near Thunderbridge and in 1961 at Tag Lock, Elland, and in 1960 C. R. Haxby saw it at an altitude of more than 1000 ft near Queensbury, in the north of the area; “a more barren place for butterflies ... it would be difficult to imagine” (Bradford NS). In 1961 it was found at Hipperholme, at Birds Royd (Brighouse), and at Elland Wood Bottom (Cain 1990). It was also known in the Gunthwaite area before 1961 (J. H. Seago, YNU excursion circular 1961) and was seen there on the subsequent excursion (Flint 1962). By 1962 it had become more widely distributed in the Halifax area and was found at an altitude of more than 1000ft at Ogden Clough just outside it (Cain 1990). In 1963 Halifax SS records show that it had “moved into the district in numbers”, Halifax and Triangle being given as specific localities. By 1968 or earlier it had established a strong colony at Copley (Collinson 1969), and in 1969 it was seen at Cowcliffe (B. & M.J.L.). It was reported from Ovenden in 1976 (I. A. Hogg, Bradford NS). It continued to extend its range and by the 1980s it had not only been found in the south east (whence the colonists probably came) at Scout Dike and Royd Moor Reservoir near Penistone (D. S. & V. A. Ives), but had spread much more widely, being seen at Aspley (Huddersfield), Almondbury, Mount (Outlane), Holmfirth and Cromwell Bottom (D.S. & V.A.I.; B. & M.J.L.), and at Northowram 1984 (Bradford NS). The spread continued in the 1990s when it was widely recorded. It continued to establish itself in the Calder Valley, as at Sowerby Bridge and beyond, being described as “common on suitable grassland sites throughout Calderdale” in 1991 and 1992 (Cain 1992, Cain & Baggaley 1994) and was found in the Luddenden Valley. It may be via the Calder Valley that it arrived at Scammonden where it was seen by J. Dale in 1991. During this decade it was Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 83 also found at sites as widely separated as Kirkheaton, Grange Moor, Shelley, Upper Denby, Ingbirchworth, Yateholme, Digley, Holme (above 1000 ft), Meltham, Linthwaite, Bradshaw, Clifton, Bailiff Bridge, E. Bierley, Low Moor, Shelf, Queensbury, Lindley Moor, Norland, West Vale, Siddal and elsewhere (various observers). Numbers continue to increase in some places, as in the Denby Dale, Denby Delf, Shelley area where it was abundant in 1999 (T. Melling). On the western side of the Pennines, L. N. Kidd recorded it near Greenfield from 1977 onwards. It probably colonised this area from the west. It was known north of Oldham from at least 1946, and L.N.K. and B. Hodson saw it annually in the Medlock Valley between Oldham and Ashton-under- Lyne from at least as early as 1969. Hedge Brown. Pyronia tithonus (L.) The Hedge Brown, or Gatekeeper, was unknown in the Huddersfield area to 19th century entomologists. After being more widespread in Yorkshire, towards the end of that century it retreated to the south-east of the county and by 1967 was “restricted to very few localities” (YNU Lepidoptera Committee 1967). However, it spread in the 1970s, “displaying a great range extension” since 1973 (Sutton & Beaumont 1989). Limbert (1975) suggested that, habitat destruction and declining temperatures since about 1940 may have contributed to its contraction of range, but added a few localities in 1973 and 1974 - an early sign of revival? Subsequent expansion has coincided, at least in recent years, with a period of warming. A possible stray, but the first ever reported in the Huddersfield area, was encountered at Lightcliffe in about 1960 by M. D. Bridge (per. B. D. Cain). There was also an unconfirmed report of a colony near Mirfield during the 1970s (per. B. D. Cain), but proven colonisation began in 1983 when two were seen in the Hall Dike Valley (the Hall Dike flows from Meltham to the R. Holme) (YNU record cards but without finder’s name). In that year there were two sightings in the Halifax area (B. D. Cain) - where no more were seen until 1996. In 1985 J. Dale saw two at Royd Moor in the extreme south-east, and in 1987 D. S. and V. A. Ives saw it at Almondbury. In 1991 J.D. saw two more at different sites near Ingbirchworth, not far from the 1985 Royd Moor site. Further north it was reported in 1992 from Wyke (Bradford NS), and J.D. saw one at Lindley Moor. It was seen at Bretton Lakes in 1994 (L. Lloyd-Evans), and near Bretton in 1995 and 1996 (R. M. Sunter). In 1996 it was present at Upper Denby (D. Elliot) and J.D. found it again at Lindley Moor and, further west, encountered two at Scammonden. In that year it was seen at three places near the town of Halifax and at Clifton (B. D. Cain). Denby Grange Colliery and two sites near Horbury were added in 1997 (R.M.S.). There were several sightings in 1998 - near Bretton and at Clifton (B. & M.J.L.), at Scout Dike (D. Manchester), Birds Edge, near Ingbirchworth (S. Hey), Shelley (S. Graham) and Denby Delf (T. Melling). In 1999 S. Graham reported it as increasing at Shelley where he saw 10 on July 20, and where it persisted into September; B. Lucas counted 38 at Brickyard Plantation near W. Bretton, B. and M.J.L. saw it near New Mill, M. Dale reported it from Skelmanthorpe, and R.M.S. from two sites at Gawthorpe. It frequented Cromwell Bottom from 1997 to 2000 (B. D. Cain). Persistence at Clifton and several places in the east and south east was confirmed in 2000 (D. Manchester saw up to 50 at Ingbirchworth, and D.S. and V.A.I. saw 30 at Scout Dike) and new sites in that region - Emley, Deffer, Kirkburton, Stocksmoor - were added (B. & M.J.L.; D.M.; D.S. & V.A.I.; D. Knight). Further west, M. Earnshaw saw it in the Ryburn Valley. Even if the first individual seen, long before the main influx, was a stray, these widely scattered sightings, some at sites occupied for more than one year, reveal that colonisation of the area has been in progress for at least 18 years. This started later than expansion into the Sheffield area, further south and further east. There it began to extend into the vicinity of Rotherham in the 1970s, apparently especially in 1976. Few sites, all to the east, were known near Sheffield by 1980 (Garland 1981), but a great increase, especially in the east, subsequently took place (Whiteley 1993). The main expansion was between 1988 and 1990 with consolidation and further expansion in 1991 and 1992. In the Harrogate area, where it was previously unknown, there was initial colonisation of one small region in 1983, where it persisted until 1987 but was not seen in 1988 and 1989. However, it then rapidly colonised 84 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire eastern parts of the district, where by 1992 it occupied a large area (Barnham et al. 1993) - a pattern that stands in marked contrast to that which prevailed around Huddersfield (see also Ringlet). It will be interesting to see if it is now on the brink of a rapid increase in numbers and range in the latter area. The spread of the Hedge Brown in the Huddersfield area has been less dramatic than that of the Large and Small Skippers and is still in progress. Its slower advance may reflect its more demanding ecological requirements - hedgerows, the edges of woods, woodland paths and glades, and lanes, especially where brambles ( Rnbus spp.) are abundant. Such habitats are less common in the area than those frequented by the two skippers. It is also rather sedentary and seldom flies across open grassland, which militates against rapid dispersal. Meadow Brown Maniola jurtina (L.) Because it is so common in many places, the loss and long period of absence or great scarcity of the Meadow Brown in the vicinity of Huddersfield are some of the most interesting biological events in the recent history of the area. In the 1850s and 1860s it was “common in meadows”, (Hobkirk 1859, 1868) and continued to be so until possibly as late as 1870, prior to which it was reported as very common at Castle Hill and Almondbury, but then disappeared (Mosley 1883). Porritt (1883) said that “at Huddersfield it seems to be extinct” and notes that although he saw it in abundance at Almondbury “many years ago”, he had never seen a single individual there since. That it disappeared over an extensive area in the vicinity of Huddersfield is indicated by a comment of Morley (1902) on a single individual taken at Skelmanthorpe in 1901 - “an insect not seen here for twenty years or more”. In his manuscript list for Skelmanthorpe (1896-1908), he says “seems to have disappeared”, and was clearly struck by the local nature of this event as he added “Swarms at Edlington, Thorne Waste etc.”, sites that lie respectively just over 20 and 30 miles east of Skelmanthorpe. It was also long absent from that part of the Halifax area included here. Indeed Collinson (1969) says there were only nine records of single individuals from the entire extensive parish between 1863 and 1955. Of these the earliest and four others were from outside the area covered here. The relevant records (Halifax SS) were near Hipperholme before 1896 (Halliday 1896), Elland 1941, 1944, and Cromwell Bottom 1955 (see below). One of those outside the area was for Crimsworth Dene, near Hebden Bridge where W. Greaves found it in 1921, which elicited the comment that “this species does not seem to have been noticed for some years in the South Western part of the county” (Morley 1922b). Rimington (1992) gives information on a decline in some northern parts of England that began around 1860 but which seems to have affected neither the Doncaster area, with which he was concerned, nor Lincolnshire nor much of north Nottinghamshire, and Dunn and Parrack (1986) say it has “always been common and evenly distributed” throughout Durham and Northumberland save in the border region of the latter county. However, as around Huddersfield, the decline was well marked in the adjoining Barnsley area. Here Brady (1884) had regarded is as “common and universally distributed” (but until when?) yet the diaries of A. Whittaker (1898), cited by Rimington, record that he had “never seen it in the neighbourhood of Barnsley”. Here it then long remained unknown, or so rare as to be unrecorded. It could have disappeared some years before Brady wrote, and probably did so. Whittaker’s experience was shared by E. G. Bayford who was 15 years old in 1880 and who never saw the Meadow Brown in Barnsley until 1940 (see below). In the Bradford area, Butterfield (1906) recorded that “it is certainly not a local butterfly at the present time”. In the Huddersfield area it was never seen by Wattam (1936) in the 38 years before 1932 when he then encountered it at Berry Brow. It had, however, been seen in 1917 when C. Mosley (1917), son of S. L. Mosley, cited a mention in the Huddersfield Examiner of July 14 1917 by G. K. Crosland of its occurrence at Farnley. That this was deemed worthy of reporting to a newspaper is indicative of the unusual nature of the event. Mosley comments on its disappearance from that locality, as recorded in 1883, and adds “Its recurrence after nearly fifty years is remarkable”. Whether a small population then persisted at Farnley and Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 85 formed the nucleus of that seen there 16 years or more later by Wattam during its recovery cannot be established, but the area was one that he often frequented, which perhaps suggests that, if it persisted, it did so only in small numbers. Subsequent to 1932, and up to 1936, Wattam witnessed a definite recovery, seeing it again at Berry Brow (“a good many”) and at Primrose Hill, Farnley and Gunthwaite, a spate of sightings that heralded recolonisation of the area. Shortly thereafter a similar recovery took place south east of the area. Dearing (1941) reported that “Mr Bayford notes as his most interesting record” of 1940 “a battered specimen” of this species which he had not previously seen within the confines of the old Borough of Barnsley. E. G. Bayford (born in 1865) had long experience of the insects of the Barnsley area and, like Whittaker’s comments, his remarks confirm its absence, or great rarity, there for 50 years or more. Here it quickly re-established itself, for two years later it was described as “abundant” in the County Borough of Barnsley “and more especially just outside it” (Dearing 1943). With fluctuations in the interim it was common there in 1946 (Dearing 1947). In the Huddersfield area two Meadow Browns were seen at Elland by H. Spencer in 1934 (YNU record cards), a record not known to Collinson (1969). Spencer did not see it there again until 1941 (Anon. 1942) which elicited the comment that “this species is not usually seen in the [Elland] district”. In the meantime singles were seen at Low Moor by W. Barraclough in 1935 and 1936. These received the comments “a very uncommon butterfly with us” and “very unusual around Bradford” respectively (Bradford NS). Another was seen at Elland in 1944, in which year Wattam saw one at Newsome and several at High Hoy land (Dearing 1945). J. Briggs discovered a small colony between Low Moor and Bierley in 1950. It was common near Newsome in 1951 (Wattam, in litt.). GF, who had never previously seen this species in the Colne Valley, clearly recalls seeing a few individuals in a field in the valley bottom between Slaithwaite and Holme Mills, Marsden in 1951 or 1952. In 1955 it was recorded from Cromwell Bottom. It was found at Lepton in 1958 (A. Steele), R. Crossley netted seven at Mollicar Wood, Almondbury in 1959, and singles were seen by B. D. Cain in the Shibden Valley in 1959 and at Scout Hall, Shibden in 1960 (Cain 1990). Colonisation was clearly in progress, but the contrast with the situation to the south-east was still marked. In south Yorkshire and adjacent parts of Nottinghamshire and Lincolnshire, the Meadow Brown was at that time described by Seago (1959) as “common and widely distributed in all areas”. Although initial colonisers of the Huddersfield area were evidently sparse, and observers few, the earliest finds - “several” at Berry Brow (mid- 1930s) and High Hoyland (1944) - indicate a spread from the east and south-east. In the second half of the 20th century there was a remarkable increase in range and abundance, slow at first then increasing in tempo. It was apparently common and widely distributed in the Cawthorne-Bretton area by the early 1950s (J.H. Seago, YNU excursion circular 1953), perhaps a northward spread from the Barnsley area. The inhospitable Colne Valley may not have been colonised until the early 1950s, though early prospectors could easily have been missed. Localities in the Calder Valley such as West Vale, Copley, Triangle and others, upstream of the sites at Elland and Cromwell Bottom, where pioneers were seen in 1934, 1941, 1944 and 1955, were evidently not colonised until some time after 1969. In 1976 B. D. Cain found a small colony in the Shibden Valley and saw it near Bailiff Bridge (Cain 1990). It then rapidly extended its range and by the 1980s was widespread in the area, from Bretton and Scout Dike on the eastern and south-eastern fringes, to Almondbury, Honley, Gawthorpe Green, Cowcliffe, Lockwood, Paddock, and the Calder Valley. Further localities were added in the 1990s, from Grange Moor in the east, Hepworth and Yateholme in the south, Queensbury in the north, and from many others between them. It was established throughout Calderdale where by 1991 it was described as “usually our most common ‘brown’ butterfly” (Cain 1992). Here it was to be found on most natural grassland throughout the valley, as well as on hillsides in upland tributary valleys, a situation remarkably different from that of 50 years earlier when a single individual at Elland was regarded as particularly noteworthy. West of the Pennines where, if the suggestion made here is correct (see below), it 86 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire probably suffered extinction for the same reason as it did nearer Huddersfield, it was absent for an unknown period, but appeared in the Medlock Valley, outside the area considered, in 1975. It began to be seen in the Greenfield - Saddleworth area by L. N. Kidd in 1983. Colonisation here was evidently from a source different from that to the east of the Pennine barrier. Both older and more recent works give the impression that the Meadow Brown, perhaps Britain’s commonest butterfly, is virtually ubiquitous. However, large tracts of the uplands to the west and south-west of Huddersfield and Halifax are ill-suited to its needs. Although recorded elsewhere from ‘moorland’, the acidic, Eriophorum- dominated Southern Pennines do not provide suitable habitats. Much land is also too high. According to Emmet & Heath (1989) it occurs at 300 m (c. 980 ft) on Dartmoor but its altitudinal limit is a little lower in Scotland, on which basis alone it is hardly to be expected on the local uplands. In the adjacent Sheffield area, where it is now well entrenched in suitable habitats even in the city, and to the north, south and east of it, Whiteley’s map (1993) shows that large tracts of land in the west know it not. Occupied sites among the western uplands are essentially in valleys that offer shelter and larval food plants. Although M. jurtina probably entered these valleys from the east, exclusion from the inhospitable uplands is clearly shown. Whiteley’s maps (1993), incidentally, show how these moorlands exclude several other ecologically tolerant species that frequent their valleys. A map of its national distribution (Emmet & Heath 1989) reveals a large area of the Pennines from which M. jurtina is unrecorded. Moreover, as a single record from a valley site on the fringes of this area is given as much weight as a large population on a lowland site, such a map can give a misleading impression of status. During the re-colonisation of the Huddersfield area from the east and south east, the lowlands were inevitably occupied first, since when climate, topography and vegetation have decreed that it spread up valleys, with incursions into adjacent terrain only where this is suitable. Notwithstanding such facts, near Huddersfield the Meadow Brown sometimes ventures onto high ground. J. Dale has seen it regularly at between 900 and 950 ft at Scammonden and once on the reservoir embankment at 1000 ft, all in the sheltered valley. In 1996 he saw about 20 at c. 1025 ft near Broadstone Reservoir (near Ingbirchworth). There is a record from c. 1080 ft near Holme, while in 2000 G.F. encountered a colony on a steep, south- facing slope at between 1000 and 1050 ft on the the exposed Scapegoat Hill, and B. and M.J.L saw several just above 1065 ft at Pule Hill, Marsden. Such observations are in keeping with those of Barnham & Foggitt (1987) who reported colonies on sheltered verges at c. 1000 ft west of Harrogate, but whose map shows that it shuns the uplands thereabouts and scarcely penetrates into Upper Nidderdale. Climatic warming may currently be permitting colonisation of higher ground, but local moorland remains ecologically unsuitable. In general, recolonisation of the Huddersfield district at first took place slowly. However, having established itself in a locality it sometimes increased rapidly in numbers, as it did more spectacularly around Barnsley. This pattern is in keeping with the basically sedentary habits of the Meadow Brown which, having located a favourable site seldom emigrates from it (e.g. Brakefield 1982). Most observers agree. Ford (1964) notes that, although powerful on the wing, even 100 yards of unsuitable terrain is an almost complete barrier to it, and that it often turns back within 10 yards after entering an unfavourable habitat. Nevertheless it has colonised offshore islands - 69 of the 73 British islands considered by Dennis & Shreeve (1997) - and, as Ford (1945) himself records, has been caught on a light vessel seven miles offshore! Some individuals of course disperse, but mark and recapture studies suggest that these are few. Of distances moved by many individuals marked by Brakefield, the greatest was 320 m. (cf. the 94 km and 150 km. recorded for the Peacock and Small Tortoiseshell respectively). However, as Dennis & Shreeve (1997) note, such studies are not appropriate for recording between-habitat movements, and low frequency events involving greater distances are not easily detected. Around Huddersfield, early sightings after long absences were, in several cases, of single individuals - Skelmanthorpe, 1901, (seemingly not a forerunner of establishment), Low Moor 1935, 1936, Elland 1942, 1944, Newsome 1944 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 87 and, just outside the area, Crimsworth Dene 1921, Todmorden 1923 (?), 1924, 1927, Barnsley 1940. Others were of small numbers, as the two at Elland in 1934. Extinctions of the Meadow Brown elsewhere do not coincide with those around Huddersfield. Dennis (1977) includes it in a group of species “noted for both local extinctions and numerical depression in the early 20th century”, recovery of which seems to have taken place in the early 1930s, reached a peak in the 1940s, and was followed by a decline in the 1950s. No specific example is cited, and such a temporal sequence is completely different from that which took place in the Huddersfield area. Here a rapid decline to virtual or complete extinction took place some 40 years before that indicated by Dennis, and the main recovery occurred later. In a popular account of the butterflies of Bewdley, Worcestershire, Hickin (1987) says of this species “Now everywhere abundant, but was not to be found in Wyre 50 years ago” which is intriguing and does not suggest synchrony with the pattern to which Dennis refers. That the timing of extinction around Huddersfield was different from that indicated by Dennis (1977) points to a local and not an all-pervasive cause. Ringlet Aphantopus hyperantus (L.) In the 19th century the Ringlet was known from Farnley Mill and Storthes Hall pastures, both in the eastern half of the Huddersfield area, (Hobkirk 1859, 1868), but by 1883 Mosley could only note, on the authority of J. Varley, that it formerly occurred at Farnley. Long known from eastern parts of Yorkshire, its range had contracted by the end of the 19th century. It then long remained unknown in the Huddersfield area, as it was to Wattam (1936) and other entomologists. There was, however, an isolated, apparently unpublished, occurrence at Elland in 1934 where one was seen by H. Spencer, (YNU record book & cards). This was unknown to Collinson (1969) in whose Halifax list the Ringlet has no place. This presumably stray individual had no consequences for the future of this species in the area. The Ringlet was also reported from Great Park, Low Moor, by W. Barraclough in 1949, the single individual being described as “almost v. obsoleta ” (Dearing 1950). On the relevant YNU record card Hewson noted that this was only seen and not taken, and C. R. Haxby regarded it as highly suspect (Bradford NS). The reason is unclear as Barraclough was an experienced lepidopterist, must even have noted the reduction of the under-surface ocelli, and would hardly confuse even a variant Ringlet with any other species. The unexpected locality (listed as Royds Hall in Bradford NS records) may have engendered doubts. A valid record from this time and place would be interesting, but of scant significance in the history of this species in the area. In the early 1970s the Ringlet began to increase again in Yorkshire and to spread westwards. The earliest records for the Huddersfield area during this epoch were from near Bretton. In 1983 it was reported from Bretton Park (YNU record cards.) and in 1984 it was seen at two adjacent sites at Bretton (D. S. & V. A. Ives). There were no further reports until 1992 when J. Dale saw one at Lindley Moor, further west than any known 19th century site. In the late 1990s it continued to occur in the Bretton area, and was found in Bank Wood by R. M. Sunter. In 2000 one was seen at Shelley by S. Graham. Its spread was contemporaneous with penetration of the Harrogate area from the east, which took place very quickly. Between 1982 and 1985 it established itself over a wide area of the lower land there (Barnham et al. 1993), since when it appears to be colonising sites at higher altitudes (Blakeley 1999). While the Huddersfield area now lies within the westward (and northward) extending range of the Ringlet, which includes the Sheffield area (Whiteley 1993), progress has, as yet, been slow with little evidence of firm establishment save in the extreme east. Its pattern of spread resembles that displayed by the Hedge Brown and it is of interest that both species colonised the Harrogate area (to the north and somewhat to the east) at about the same time as they did the Huddersfield area. In both cases, however, colonisation around Harrogate was both more rapid and more extensive than in the Huddersfield area, a possible inference being that habitats in the latter remain sub-optimal. Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire Small Heath Coenonympha pamphilus (L.) Because it is so common, the history of the Small Heath in the Huddersfield area is particularly significant. It was clearly well known in the mid- 19th century when Hobkirk (1859, 1868) simply noted that it was common on dry banks, but in 1883 Mosley was unable to cite a single then frequented locality and only reported it, on the authority of J. Varley, as formerly known at Farnley. Porritt’s (1883) comment that it was distributed all over Yorkshire but was rare in some parts of the West Riding is further evidence of its status at that time, and this is borne out by what would otherwise be its surprising absence from the list of Halifax species (Halliday 1896). The situation there was confirmed by Collinson (1969) who says it was recorded in that area only five times up to 1945. Halifax SS records show that one of these was for Luddenden in 1866 and that the three others in the area concerned, were all post- 1940 - Elland and Copley in 1942 and Cragg Vale, Mytholmroyd in 1945. This emphasises its absence or great rarity over a long period of time. In the early 20th century it was also absent from, or very rare in, the Bradford area. Butterfield (1906) said it had “almost disappeared from the neighbourhood” and, equally categorically, Butterfield (1911) said “does not occur in this immediate neighbourhood”. Just south of Huddersfield, Brady (1884) noted that “this generally abundant species does not appear to be common around Barnsley”, and cited only four known localities but gives no dates. For the more distant, lower-lying Doncaster area, Rimington (1992) noted a lack of records for the 80 years prior to 1923, (an entry for 1919 has since been located in the record book of the S.W. Yorks. Entomological Society). This he found surprising but felt that it tended to confirm the poor 19th century recording there. This can hardly hold good for the Huddersfield area, where it had been regarded as common and where, had it not gone through a period of absence or great rarity, its presence would surely have been detected by entomologists of the calibre of Porritt, Mosley, Varley and others. Moreover Mosley specifically drew attention to species, of which the Small Heath was one, that, formerly abundant, declined dramatically or disappeared. Even in the Sheffield area not far to the south-east, however, Garland (1981) believed that it remained more or less constant in numbers and range in the 19th century, which suggests that the decline - to virtual if not complete extinction - was restricted to a relatively small area. Indications of what may be the first signs of recovery are records from two localities given in Morley’s manuscript list for Skelmanthorpe and vicinity (1896-1908) - Gunthwaite and (in the border area) Langsett. In his notebooks Morley recorded it in 1917 from “Woodhead Moors”, an imprecise locality, again in the border region, and, without dates. He also recorded it as common in some years on the railway side at Gunthwaite, and from grassy places on moors near Langsett, all of which are also mentioned in his updated manuscript. Its establishment, at least at Gunthwaite, in the first two decades of the century is implied. These sites are repeated in a YNU record book, that also gives Broadstone (near Ingbirchworth) in 1922 (H. D. Smart) and Coxley (undated). The last is also reported in the record book of the S.W. Yorks. Entomological Society as “rare” (J.H. = J. Hooper) and, although undated, was probably before 1919. If records from no later than 1908 and onwards mark the early stages of a recovery in the south and east of the area they were apparently paralleled by events further north, though precise dating is not possible. The recorder’s report for 1925 (Bradford NS) said “may now be considered to be well established in our area”. Whether this refers to the Bradford area as a whole or to the vicinity of Low Moor, where the recorder (W. Barraclough) lived, is not clear. The wording implies recolonisation sometime before 1925, by which time the situation was strikingly different from that which prevailed in the early years of the century. Especially if Low Moor was implied, arrival from the east is indicated. Information on its recovery was given by Wattam (1936). Comparing its status with that reported by Mosley (1883), he made it clear that “this is another of our local species which is now abundant, especially along the moorland areas at Meltham, Holmfirth, and the Chew Valley”. The last locality, on the western slopes of the Pennines, lies just outside the area covered but is continuous with similar terrain that falls within it. It is not known when Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 89 recovery began hereabouts but Wattam had already mentioned in the appropriate Annual Report of the HNS that the Small Heath was present on Holme Moss in 1926. It was to be seen in parts of the Colne Valley in the 1940s and early 1950s (G.F.), and R. Crossley noted it as abundant near Deanhead Reservoir, Scammonden, in 1957 (Hewson 1958). It seems to have been slower to recover in the vicinity of Halifax where the records for well scrutinised Elland and for Copley in 1942, and Cragg Vale 1945, were the earliest. It was, however, found in the Hebden Valley, outside the area, in 1946, 1948 and 1949, and was common there in 1955 (and had been seen at Todmorden in 1926), so it is possible that it spread into the western parts of the Halifax area from this region - a route unsuitable for most colonising species. The Cragg Vale population, further west than others, and remote from eastern sites, fits such a scenario. By 1955 it was well established on Norland hillsides (Halifax SS), a region that could have been colonised either from the north-west or from the Colne Valley and Deanhead - Scammonden areas in the south. It was found in the Shibden Valley in 1956 (B. D. Cain). By 1969 Collinson was able to say that “it is now strongly established through the [Halifax] area especially on hills and moors”, a truly momentous change from the situation at the end of the previous century when it had vanished from the area. It continues to be common there (Cain 1991 and subsequent reports), but Cain & Baggaley (1997, 1998) suggest that it may have suffered recent declines in some places near Halifax as a result of habitat loss. It has been common in the southern parts of the Huddersfield area from the 1960s to the time of writing and is known from many sites. Here B. and M.J.L. did not notice any obvious changes in status in 40 years of local acquaintance with this species until a possible decline around the bi-millennium. The Small Heath favours rough fields with short grass and an admixture of nectar- yielding flowers, grassy moorland, hillside slopes and upland doughs and, in contrast to most local species, shows a preference for the higher ground in the western part of the area. It has been found at about 1500 ft at Wessenden Head. According to Lees (1962, 1965), some Pennine populations at least are univoltine whereas those from southern England are usually bivoltine. Voltinism in the populations tested was under genetic control but could be modified by temperature. By artificial selection the percentage of individuals of a Pennine population producing two broods per year was increased over two successive years. Nevertheless a long emergence period, even in the upland parts of the area, indicates, as Blakeley (1998) suggests from observations elsewhere, that the pattern of emergence and overwintering may be more complex than generally supposed. Altitude, and therefore climate, doubtless exert an effect and, as the differences between Pennine and S. Devon populations described by Lees confirms, local selection occurs and different areas are populated by genetically different stocks. This may have been involved in some unexplained way in the 19th century decline in the area. It is impossible to prove that the Small Heath, previously common there, disappeared completely from the Huddersfield area in about the mid- 1860s, but its apparent demise was recorded and it was certainly brought at least to the brink of extinction. A few individuals may have persisted but there was no record for the next 30 years or more, and only one positive site, Gunthwaite, is mentioned in the next 20 years. Beginning towards the end of the first decade of that century, and showing a definite increase in the 1920s, it then recolonised suitable habitats, was common in many places by the 1940s and 1950s and seemingly widespread throughout the area in the last four decades of the century. Monarch Danaus plexippus (L.) An individual of this rare migrant was found at Kirkheaton in 1917 (Morley 1928), not 1927 as reported by YNU Lepidoptera Committee (1967) and Rutherford (1971). The confusion over the date arose because details were not generally available until the specimen was given to the Tolson Memorial Museum a decade after its capture. 90 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire Changing Patterns of Diversity In little more than 150 years there have been enormous changes in the composition and diversity of the butterfly fauna of the Huddersfield area (Figure 2). The number of species present at different times is indicated in Figure 4. There was a decline in diversity (here defined simply as the number of species: what is now often referred to as species richness), which included a dramatic episode of several more or less contemporaneous extinctions between about 1865 and 1870. The ensuing period of very low diversity then continued into the early decades of the 20th century. Similar faunal impoverishment occurred in nearby Bradford and, as shown by the absence, or extreme rarity, of such generally common species as the Meadow Brown and Common Blue, at least to some extent in the Barnsley region, but was nevertheless confined to a limited area. At the time of minimum diversity at the turn of the century, only 1 1 species, including the Red Admiral but not intermittent migrants, were present, and of these the Silver-washed Fritillary and Large Tortoiseshell were very rare. The fauna was then probably as impoverished as that of any part of England of similar size that offered a comparable array of habitats, either then or at any time in the previous or subsequent century. In the 20th century diversity increased slowly until, during about the last two decades, it did so more rapidly. At its point of lowest diversity the fauna consisted of only half as many species as it did in the mid- 19th century, but by the bi-millennium it included slightly more species, 23, than ever previously recorded. Increased diversity was not achieved solely by recovery of species that previously suffered extinction in the area: several disappeared, never to return. The Dingy Skipper, No of Species FIGURE 4. Changes in species diversity of the butterfly fauna of the Huddersfield area. Although a migrant, the Red Admiral, which has been present throughout, is included. Irregular migrants (Clouded Yellow, Painted Lady, and Camberwell Beauty) are omitted, as are the Small Pearl-bordered Fritillary and the vagrant Monarch, each recorded once only. Two of the 11 species present in 1900 were very rare. Some relevant conditions and events are indicated. A: smoke and soot levels high. S02 lower than later. B: Small reduction in industrial smoke emission. C: Marked reduction in soot and smoke. D: Reduction in S02, especially towards end of period. E: Large reduction in smoke and soot. F: Reduced acidity of rain. X and Y: Clean Air Acts. Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 91 Wood White, Pearl-bordered Fritillary, and somewhat later the Silver-washed Fritillary and probably the Large Tortoiseshell, were lost from the fauna. These losses were more than outweighed by the arrival of species never previously recorded, some of which became common and widespread. Species never recorded until the second half of, and often not until the late, 20th century, are Large and Small Skipper, Purple and White-letter Hairstreak and Hedge Brown. The Green Hairstreak, that also became firmly established in this period, had been recorded only once - before 1884. Excluding migrants and the once-only recorded Small Pearl-bordered Fritillary, of 27 species recorded during the entire period, no fewer than 13 (c. 48%) suffered extinction, (the Holly Blue twice and the Comma apparently three times) in less than 150 years. If the Red Admiral, which depends on migration but has been present throughout, be included, then 13 of 28 species (c. 46%) suffered extinction during this period. As six species were added to the fauna in the last 50 years or less, this masks the true extinction rate in the 19th century when, of 22 species (residents plus Red Admiral) present in 1850, exactly half were extinct by 1890. Of these, all save perhaps the Comma were still present in 1860, and Mosley’s (1883) account suggests that most extinctions happened between about 1865 and 1870 - a biologically cataclysmic event. Only nine species (c. 32%) persisted throughout, some tenuously at times. These are the bare facts: possible explanations are considered later. Faunal differences at the beginning and end of the 20th century are thrown into strong relief by noting that, including the Red Admiral but excluding intermittent migrants, the entire area could muster only 11 species in 1900, whereas in 1998 and 1999 S. Graham recorded 15 such species in, and in the near vicinity of, his garden at Shelley. In 1999 he recorded 13 species, plus the Painted Lady, in his garden alone, as did D. S. and V. A. Ives in their garden at Almondbury on the fringe of Huddersfield. They had previously done so also in 1992. Between 1983 and 1999 they recorded 16 species, plus the Painted Lady, in their Almondbury garden. S. Graham had earlier lived in Almondbury where he recorded 15 species, plus the Painted Lady, in his garden between 1991 and 95. In the last few years of the 20th century 17 species, plus the migrant Painted Lady, were seen in these two gardens. This is more than half as many again as were known in an area of 314 square miles in 1900! This recovery, and the rapid 19th century extinctions, are perhaps the greatest changes ever recorded in any local butterfly fauna in Britain. Some Aspects of Invasion and Colonisation Much attention has been directed to island populations of animals, and what is often called the theory of island biogeography has considered rates of extinction and invasion of islands, but similar events take place in localised mainland areas as the rapid changes in the butterfly fauna of the Huddersfield area make plain. These shift the emphasis to local invasions, often by species that are common nearby. There are no such barriers around this area as the sea that confronts potential invaders of islands, though the Pennines are such to some species in the south and west. There is largely unimpeded access from the east. Theoretically, invasions should be easier, and extinction rates lower, than on islands. Contrariwise, emigration should also be easier but, provided conditions remain similar on both sides of the border, should be cancelled out by immigration. Invasions, whether the result of deliberate or inadvertent human intervention, or of natural extensions of range, have generated much theory. Attempts have been made to ascertain what determines whether a given species will become a successful invader and what properties of the invaded environment facilitate, or resist, invasion. Attention has been concentrated on such matters as the intrinsic rate of population increase as a predictor of the ability to invade. It has been deemed “clearly important” to a colonising species that this should be large (Mac Arthur & Wilson 1967). High rates of population increase are supposed to reduce the initial risk of random population extinction, and of course to provide recruits for colonisation, but clearly are not always necessary. The Fulmar, Fulmarus glacialis, lays a single egg, does not breed until it is between 6 and 12 years old (mean 9.2 years), yet has been a spectacularly successful bird invader. 92 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire While it is easy to suggest things that may facilitate invasion by butterflies - mobility and good powers of dispersal, the production of several broods per year, long flight periods, wide ecological tolerance, polyphagous larvae and so on - these need not necessarily be effective or, like high fecundity, even necessary, and what is important to one species may be less so to another. Much nonsense, sometimes couched in “scientific” or mathematical language, has been written about invaders. Examples are cited by Fryer (1987, 1993a). In a consideration of the ecology of invading insects, Lawton & Brown (1986) found it difficult to be sure of hardly anything discussed and concluded that the “ only solution (our italics) to this empirical uncertainty is more, and better, theory”. We strongly disagree. Better understanding of what the animals really do may tell us far more. On the other hand we completely agree with Lawton and Brown when they say that it is not clear what makes an insect species a good invader. The biology of individual species is highly relevant to changes in status, and colonising abilities presumably differ between species, but details are not easy to elucidate. The Small Tortoiseshell and Peacock, that never became extinct though they passed through periods of great scarcity, are highly mobile - an attribute to which their persistent, if at times tenuous, presence may in part be due. The invasive abilities of the Holly Blue are self-evident. Quite different is the behaviour of the Hedge Brown, adults of which are less mobile, live in colonies, seldom fly over open grassland, and in several respects have more precise ecological requirements than some of the mobile species. It is only slowly establishing itself at the bi-millennium. A correlation of its habits with slow colonisation may seem self- evident - but is certainly suspect as a comparison shows. The Hedge Brown invaded the Huddersfield and Harrogate areas at the same time, but while it made slow progress around Huddersfield it colonised the Harrogate area both quickly and extensively (Barnham et al. 1993). Differences in habitat quality in the two areas may be involved, a possibility strengthened by the fact that the Ringlet shows the same difference in the pattern of colonisation of these two regions. Larvae of the Hedge Brown feed on a range of grasses, some of them common, so their food requirements are easily met, though they may have subtle micro-habitat requirements of which we are largely ignorant. Larvae of the Small Tortoiseshell and Peacock are more demanding food-wise and feed exclusively on nettles ( Urtica spp.) which may impose some restrictions. Thus, at least in theory, adult and larval ecologies may respectively impose restrictions on, and facilitate, dispersal and invasions, or vice versa. It is not easy to assess in which way a particular habit will affect invasive abilities. The Large and Small Skippers are both colony-formers - a habit that might be expected to restrain invasion - yet, as newcomers, they quickly occupied large tracts of the Huddersfield area. On the other hand the colony-forming Hedge Brown and Ringlet made slow progress in the same area - but rapid progress around Harrogate! Generalisation is difficult and dangerous. Only more intensive study of the habits and ecology of individual species (preferably in several localities) will clarify these matters. Invasion also involves coping with a series of habitat islands. To a woodland species a wood is a habitable island, and open terrain a barrier: for species of open habitats the reverse is the case. The efficacy or otherwise of such barriers differs much in different species according to their mobility, or tolerance, which adds to the complex of interacting factors involved. Local Extinctions and Recolonisation: Are Explanations Possible? Preamble Extinctions of butterfly populations are not rare. Colonies of colony-forming species not infrequently suffer this fate, and persistent contractions of range that extinguished several species over extensive areas are well known in Britain. The striking feature of the 19th century extinctions in the Huddersfield area is that they happened more or less simultaneously in several species, and in several species of moths, and were accompanied Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 93 by declines in the abundance of others. Such historical events in biology are often not easy to explain. As different species may respond in different ways to a common factor, and each is affected by a medley of influences whose importance differs from species to species, such a near synchronous decline of several species, in some cases to extinction, is particularly intriguing. It points to some potent deleterious factor. Of particular significance is that the events were localised: some affected species continued to flourish in nearby areas. Like invasions, extinctions have tempted speculation and model making. For butterflies, Thomas (2000) at least attempted to analyse actual situations, but presented the results as generalisations, only three species being mentioned by name, and these only in passing. He sought correlations between adult mortality and the decline and extinction of populations in fragmented habitats. His conclusion was that species of intermediate mobility were most susceptible to decline, and ultimately to extinction - the opposite of what one model had predicted - but that under some circumstances such species may survive best! He also concluded that a population “will not persist if the total mortality due to migration is higher than can be sustained by the reproductive output of individuals within breeding habitats” and even says this twice in rather different words ! Of extinctions around Huddersfield reported by Mosley (1883), all save those of the Comma and Holly Blue were of colonial species - but as most British butterflies are colonial little can be deduced from this. Furthermore, the Comma may have succumbed before the main extinctions, yet, being mobile, belongs to the category supposedly best fitted to survive - if other aspects of its biology are ignored. The most striking feature of these extinctions is the rapidity with which several species with a wide range of habits and ecological requirements disappeared essentially simultaneously. However it may be related to susceptibility to extinction, it is difficult to see how degrees of mobility may have been implicated here. Animals are continuously adapting to local conditions: natural selection is always operating to this end, and often a measure of phenotypic plasticity is also available. A striking discovery of biology in the 1960s, revealed by electrophoresis, was that a large amount of variation exists in most populations, which enables a species to respond rapidly to changing environmental stresses. Genetic adaptation is often cryptic, but dramatic visible evidence among the Lepidoptera of the Huddersfield area is provided by the development, increase, and spread of industrial melanism in more than 40 species of moths that took place from about 1880 or a little earlier, (perhaps 20 years earlier in the case of the Peppered moth, Biston betularia and a few others) and in some cases even later. Porritt (1906) gave a graphic account of these changes, some of which took place “during the collecting experience of many of our present-day lepidopterists”, and documented the case of the Mottled Grey, Colo sty gia multistrigaria in which the melanic form largely replaced the normal in ten years. Aspects of the classical story of selection of melanic and typical forms of B. betularia by bird predators on the basis of resemblance to background have been questioned in recent years (Majerus 1998) - which would not have surprised Porritt (1906) - but Cook (2000) regards the essence of the story as valid. Certainly, the spread of melanism in many species in the same area is a dramatic example of evolutionary change. Furthermore it is an example of change that took place convergently and simultaneously many times independently. This implies that it confers a strong selective advantage on several species, even if full understanding of the advantage is illusive. It also provides, many times over, particularly pertinent examples of the rapidity with which genetic changes can spread through a population. The existence of local races is further evidence of genetic adaptation. Differences in voltinism (the number of generations in a season) may be indicators of racial distinction and, as in the Green-veined White, may perhaps even reveal the presence of altitudinal races (or even more distinct taxa) within a small area. Seasonal forms, as of the three “Whites”, Pieris spp., provide further evidence, and are even more striking in some exotic species. The latter provide other impressive evidence. In the African Papilio phorcas a difference of a single 94 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire gene gives rise to females that differ in both colour and colour pattern, and the amazingly different mimetic females of P dardanus in different geographical regions are remarkable adaptations to local conditions. Such evidence suggests that the populations of those species that recolonised the Huddersfield area after an absence of several decades or, in the case of the Speckled Wood was recorded again after more than a century’s absence, are in subtle ways different from their extinct predecessors, and perhaps better able to cope with prevailing conditions. They may even exploit niches slightly different from those utilised by their predecessors. What caused the late 19th century declines and extinctions? Notwithstanding the possibility of different responses by different species to the same influence, the local extinction of several species and the drastic reduction in abundance of others in the same short period of time in the second half of the 19th century suggests that some over-riding common factor was involved. The species concerned belong to different families, frequent a variety of habitats, overwinter in different stages, have larvae that use different food plants, and adults with different flight periods. They have different predators and parasites at all stages of the life cycle, and often differ much in habits. Furthermore various moths disappeared in the same period. Had only a decline, or loss, and no recovery, been recorded, an apparently plausible explanation might have been habitat loss or deterioration. Deterioration indeed occurred yet these habitats were later recolonised by several species. After considering the national picture since about the 1830s, Heath et al. (1984) concluded that “the single main cause of change in the abundance and distribution of butterflies has been change and destruction of their habitats”. This is clearly inapplicable to some of the changes in the Huddersfield area. Here, after loss, in several cases recovery took place during periods when, as measured for example by loss of plant species, habitats continued to deteriorate, and also became more fragmented and reduced in availability. Moreover, the extinctions were confined to a restricted area, took place with great rapidity, and included several species that between them frequented a variety of habitats. To be convincing as an explanation, habitat destruction must be shown to have taken place in several habitats in a relatively small area, to have reached a critical stage in all cases essentially simultaneously, and to have led to permanent loss of the species affected. These demands cannot be met. Even on a national scale, 20th century fluctuations in the abundance and range of the Speckled Wood and Comma can hardly be attributed to this cause. While habitat destruction can be ruled out as an explanation of the extinctions, the deleterious effects of a particular factor or complex of factors that for a time reduced the suitability of the environment without destroying it is a different matter. If one can identify a factor that operated with such severity as to cause declines and extinctions but was later removed, leaving the environment sufficiently similar to its original condition to allow some of the exterminated species to return, it may be possible to suggest a cause for the declines and extinctions. Two candidates suggest themselves, climatic change and the deleterious effects of industrialisation and urbanisation, especially air pollution, and the associated acidic precipitation and deposition of soot. Climatic change is easily ruled out. How this could be responsible for local extinctions is difficult to imagine, and such events seem improbable. Some species that became extinct around Huddersfield continued to flourish in adjacent areas. Thus while the Meadow Brown became extinct in the second half of the 19th century it was apparently not affected in nearby Doncaster, from which area Rimington (1992) cites a series of records for the two decades on either side of the change of century, and in whose vicinity Morley noted that it “swarmed” at a time when it was absent even around Skelmanthorpe, in the east of the Huddersfield area. The 19th century declines and extinctions were also out of synchrony with those that sometimes occurred in other areas (see for example under Meadow Brown). This renders all-embracing explanations untenable and emphasises that local factors were operating. Ironically, at least temperature-wise, the climate seems not to have been adverse during Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 95 the critical period. Indeed, following a cold period of about 250 years - the so-called “Little Ice Age” - warmer conditions returned around 1830 and continued until the 1950s. This might be expected to have been conducive to the well-being of most British butterflies. On the other hand this general amelioration masks the fact that summer temperatures were generally low in the second half of the 19th century and fell during the period 1870-1895. Heath (1974) thought that this may have tended to cause contractions in range, and Heath et al. (1984) argued that the decline of certain species in northern Britain at that time may have been related to this but were nevertheless “certain that climate has not been the main factor in recent declines in abundance and contractions of range of many species”. In any case the extinctions in the Huddersfield area had already occurred by about 1870. That environmental degradation occurred in the area prior to, during, and subsequent to, the late 19th century decline of butterflies is well documented. The loss of many plant species provides concrete evidence of such changes. Important because it is so specific is the list of some 30 species of angiosperms typical of limestone regions of Yorkshire that also grew in the non-calcareous Huddersfield area before 1878 but no longer did so in the 1940s (Grainger 1942). Such a loss hints at a role for acidic precipitation, which long prevailed at high levels as a consequence of the Industrial Revolution. Cohen & Ruston (1925) described how soils in polluted Leeds became depleted of calcium carbonate. Moss (1901) had earlier noted the loss of several angiosperms around Halifax since 1775. The disappearance of lichens is a well known indicator of such adverse conditions, especially of high levels of sulphur dioxide, and the loss of Sphagnum and various angiosperms on the southern Pennine moorlands has also been convincingly attributed to atmospheric pollution, especially by oxides of sulphur and nitrogen. The period of extinctions and declines was one of increasing industrial pollution. The early 19th century saw an increase in the number of textile mills and the replacement of water power by steam engines that became dominant by the 1830s. As early as 1805 the Dartmouth Estate Book (cited by Crump & Ghorbal 1935) referred to the “mills and steam engines” at Farnley that “almost continually contaminate those pleasing features of picturesque beauty, water and air”. By 1850 many large textile mills had been constructed in the area (examples are illustrated by Giles & Goodhall (1992) whose fig. 274 shows how a large area south of the town of Huddersfield became dominated by textile mills in the mid- 19th century). Similar developments occurred in Halifax and adjacent villages and small towns, and considerable expansion continued in the 1860s. The number of power-looms in Yorkshire woollen mills increased almost eight-fold between 1840 and 1874, which gives some measure of the expansion. The inventory in Giles & Goodhall (1992) lists well over 800 textile mills, mostly built in the 19th century, in the area covered. Some were small: others very large. Increasing engine power meant increased coal consumption and higher levels of smoke pollution, as did an increasing human population, open coal fires being the norm. Contemporary illustrations of the many textile mills and the profusion of their smoking chimneys give a vivid picture of the situation that arose. In 1830 Read Holliday set up a small chemical works just east of the town and distilled ammonia from ammoniacal liquor wastes from the Huddersfield Gaslight Company. Such was the opposition to the pollution caused that within a few years it moved to Turnbridge where a huge chemical complex eventually developed. Small at first, it produced naphtha, creosote and later benzene, and by 1857 was one of the country’s largest coal-tar distillers (Jenkins 1992). Again there was local opposition to the pollution caused, and in the 1860s emphasis changed to the production of analine dyes. (By the end of the 19th century this chemical complex, eventually part of ICI, had a perimeter over two miles in length.) Other chemical enterprises also polluted the area. In 1867 the Rivers Pollution Commission reported that the Colne and Holme were extremely polluted and that four miles of the Ramsden Canal, fed from the Colne, was polluted and a great nuisance to part of the town. No fewer than 22 manufacturing chemists were found to be polluting local rivers. The 1871 report showed the much discoloured Calder at Wakefield to be suffering from pollution originating upstream that consisted of “many kinds of liquid refuse”. As evidence it 96 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire produced a letter written in 1 868 with water taken from the river, of which Woodhead (1939) gives a photograph. While water pollution has little direct effect on butterflies it reflects the general and extensive contamination of the environment in the mid- 19th century. Some of the offending enterprises doubtless produced volatile wastes and smoke. Coal mining was also widespread. In 1859 there were 116 coal pits in the immediate vicinity of Huddersfield, Halifax, Dewsbury and Holmfirth - which area then produced about a quarter of the coal mined in Yorkshire (Wray 1929). The number dwindled as shallow seams were worked out and mining moved east. Of at least 20 early 1 9th century pits in the Holme Valley between Holmfirth and Huddersfield none remained by the early 20th century. Coal mining contributed to environmental degradation by dumping waste, adding to air pollution by burning coal to power steam engines and sometimes firing spoil, and to water pollution by pumping out mineral-rich water. Other contributors to pollution were iron works, foundries, engineering works, dyeworks, and railways. What they believed to be the deleterious effects of such pollution on living organisms did not go unnoticed by naturalists. Nowell (1866) connected the loss of a considerable number of species of mosses in the vicinity of Todmorden (outside the area but nearby and very relevant) with “the super- abundance of factory smoke”. Moss (1901), commenting on conditions that had prevailed around Halifax for more than half a century, attributed the disappearance of various flowering plants to “the dense black smoke” that belched forth from “scores and scores again” of mill chimneys in SE Lancashire, which produced “great smoke drifts” that could be seen crossing the Pennines and mingling with locally produced smoke. ‘Black rain’ was frequent. As early as 1 842 the situation in Manchester, a prime source of such pollution, of which the Huddersfield area was a recipient (see below), provoked a leader on “The smoke nuisance” in the Manchester Guardian (May 28) which not only complained of the density of the smoke that was “palpable to the senses”, but alleged that trees were dying as a result of it and that even in open spaces in the city flowering shrubs would “not grow at all”. In the 1860s things were even worse in St. Helens where one “might look around for a mile and not see a tree with any foliage whatsoever”. Early observations on the detrimental effects of similar pollution in Leeds, only about 1 6 miles from Huddersfield, where in 1874 smoke was said to form “a dingy perpetual cloud”, are admirably summarised by Seaward (1975) who cites various pioneer studies and observations. Conditions were sufficiently bad as early as 1842 for meetings to be convened in both Leeds and Manchester to discuss smoke abatement and for an Inspector of Smoke to be appointed shortly thereafter. In the second edition of a book first published in 1912, Cohen & Ruston (1925) vividly described the effects of pollution in Leeds and provided abundant evidence, both observational and experimental, of the effects of air pollutants on plants. They showed how yields of various vegetables, wallflowers and grasses were reduced, most drastically in the most polluted areas where the growing of vegetables was pointless. Modern studies confirm these results and show how pollution reduces photosynthesis, curtails growth, causes crown thinning in certain trees and has other effects, though experimental results have not always been clear cut. Lor a summary and many references see DoE (1988). Mixtures of pollutants, such as oxides of sulphur and nitrogen, are usually more damaging than are either alone. Less is known about the effects of pollution on animals. Alstad et al. (1982) review information relating to insects. Many studies are inevitably retrospective and describe effects in terms that are correlative rather than causal. Relevant deleterious effects of sulphur compounds, however, include reduced flight and pollination activity in bees. Pertinent to present considerations is that silkworms react by a reduction in feeding rates and in activity in general, and suffer softening of the cuticle and delayed formation of cocoons. Dust particles, as in smoke and soot, disrupt the epicuticle. Delicate insects are more affected than large, heavily sclerotized forms, but highly absorptive dusts, that take up wax lipid secretions, can cause rapid desiccation and death even in large insects. Simultaneous exposure to different kinds of pollution in the past is a complicating factor. Our analysis of Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 97 the evidence suggests that smoke and soot may have had more adverse effects on butterflies than the more studied sulphur dioxide and oxides of nitrogen, but it must be remembered that the latter were also exerting an influence, perhaps synergistically, at the time of greatest production of smoke and soot. Surprisingly little attention has been paid to the possible deleterious effects of atmospheric pollution on butterflies in Britain, though concern has been expressed about severe declines of several species in major industrial areas in a number of European countries. Thus, in a discussion on the conservation of British butterflies, J. A. Thomas (1984) devoted less than seven lines to the topic. Although he noted that many Europeans, especially in Germany and Italy, blamed air pollution for the dearth of certain species near some industrial areas, and that a similar situation prevailed in the English Midlands, he concluded that “no more positive evidence links the two phenomena, and research is necessary to elucidate the matter”. Contrary to a generally dismissive attitude to the possible effects of pollution in Britain, however, Barbour (1986) considered the regional distribution of extinctions by comparing status in 1984 (Heath et al. 1984) with pre-1970 records - many of them much earlier than 1970. Many of the highest extinction rates were in areas close to major centres of industrialisation, the two highest being in south Yorkshire and Nottinghamshire, and north-east of London. In general, areas with most extinctions corresponded to a marked degree with those that comprised the zone in which lichens showed moderate to severe depletion- the zone in which atmospheric SO2 levels were 60pg m-3 or more. (In the classification followed, the zone in which the most serious depletion occurred was characterised by SO2 levels >170pg m-3 ). Higher levels in fact long prevailed in some places, including the Huddersfield area. In 1962-1963, by which time local levels of SO2 had declined, the winter mean at Holmfirth was 263pg nr3 and in 1969-1970 was 188pg nr3. Moreover, on a substantial number of days it exceeded 500pg m"3 (Ferguson & Lee 1983). Barbour (1986) also considered the effects of “dry deposition” of sulphur, resulting from atmospheric SO2 (not to be confused with “wet deposition” - “acid rain”) and found a strong correlation between extinctions and the amount of sulphur deposited. Indeed the correlation accounted for 79% of variance in number of extinctions. The Huddersfield area lies in the highest of the five zones of annual sulphur deposition, >4g mr2. More recent figures (DoE 1990) show that it lies in the area (E. Lancashire, S. Yorkshire and the N. Midlands) with the then highest sulphur deposition in Britain, - 4.0 - >6g m 2 per year - and also in the region receiving among the highest levels of oxidised nitrogen species. Average figures are not necessarily the most significant. Deposition is episodic and as much as 30% of the acidity deposited in a year may be received on <5% of wet days. Occult deposition (in mist and fog) of sulphur and nitrogen is also important and the amounts deposited in this way were often underestimated in the past. Notwithstanding the comment of Pollard et al. (1995) in one of the few papers to make even passing reference to the topic, that Barbour (1986) suggested that “atmospheric pollution (smoke and sulphur dioxide) may have played a role in the nineteenth-century contractions of range of butterflies”, he referred only to SO2, and made no reference to smoke - an important distinction (see below). Although he does not refer to the effects of industrialisation, Turner (1986), who considered the influence of climate on butterfly distribution in Britain, noted that in relation to their climates the industrial areas of Yorkshire and Lancashire are exceptionally unfavourable. (He used data on the maxima of species distributions between about 1910 and 1982.) What he did not know, however, was that the pre-industrial butterfly fauna of the Huddersfield area boasted twice as many species as it did after the 19th century extinctions. Both facts are in keeping with the suggestion developed here that pollution was a prime cause of the local extinctions. The effects on animals of particulate pollutants, especially soot which was formerly deposited in large quantities, is poorly understood. Soot includes toxins such as heavy metals. A pointer towards the possible deleterious effects of such toxins is that several freshwater crustaceans have been found living in more acidic conditions in pollution-free Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire areas, such as the Hebrides, than those in which they have ever been found in Yorkshire. Furthermore, they were doing so even though other conditions - more ‘dilute’ water and lower calcium levels than generally prevail in Yorkshire - make life physiologically more difficult there. They may owe this ability to the absence of the toxins that accompany pollutants in Yorkshire (Fryer 1993b). Heavy soot-falls were for long a feature of pollution in the Huddersfield area, falls of hundreds of tons per square mile each year being the norm in some places. However, soot was not uniformly scattered. Just outside the area, Cohen & Ruston (1925) showed that, in the early 20th century, 539 tons per square mile fell on industrial parts of Leeds, but the city centre received only 243 tons, and as little as 29 tons was recorded three miles north of the city. Soot-fall in the four square miles of central Leeds was roughly one ton per square mile per day. Much of the soot produced in industrial and urban areas was dispersed. For example Cohen and Ruston calculated that in 1897 every 12 hours saw the discharge of 80 tons of soot into the air above the 16 square miles covered by the city of Leeds. That is 10 tons per square mile per day. Some of this would fall on the city; most would be blown elsewhere. During the soot-producing era, many industrial sites not only in Huddersfield and Halifax, but in the Colne and Holme valleys and such towns as Elland, Brighouse, Cleckheaton, Batley, Dewsbury, Horbury and Ossett, ensured a generous dispersal of soot, whose fall was affected by the vagaries of wind direction. More was received from the Manchester area (see below). The cleanest region was probably always the Denby Dale, Cawthorne, Skelmanthorpe, Bretton area - which is also biologically richest for other reasons. Smoke reduces the amount of sunshine and light that reaches ground level, and formerly did so to such an extent as to be detrimental to such sun- and warmth-loving insects as butterflies. Cohen & Ruston (1925) gave examples of the amount of sunlight and light excluded by smoke in Leeds. In 1907 the duration of sunlight in the centre of the city was reduced by 17% compared with that at Adel about 4 miles to the north - and the amount received at Adel must itself have been reduced. That it received its share of smoke is delightfully demonstrated by a still smoke-blackened pair of Raven heads in the corbel table of its Norman church, that thereby achieve a remarkable realism never envisaged by their medieval carver. Light is restricted even more by smoke. Cohen and Ruston found that over a period of 19 days in May and June 1910 an industrial area of Leeds received more than 40% less light than did Garforth 7 miles distant - and Garforth must also have lost light because of smoke. Unlike insects such as bumble bees that can increase their body temperature by shivering or substrate cycling, butterflies are markedly ectothermic animals. All stages are much affected by summer temperatures and sunshine. As they realise, both Turner (1986) and Dennis & Williams (1986) are only confirming the obvious when they show that multiple regression analysis reveals that, between them, two factors, sunshine and temperature, account for almost 80% of the variance of butterfly diversity. As Dennis and Williams point out, this hides a multiplicity of complications. Nevertheless, biologically (as opposed to statistically) these factors are of prime importance. Many butterflies use their outspread wings as solar panels to absorb the sun’s heat, and many fly, and feed, only in sunlight. Larvae of various species also bask in sunshine, which increases their metabolic and growth rates, and warming of the ground by sunshine, when its temperature may greatly exceed that of the air, increases the rate of development of both eggs and pupae. Screening by smoke has the opposite effect, and thus increases exposure time to predators and parasites. Reduction of sunlight by smoke, even by amounts less than those recorded by Cohen & Ruston (1925) must therefore have had deleterious, and perhaps seriously adverse, effects on butterflies in the Huddersfield area during the epoch of heavy smoke pollution. Statistically every 15 minutes per day of summer sunshine adds one species to the butterfly fauna to be expected in an area of 10 miles radius - the unit used here! - (Turner 1986). The average total of sunshine received in May, June and July in Bradford (the nearest convenient station) in 1951-1980, mostly after marked reductions in smoke levels, was 509.3 hours, or just over 5h. 32 min. per day. Although 15 min. is only about 4.5% of this Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 99 amount, its loss would be sufficient, statistically, to remove one species from the local fauna. Losses of sunshine of the magnitude measured by Cohen & Ruston (1925) - admittedly in the centre of Leeds - were clearly detrimental. The brief adult life span of non-hibernating British butterflies, often no more than 3 weeks, and sometimes less than generally believed, emphasises the importance of exploiting all available sunshine. If reduced sunshine did in fact affect the butterflies of the area it might be expected also to have affected day-flying moths. Perusal of Mosley’s (1883) catalogue shows that day- flying species were indeed disproportionately represented among those that suffered extinction or became very rare. About half the moths that fall into this category were day- flying - a much larger proportion than of moths as a whole. Although their disappearance, especially in the case of common species, would be more quickly detected than would that of all but the commonest nocturnal species, their preponderance is too great to be attributed solely to this reason. Those lost included the Forester, Adscita statices, and the Narrow- bordered Five-spot Burnet, Zygaena lonicerae, both of which formerly “simply swarmed” in some places but were reported as “now entirely gone”. Also affected were the Wood Tiger, Parasemia plantaginis, that perhaps just escaped extinction, and the large and conspicuous Fox Moth, Macrothylacia rubi, and Northern Eggar, Lasiocampa quercus callunae, of which the last was described as extremely abundant until about 1873. All these have males that fly actively in sunshine. These three are predominantly upland species that live where cloud cover is greater, and the amount of sunshine less, than in the lowlands, and may therefore have been particularly vulnerable to any reduction in sunshine by smoke. Moreover, larvae of the Fox Moth and Northern Eggar bask in sunshine. The effect and significance of warmth is nicely demonstrated by the two-year life cycle of the Northern Eggar as compared with the one-year cycle of its southern sib the Oak Eggar L. q .quercus. Other species that fly actively in sunshine and were apparently lost were the Small Yellow Underwing, Panemeria tenebrata, Mother Shipton, Callistege mi, Burnet Companion, Euclidia glyphica, and Small Purple Barred, Phytometra viridaria. So was the Cinnabar, Tyria jacobaeae, that flies both by day (when its warning coloration is effective) and at night. Its larvae too are often conspicuously exposed to sunshine. That day-flying moths suffered proportionately more extinctions than night-flyers strongly supports the view that reduction of sunshine by smoke was a significant contributor to the extinction of butterflies in the late 19th century. Thanks to its tar content, soot adheres to vegetation. As a contaminator of the food of butterfly larvae, and as a possible irritant or abrasive of their delicate cuticle, it may have had deleterious effects, but little information on this has been found (see Alstad et al. 1982). Smoke and soot, however, may reduce larval growth by impairing the quality of their host plants - as does SO2 For use in reproduction most female butterflies store in the abdomen, nutrients that have been accumulated by their larvae. If conditions for the larvae are sub- optimal these reserves are liable to be reduced and reproductive capacity impaired. Moreover, in polyandrous species males transfer nutrients, as well as sperms, in the spermatophores at mating. In the Green-veined White the material transferred at a single mating may exceed 20% of body mass. While not all of this provides nutrients to the female, the amount is significant, and, with intervals, the female may be mated several times. If males cannot produce full-sized spermatophores, or if the females receive too few spermatophores, they are unlikely to realise their potential fecundity. Such insidious effects may have contributed to extinctions. The location of the Huddersfield area is very relevant here, as is made clear by a comparison with the Sheffield and Doncaster areas. It abuts the eastern slopes of the Pennines at their narrowest point. On the western side, and to the south, lies the Manchester conurbation including Stockport, Oldham, Rochdale and other towns, west of which is Merseyside. Manchester was one of the earliest areas of development during the Industrial Revolution and for long continued to contaminate the atmosphere. (Even in 1970 it produced greater amounts of sulphur dioxide than anywhere else in Britain.) Prevailing winds are south-westerly. These carry polluted air from SE Lancashire, and formerly the 100 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire smoke so graphically described by Nowell (1866) and Moss (1901), across the narrow Pennine ridge into the Huddersfield area where it joins locally produced pollution. Such polluted air passes to the north of Sheffield and Doncaster, whose south-westerly winds come from a much less polluted region. On the fewer occasions on which westerly winds with a northerly component carry Lancashire air towards these towns it has to cross the wider High Peak region of the Pennines, whose altitude is somewhat greater than that of the uplands west of Huddersfield. In the past a greater proportion of the particulate soot, which travels for shorter distances than gases, would be deposited before it reached Sheffield, whose centre lies 32 miles from that of Manchester, than would be the case of that carried to Huddersfield only 21 miles away. Less smoke would also survive the longer journey. Wet deposition of sulphur is also inevitably reduced in the dryer country from Sheffield eastwards. Sheffield and Doncaster produced their own smoke and other pollutants but were regularly refreshed by purer air from the south-west: the same winds added to the pollution load around Huddersfield. Sheffield suffered some losses of butterflies, but the Dingy Skipper and Small Heath, for example, persisted throughout, and species such as the Orange Tip and Common Blue, that declined in the Huddersfield area, appear not to have been affected (Garland 1981). If pollutants, including smoke and heavy loads of soot, had the effects on butterflies suggested here, these facts are consistent with the late 19th century extinction of several species around Huddersfield and their continued presence in the Sheffield, and particularly the Doncaster, areas. As in other cities pollution levels differed in different parts of Sheffield. Industrial Attercliffe received increasing amounts of sulphate sulphur between 1929 and 1954 but Nether Green generally received less of this contaminant between 1934 and 1949 than Marsden in the Huddersfield area (Ferguson & Lee 1983). It may be more than coincidence that it was in the south-east of the Huddersfield area, least affected by smoke and soot, that the Common Blue and, for the most part, the Orange Tip persisted, albeit in small numbers, during the period of lowest faunal diversity. Both extended westwards when levels of these contaminants declined. That atmospheric pollution affected the Lepidoptera of the Huddersfield area around the time of the extinctions is shown by the spread of melanic forms among the local moths. The carbonaria form of the Peppered moth Biston betularia appeared and began to spread at about this time, and the appearance and increased incidence of melanics in many other species subsequently became a feature of the area. Some time lag between cause and effect was inevitable, so the increased incidence of melanism would be after the stimulus of pollution, particularly smoke and soot, began to make itself felt. Of species of which melanic forms later became common, Porritt (1906) noted the breeding in the 1860s of a single black individual of the Mottled Grey, Colostygia multistrigaria, and knew of two locally obtained black individuals of the Mottled Beauty, Aids repandata from at least as early as the 1850s, and suggested that the melanic form of this species “may even have been common long before we noticed it”. The increased incidence of melanic forms was clearly correlated with the deposition of soot. Melanism favoured survival. It may be argued that melanisation was not a direct response to pollution but an adaptation driven by predation. Even if this were so the spread of melanic forms would not have happened had the environment not been contaminated by soot. It was therefore a response to environmental change, brought about by pollution, even if the mechanism was indirect. The effects of pollution were deleterious: they rendered the moths conspicuous. Melanism granted protection against predators. In fact larvae of the carbonaria form of Biston betularia enjoy a 20% selective advantage over those of the pale form and, in contaminated environments, melanic forms of several species are hardier than the normal form which suggests some physiological advantage in these situations. It is not proven that acid rain or the dry deposition of sulphur have deleterious effects on butterflies, though, acting mostly on their larvae, this would not be surprising. Soot, with its attendant toxins, is a more obvious candidate. Although the amount of smoke had been reduced by the 1930s and 1940s, even in 1947 some 242 tons of soot per square mile were Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 101 deposited in the Borough of Halifax. In more comprehensible units this was 2.8 oz. per square yard (c. 95 g nr2). Vegetation in many parts of the area was still covered by a layer of soot, being particularly in evidence on evergreen leaves, such as those of the Holly, to which its tar content ensured tenacious adhesion. It is worth noting that the extinction of a species such as the Meadow Brown was not confined to the immediate vicinity of the many industrialised and urbanised parts of the area but occurred also in a place like the semi-rural Skelmanthorpe, which received its share of soot, the amount varying according to weather conditions. A matter we believe to be of great importance is the relative influence of two kinds of pollution. SO2 levels were lower in the 19th than in the 20th century, during which they continued to rise for many years, power stations being major contributors in some areas. Levels of smoke and soot, however, were very high in the 19th century, and dominated the pollution scene. Although extinctions in industrialised regions can be correlated with high levels of SO2, as they were by Barbour (1986), those around Huddersfield can just as readily be correlated with high levels of smoke and soot, though these now defy precise quantification. Timewise, the correlation is indeed better than with SO2 levels. In fact recovery of the butterfly fauna began, and continued, during a long period of increasing levels of SO2 (Figure 4). This suggests that smoke and soot had more deleterious effects than SO2 and oxides of nitrogen, a matter elaborated below. It was also to soot-blackening of the environment that many moths responded by becoming melanic. Furthermore, and highly significant, is that the localised nature of events is much more readily attributable to smoke and soot, than to gases such as SO2. The former disperse for relatively short distances: the latter more widely, as Scandinavian recipients of SO2 produced in Britain and elsewhere in Europe are painfully aware. Acid yielding gases may, of course have aggravated the situation. That pollution may have been a major cause of the extinctions need not deny the vagaries of climate a role in the decline. A period of particularly heavy pollution, with a persistent, dense smoke pall, in a windless but wet period, at a time of year when several species were particularly vulnerable, e.g. when egg-laying, could have had adverse effects. If this happened several times within a few years the effects may have been lethal. Of this however, there is no proof. Further east, warmer and drier summers already provided less stringent conditions, sunshine reduction by smoke, and soot deposition, were less, and somewhat lower doses of acid rain (a reflection of lower rainfall) may have been more easily counteracted by more alkaline soils. The 20th century recovery and late resurgence The recovery of several species, if not in such a dramatically synchronous manner as their decline, and the colonisation of the area by others, also suggests the involvement of a common factor or complex of factors. Both climatic amelioration and reduced levels of atmospheric pollution may have been involved. Although the vegetation remained less diverse than a century or more earlier, recolonisation showed that it remained suitable for many of the species that had become extinct. Air quality improved in some respects, but not in others, even before the end of the 19th century. At that time emission of black smoke from mill chimneys was restricted to 5 minutes per hour but, while this had some effect, the regulation was often flouted. Nevertheless, coal-smoke pollution had already passed its peak by about 1927, and Ferguson & Lee (1983) record that in several towns the “total deposited matter” in the gauges installed to measure this had fallen by 50% or more since 1914. Levels of smoke and soot deposition continued to fall following the gradual replacement of steam engines by electric motors, the implementation of the Clean Air Acts of 1956 and 1968, and later by a significant change from coal to other fuels and a decline in the textile industry. To those who remember conditions in the still recent past, today’s clean air is little short of a miracle. Notwithstanding reductions in the levels of smoke and soot, there was not much change in the deposition of sulphur as sulphate in the area between 1929 and 1954, even though by 102 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 1954 SO: levels had fallen nationally about 10% below the 1939 level (Ferguson & Lee 1983). In the next 25 years, however, SO2 levels fell rapidly in the area but, as indicated by the figures for Holmfirth in 1962/3 and 1969/70 cited in the previous section, they remained substantial. Levels then fell markedly. For example, in 1952 the mean SO2 concentration at Slaithwaite was in excess of 250 pg m 3; by 1978 it was about 80 pg m-3 and at Holme Moss in 1979/80 the mean concentration was 46.8 pg nr3. There was a slow net gain of five species of butterflies between about 1900 and 1980. The comings and goings of the Holly Blue and Comma also showed that environmental conditions were suitable for one or both of them at times in the 1940s, 1950s and 1970s (Figure 2). By the end of the century, by which time pollution by both soot and smoke and acid-producing gasses had been reduced, additional colonisers, would-be colonisers, and re- colonisers brought the total to 23 species (Figure 4). Various lichens have returned to the West Yorkshire conurbation as a direct result of these improvements in air quality (see Seaward & Henderson 1999 and references therein). The relationship between butterflies and air pollution is likely to be more complex than that involving these sensitive and directly influenced organisms, but improvements could hardly have been other than beneficial. We suggest that early returning species, such as the Meadow Brown and Small Heath, were helped by a reduction in the amount of smoke and lower levels of soot deposition. Conditions in some respects were better when species such as the Large and Small Skippers colonised the area than they were 30 years earlier, which may have facilitated their spread. (Note, however, that these two species were not present even in the 1840s.) Ferguson & Lee (1983) believed that, notwithstanding the fall in sulphur dioxide levels, the acidity of rainwater in the southern Pennine area was still increasing in the early 1980s, being “substantially more acidic than it was in 1950-1954” and that levels of sulphate in rural rainwater had not declined. This suggests that smoke and soot were more serious hazards to butterflies in the past than was SO2. More recently there has been a reduction in the acidity of rainwater. A conspicuous sight in the 1940s, now gone, was a brown plume of nitrogen dioxide issuing from a chimney in a chemical works at Deighton just outside Huddersfield - and adjacent to it the vegetation-denuded Kilner Bank. The latter is now wooded. Tree planting took place elsewhere and there was regeneration of Birch and Willows in some places. Whether most butterflies have yet benefited is questionable, though the Speckled Wood in the Colne Valley may be an exception. Such changes are, however, indicators of improved environmental wellbeing. The adverse effects of house building were to some extent offset by an increase in the number of gardens. On the debit side were increased emissions from motor vehicles, now a major source of oxides of nitrogen, which for some years deposited lead, particularly near roads. Indeed there was a sharp increase in the amount of lead in the environment from about 1940 onwards, due mainly to the combustion of Pb alkyls in petrol, but levels have now been much reduced. There was also an increase in organochlorines until their withdrawal, but levels were probably much lower than in predominantly agricultural areas. Even now, however, there is probably a greater use of pesticides in the area than was the case at the time of the 19th century decline. The climate of Britain showed a slight but steady amelioration between about 1 895 and 1950, and Heath (1974) suggested that this may have been conducive to range expansions of butterflies. As Dennis (1977) pointed out, when short periods of national expansion of several species are coincident, as of the Wood White, Comma and White Admiral that culminated in the 1940s, purely local events are unlikely to be the cause, and climate probably played a part. Nationally there were extensions of range and increases in abundance of several species in the 1970s and 1980s. Not all species that extended their range increased generally in abundance, while some that showed no expansion of range did so. Pollard et al. (1995) provide details and suggest that changes in weather were involved. An obvious candidate for association with the late 20th century colonisation and recolonisation of the Huddersfield area is global warming, the reality of which is now Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 103 widely accepted. A general increase in temperature in the last two decades of the 20th century is now an established fact, and biological consequences in Britain, such as breeding earlier in the year by birds, are already apparent. Particularly pertinent is that, of 35 non- migratory species of European butterflies for which adequate information is available, 63% have ranges that shifted northwards between 35 and 240 km during the 20th century, and only 3% shifted to the south. The southern boundary moved northwards in the previous 30- 100 years in 22% of these species, remained stable in 72%, and extended south in 5% (Parmesan et al. 1999). It is a reasonable inference that a warming climate assisted recent colonisations of the Huddersfield area, though, save in the case of the Comma, this was not a direct result of a northward spread. While climatic amelioration may have helped such species as the Hedge Brown, Ringlet, and White-letter Hairstreak to extend their range, (even though the last-mentioned species is suffering reduction of its larval food plant - Elms), recolonisation began by some species before global warming gained impetus. The Meadow Brown and Small Heath anticipated this phenomenon by 50 years or more. A prerequisite for recolonisation is a reservoir of the species concerned in reasonable proximity to the area involved. What constitutes “reasonable proximity” depends on mobility and other biological attributes and differs from species to species. Thomas et al. (1992) showed that several uncommon British species, none represented in the Huddersfield area, disperse slowly. The Silver-studded Blue, Plebejus argus was found only in habitat patches within 1km of other populated patches, and the Black Hairstreak, Satyrium (= Strymonidia) pruni, often takes two to three years to colonise patches within 300 to 700 m of occupied areas, the maximum one-step colonisation recorded being only 1 .4 km. Several of the species lost from the Huddersfield area, or that have recently arrived there, were long resident not far to the east. Their failure to colonise until recently seems not to have been because of slow crossing of barriers between habitat pockets as they often spread rapidly when expansion began. This suggests former exclusion by locally prevailing conditions rather than limitations of dispersal ability. Recolonisation by such a species as the Wood White, whose range greatly contracted in the 19th century, then expanded, but of which there is no reservoir within 80 miles ( c . 130 km), is unlikely at present even if suitable habitats are available. The ability of the Holly Blue to migrate has clearly enabled it to return and to spread rapidly within the area. Returning species may not only have different, and better adapted, genetic constitutions than their lost predecessors, but may even exploit slightly different niches. Possible examples have been described among the lichens currently recolonising the conurbation of which some of the area is a part. Some species are extensively occupying microhabitats from which they were previously not recorded (Seaward & Henderson 1999). Similar subtle changes may have taken place in the butterflies involved. Testing the hypothesis A great problem when seeking explanations of ecological changes is that often no control is available. A control exists in the case of the 19th century extinctions of butterflies in the Huddersfield area. Most of the extinctions were a local, not a national, or even a regional phenomenon. Some of the species that were lost over a short period continued to exist in areas less than 30 miles to the east and south-east. Comparison of events in the Huddersfield area with the recent history of the White Admiral, Ladoga Camilla (L.), a species never recorded there, throws light on the matter. The range of this species contracted and by 1 930 it was largely confined to Hampshire and a few sites in Essex. Between then and 1942 it greatly expanded. An explanation was sought by Pollard (1979). The expansion coincided with the longest run of warm Junes in the century up to that time. These high temperatures led to rapid development of late instar larvae and pupae and reduced the time they were available to predatory birds (the reverse of the effects of smoke). Pollard also believed that the abandonment of coppicing in woodland had increased the area of suitable habitat, which contributed to the spread. June 1972 was as cold as any previous June in the century yet the White Admiral was plentiful in 1973 at 104 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire Monks Wood (where its ecology was studied). Pollard suggested that, just as he believed habitat improvement and favourable weather were necessary for colonisation, “local extinction is likely only if habitat suitability declines, when the timing of extinction may be determined by a period of unfavourable weather”. No controls were available to test these suggestions. In the Huddersfield area extinctions and declines of several species occurred quickly and more or less simultaneously. As shown by subsequent events habitat destruction was not responsible: even though degraded, habitats remained suitable long after the extinctions. However, their quality certainly suffered temporary deterioration as a result of reduced sunlight, the deposition of soot, and perhaps the effects of other pollutants. A control exists that rules out climatic changes as having had an over-riding influence, and which appears to implicate the deleterious effects of pollutants. In nearby eastern areas subjected to the same climatic regime (though always drier) but where levels of pollution and soot-fall were lower, no such extinctions occurred. Recolonisation took place at a time when, in terms of plant diversity, habitats were less satisfactory than those originally frequented, and availability was less, but pollution had been reduced. In the case of some species this happened before global warming began. It was suggested that range expansion of the White Admiral was facilitated by habitat improvement. The only habitat improvements in the Huddersfield area were reduced levels of pollution, initially of soot- fall, and increased amounts of sunshine as the amount of smoke declined. Such facts strongly suggest that atmospheric pollutants, especially smoke and soot, played a major role in the 19th-century butterfly extinctions around Huddersfield, and that reductions in pollution levels subsequently facilitated recolonisation of what in other respects were somewhat degraded habitats. In support of this belief are: (1) The virtually simultaneous extinction of several species of butterflies in a restricted area is an unusual phenomenon, and cannot be related to climatic or natural ecological changes. (2) Barbour (1986) showed that extinctions of butterflies in Britain can be correlated with atmospheric pollution, and the dearth of certain species in industrial areas in Europe has been attributed to pollution by some continental workers. (3) 19th-century industrial development in the Huddersfield area was accompanied by atmospheric pollution, including sunshine-reducing smoke, and additional pollution, not least smoke and soot, was received from south-east Lancashire. Contamination from the latter source is generally acknowledged as having exterminated lichens, mosses, liverworts and various angiosperms. Within the space of a few years during this period several species of butterflies became extinct (as did certain moths) and others declined in abundance. (4) These extinctions were local in nature: the same species persisted in not too distant, less polluted, areas to the east and south-east. (5) The moths that suffered extinction included a disproportionately large contingent of day-flying species. Like sunshine-dependent butterflies, these may have been affected by reduced sunshine levels caused by a persistent smoke pall. (6) That industrialisation had marked effects on Lepidoptera is shown by the reaction of more than 40 species of moths in the area that produced melanic forms. (7) In relation to their climatic merits industrial Yorkshire and Lancashire were deemed “exceptionally unfavourable” to butterflies in the 20th century (Turner 1986). However, before the industrial revolution the Huddersfield area had twice as many species as it did at the beginning of the 20th century. (8) Timewise there is a better correlation of extinctions with high levels of smoke and soot than with those of SO2. (9) Smoke and soot are dispersed for shorter distances than acid-producing gases, and more readily explain the localised nature of the extinctions. Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 105 (10) Several species of butterflies recolonised the area as pollution levels, especially of smoke and soot-fall, declined. (11) Colonisation began when levels of smoke and soot-fall began to decline, but while those of SO2 were still rising, indicating that smoke and soot had more deleterious effects than SO2 and oxides of nitrogen. (12) Reduced smoke levels led to an increase in the amount of sunshine received, and therefore also to higher temperatures at ground level during periods of sunshine. (13) Colonisation and recolonisation occurred in the face of un- ameliorated environmental deterioration in terms of reduced plant diversity, and even though habitats had been reduced in extent and fragmented. (14) The recovery parallels a similar recovery of lichens that has been convincingly attributed to reduced pollution. (15) Towards the end of the 20th century, when atmospheric conditions were much improved, several previously unrecorded species colonised the area, some becoming common. The latter part of this phase may have been facilitated by global warming. Consistent with this belief is the tolerance by several freshwater crustaceans of high levels of acidity in pollution-free areas (p. 97). While climatic warming may have assisted recovery towards the end of the 20th century, it was clearly not the key factor in this process, and certainly not in its early stages. This explanation is not proven, but there are better controls than is usually the case when explanations of ecological changes are sought, and it appears to be consistent with the facts. No feasible alternative explanation has suggested itself. If the suggestion is correct, it is possible that similar extinctions took place in other industrial areas at the same time as they did around Huddersfield. Places near Manchester and Oldham suggest themselves. Events around Greenfield-Saddleworth hint at this possibility. Here, isolated from the rest of the Huddersfield area to all save the Small Heath and migrants, some common species were for long lacking. The area was exposed to the smoke, soot, and other pollutants of the adjacent Manchester conurbation rather than to those of the West Riding. Here the Wall appeared in 1977, the Orange Tip about 1981 or a little earlier, the Meadow Brown in 1983, and the Speckled Wood in 2000 (L. N. Kidd). Is it possible that, just as Mosley reported his observations in a local journal, some contemporary naturalist did the same for another area? If such a report can be found, either here or, perhaps a little later, in Germany, Belgium or Poland, where similar events may have taken place, proof would be provided. Relevant range changes in other insects Whatever factors have been responsible for the resurgence of butterflies may also have affected other insects. A study of the moths that suffered local extinction would be enlightening. Some have certainly returned. For example the Forester was seen near Flockton in the early 1950s (E. W. Aubrook & G.F.). The Common Field Grasshopper, Chorthippus brunneus, invaded the Huddersfield area from the east in the late 1940s (Fryer 1953). Whether it previously occurred in the area and disappeared at the same time as several species of Lepidoptera is unknown. No records appear to exist for this period. Certain dragonflies have also colonised the area in recent years, or are currently doing so. A survey from 1984 to 1991 showed the Brown Hawker, Aeshna grandis, to be established there (Lucas 1992). It was indeed described by Cain (1997) as “the most common large dragonfly” in the Halifax area, yet, as he remarked, in 1901 the occurrence of a single individual merited special mention. The Common Emerald, Lestes sponsa was also found in the Huddersfield area for the first time (Lucas 1992), and a breeding colony was located at Cromwell Bottom in 1996 (Cain 1997). The Ruddy Darter, Sympetrum sanguineum, a southern species in Britain, (but known north of the area before 1961) was also found for the first time, at two sites, in 1992 (Cain 1997). If an overall controlling factor is involved in recent range extensions, dragonflies, which have aquatic larvae, may be particularly 106 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire informative. Industrial pollution affected aquatic as well as terrestrial environments in the area. Colonisation of the area by the Emperor, Anax imperator, a southern species (Lucas & Lucas 2000) may, however, be related to global warming. Some Linal Observations and Reflections This history of a localised butterfly fauna dramatically reveals its dynamic nature. Within a century and a half several species declined to extinction and, while some never returned, others recolonised and became common and widespread. Others, previously unknown in the area, colonised it. Yet others have been present throughout its recorded history but have undergone fluctuations in abundance. The enquiry also revealed that some of the changes were confined to a relatively small area. Had a larger unit been the focus of attention, losses, gains, or changes in abundance in adjacent areas may have obscured a more dynamic series of local events. The exercise would have been impossible had not several generations of naturalists recorded their observations and had these not been preserved for posterity by a variety of organisations. In particular, had it not been for summaries by Hobkirk (1859, 1868), Mosley (1883) and Wattam (1936), and manuscript notes such as those of Morley, it would have been scarcely possible to reconstruct changes in the fortunes of individual species with the degree of precision that has been achieved. In particular, Mosley (1883) recorded what must be among the most remarkable and rapid series of changes in local populations of several species of British butterflies ever to be observed. It is particularly fortunate that, more than half a century later, Wattam (1936), using Mosley’s work as a landmark, recorded some of the subsequent changes. If it does nothing else, this overview emphasises the value of accumulating over long periods of time, records of the status of the species that occur in restricted areas. The commonest species are often particularly informative. Continuous monitoring can reveal changes as they take place and enable explanations to be sought as events unfold, perhaps aided by experiments such as habitat manipulation. It is also potentially valuable for conservation, and throws light on the biology and ecological requirements of individual species. While not unexpected, the changes demonstrate the effectiveness of the Pennines as a barrier to the dispersal of various butterflies. They also indicate that, save for the small area west of this barrier, whenever colonisation or recolonisation, of the Huddersfield area took place it was essentially from the east and south-east. The records also point to the importance of the Calder Valley as a highway for butterflies. Prom Wakefield westwards it is readily accessible from the south to the point where it receives the Colne, west of which urbanised, industrialised Brighouse is the only substantial barrier, and one that was less so when, for example, the Meadow Brown extended its range. It can still be circumvented to the south. Here, not only are habitats of various kinds provided by terrain adjacent to the river, but the valley hosts a canal and a railway with accompanying tow paths, embankments and open spaces, that to some extent at least form corridors even in built-up areas. West of Brighouse a variety of terrain extends to Sowerby Bridge, approaching which, although the valley becomes narrower and steeper-sided, various low-lying, sheltered habitats are available. Indeed, here it probably acts as a funnel whose steep sides tend to confine certain butterflies to the valley bottom. The increase in butterfly diversity towards the end of the 20th century, while known elsewhere, runs counter to the trend of impoverishment now often reported for many plant and animal groups. While offering no grounds for complacency, it at least suggests that in areas formerly subjected to environmental degradation and afflicted by heavy pollution, amelioration of these conditions, perhaps aided by climatic warming, provides the opportunity for at least some groups of animals to increase in diversity, and for some species to re-occupy habitats from which they earlier retreated, just as lichens are currently doing. Habitats may be less diverse than originally but can still support many of the butterflies that formerly occurred there. They can also accommodate species hitherto unknown in the area. Both recolonisation and establishment for the first time demand the presence of populations Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire 107 in reasonably close proximity, from which colonists can be drawn, though highly mobile species are to varying degrees free from this constraint. The situation shows how little we understand the biology of individual species. As recently as 1984 three experts (Heath et al .) suggested that without improved knowledge of their ecology and the means to apply it “another twenty years will see a further drastic decline” in our butterflies. The need for knowledge remains but their prediction has proved very wide of the mark. Many challenges remain. It may be too late to solve conclusively such problems as why the Large and Small Skippers, both of which occurred very near the eastern borders of the area, never colonised it (and were never even recorded as strays) in the 19th century, yet did so more than a century later. Others may be more tractable. For example, it may be possible to ascertain why the Hedge Brown and Ringlet were able quickly to colonise the Harrogate area but made slower progress around Huddersfield. Were the environmental consequences of industrial pollution still exerting deleterious effects around Huddersfield? Or is the somewhat sunnier climate of Harrogate (almost 14 minutes a day more than Bradford in May, June and July) sufficient to explain the difference? Another challenge is to explain why so many species colonised the area towards the end of the 20th century in spite of a continuing loss of habitats as human populations grew. Is it possible to determine the roles of global warming, continued recovery of the environment from contamination, or to pin- point other changes? There is also no species in the area about whose ecology and habits in relation to local conditions more cannot be learned. The enquiry has exploited a group of insects with a manageable number of species that has attracted continuous interest and the accumulation of information over a period of more than 150 years. This made it possible to consider the entire local fauna. While other groups may provide more difficult material, comparable studies, even of single species, would be valuable. Information on how they reacted at particular times, compared with what has been gleaned for butterflies, may cast light on some of the matters that remain unresolved, or about which we can make only guesses. We hope too that future recording might be encouraged with a view eventually to assessing changes in the butterfly fauna during the next 150 years. Such recording should aim to provide those who might assay this task with more comprehensive data than those with which we have had to work. Conclusions The butterfly fauna of the Huddersfield area has undergone many changes since the mid- 19th century. Probably always depauperate, it suffered a dramatic series of extinctions between about 1865 and 1870, remained much impoverished for the next half century, then slowly increased in diversity before doing so at an accelerating rate towards the end of the 20th century. Some, but not all, of the species that became extinct in the 19th century returned, and several species not previously recorded colonised the area, and in some cases became abundant. Many of the extinctions were restricted to a very limited area. Assessment of the evidence suggests that, contrary to what most British lepidopterists have believed, a prime cause of these events was the pollution that accompanied industrial developments in this and adjacent areas. They were not the result of environmental destruction (the most commonly suggested explanation of extinctions of butterflies in Britain) because, even though degraded, fragmented, and reduced in area, the habitats concerned proved suitable for a wide range of species that colonised or recolonised the area as levels of pollution declined. The evidence suggests that smoke and soot were more deleterious than sulphur dioxide and oxides of nitrogen, which may, however, have contributed synergistically. Because of the localised nature of the changes, events in nearby, less polluted areas, serve as controls against which to test this hypothesis. Such controls are seldom available when explanations are sought for historical changes in ecology. Soot and smoke do not disperse as readily as gases and their effects are more localised. Climatic warming may have assisted colonisation towards the end of the 20th century. 108 Changes in the Butterfly Fauna of the Huddersfield Area of Yorkshire Acknowledgements This investigation was made possible only by using information collected by many individuals over a long period of time. Historical records are documented as appropriate. Our indebtedness to those concerned is self-evident. We have also benefited from the labours of past recorders of the current status of Yorkshire Lepidoptera, such as Ben Morley, Ernest Dearing, Frank Hewson and Maurice Jackson. For more recent help we are grateful to many others. Philip Winter made available the YNU record cards of Fepidoptera and was very helpful, Margaret Hartley and Chris Yeates gave access to information in their care in museums in Keighley and Huddersfield respectively, and Michael Sykes and Brian Cain went to considerable trouble to provide information accumulated by the Halifax Scientific Society. B.D.C. also put his great knowledge of the butterflies of the Halifax area at our disposal. Ted Rimington and Howard Frost answered questions on specific points, Derek Whiteley procured some old literature, Prof. Terry Mansfield provided a copy of the book by Cohen and Ruston, and Bernard Skinner a copy of Morley’s manuscript list of c. 1928, while Vivien Fryer rendered invaluable clerical and other assistance. Many naturalists supplied records and other information for recent years or, in some cases, for earlier periods. Particularly helpful have been Roy Crossley, John Dale, whose notes on Findley Moor provided welcome continuity, Duncan Elliot, Steve Graham, Douglas and Vera Ives, Brian Fucas, Feonard Kidd, Derek Manchester and Richard Sunter. The last-mentioned also made available an unpublished manuscript on the status of butterflies in the Wakefield area, 1995- 1999. Others who provided information and records are S. Anscome, G. Baldwin, R. Bedford and other members of the Yorkshire Branch of Butterfly Conservation, Paula Booth (daughter of the late Arthur Steele), B. Brassington, B. Cocking, Stephanie Coughlan, M. Denton, S. and H. Dyson, N. Gill, S. Hey, J. Keenlyside, D. Knight, L. Lloyd-Evans, M. Mail, T. Melling, M. Pratt, B. Stanley, and R. Sykes. To all who helped we express our grateful thanks. Fastly we are particularly grateful to Harry Beaumont who read the manuscript with great care and understanding, made helpful comments, and saved us from one error! References Anon. (1942). Fepidoptera. YNU Annual Report. 1941. Naturalist 67: 22-24. Alstad, D. N., Edwards, G. F. and Weinstein, F. H. (1982). Effects of air pollution on insect populations. Ann. Rev. Entomol. 27: 369-384. Barbour, D. A. (1986). Why are there so few butterflies in Fiverpool? An answer. Antenna 10: 72-75. Barnham, M. (1997). Purple Hairstreak, Quercusia quercus. 1996. Yorkshire Fepidoptera Report. Argus No. 31: 22-23. 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Index to main entries for the species recorded Brimstone Camberwell Beauty Clouded Yellow Comma Common Blue Dark Green Fritillary Dingy Skipper Green Hairstreak Green- veined White Hedge Brown Holly Blue Large Skipper Large Tortoiseshell Large White Monarch Meadow Brown Orange Tip Painted Lady Peacock Pearl-bordered Fritillary Purple Hairstreak Red Admiral Ringlet Silver- washed Fritillary Small Copper Small Heath Small Pearl-bordered Fritillary Small Skipper Small Tortoiseshell Small White Speckled Wood Wall White-letter Hairstreak Wood White Aglais urticae 73 Anthocharis cardamines 63 Aphantopus hyperantus 87 Argynnis aglaja 78 Argynnis paphia 79 Boloria euphrosyne 78 Boloria selene 78 Callophrys rubi 65 Celastrina argiolus 71 Coenonympha pamphilus 88 Colias croceus 60 Cynthia cardui 73 Danaus plexippus 89 Erynnis tages 59 Gonepteryx rhamni 60 Inachis io 75 Lasiommata megera 82 Leptidea sinapis 60 Lycaena phlaeas 67 Maniola jurtina 84 Nymphalis antiopa 74 Nymphalis polychloros 74 Ochlodes faunus 56 Pararge aegeria 80 Pieris brassicae 61 Pieris napi 61 Pieris rapae 61 Polygonia c-album 76 Polyommatus icarus 68 Pyronia tithonus 83 Quercusia quercus 66 Satyrium w-album 67 Thymelicus sylvestris 57 Vanessa atalanta 73 60 74 60 76 68 78 59 65 61 83 71 56 74 61 89 84 63 73 75 78 66 73 87 79 67 88 78 57 73 61 80 82 67 60 Latest publication of the Yorkshire Naturalists’ Union ALIEN PLANTS OF YORKSHIRE GEOFFREY WILMORE The first comprehensive modem publication dealing with the alien plants of any major British county, detailing the flora (almost 1300 species) of Yorkshire’s wool waste dumps, railway sidings, sewage works, shoddy fields, industrial wasteland, dockland and oil mill sidings. Pp. 316, including 15 line drawings; published January 2000 Normal price: £15.00 (plus £2.00 per copy p.&p.) Reduced price for members of the YNU: £12.00 (plus £2.00 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Mr John A. Newbold, Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS THE FRESHWATER CRUSTACEA OF YORKSHIRE a faunistic & ecological survey GEOFFREY FRYER The crustacean fauna of Yorkshire reflects the great physiographic diversity of the region. Adopting an ecological approach, this book considers the Yorkshire fauna in relation to climate, topography, geology, soils and water chemistry, always keeping in mind that it is dealing with living organisms whose habits, requirements and physiological limitations determine exactly where they live. Matters covered include the ecological background; faunal assemblages and their regional attributes; an analysis of the factors that determine distribution patterns, many of which are mapped; wide geographical aspects; and conservation. Large areas, such as the Pennines, Howgill Fells, North Eastern uplands and the lowland plains are surveyed. So too are localised regions including Whemside, the Malham area, lowland heaths, and the largest lakes, as well as habitats such as upland tarns, seepages, cold springs, small lowland ponds, inland saline waters. Notes are given on every species recorded, including parasitic forms. Price £9.00 (plus £2.00 per copy p.&p.) Special offer to members of the Yorkshire Naturalists’ Union £7.00 (plus £2.00 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Professor M. R. D. Seaward, Department of Environ- mental Science, University of Bradford, Bradford BD7 1DP. Proceedings (in part) of the Yorkshire Naturalists’ Union Conference on ‘Recording and Monitoring Yorkshire’s Natural Environment’, Harrogate, 26 February 2000: The Composition and Distribution of Mixed Hedges in East Yorkshire - D. J. Boatman Recovery by Aquatic Vegetation in a Wolds Headstream Subsequent to the 1988-1992 Drought - R. Goulder Mapping Dryopteris submontana (Rigid Buckler Fern) to the Metre: Evidence and Implications for Fractal Distributions - Stephen Hartley and William E. Kunin Notes on the Distribution of Large-leaved Lime (Tilia platyphyllos Scop.) on the Magnesian Limestone in South Yorkshire - J. A. Newbould Bryophytes and Lichens of Colliery Spoil Heaps in Yorkshire - Peter Middleton and JeffLunn The Lichen Herbarium at Liverpool Museum - John Edmondson and Michael Palmer Published by the Yorkshire Naturalists’ Union 1 Editor M. R. D. Seaward MSc, PhD, DSc, FLS, The University, Bradford BD7 1DP Notice to Contributors to ‘The Naturalist’ Manuscripts (two copies if possible), typed double-spaced on one side of the paper only with margins at top and left-hand at least 2.5cm wide, should be submitted. Latin names of genera and species, but nothing else, should be underlined. S.I. Units should be used wherever possible. Authors must ensure that their references are accurately cited, and that the titles of the journals are correctly abbreviated. Volumes of The Naturalist for the years 1886 to 1975 have been retrospectively numbered 11 to 100 to accord with numbering before and after this period (see YNU Bulletin no. 3, pp. 21-22 1985); please cite these volume numbers in all references. Table and text-figures should be prepared on separate sheets of paper. Drawings and graphs, drawn about twice the linear size they are to appear, should be in jet-black Indian ink, and legends should not be written on the figures. Publishable manuscripts not conforming to the above requirements will be returned for alteration. Photographic Plates Readers of The Naturalist will have noticed that the number of photographic illustrations has increased in recent years. Good clear photographs, suitably captioned, to accompany articles or as independent features are always welcome. To encourage this development, a long-standing member of the YNU, who wishes to remain anonymous, has most generously offered to make a donation, the income from which would finance the publication of a plate or equivalent illustration in future issues whenever possible. The editor, on behalf of the YNU, wishes to record this deep appreciation of this imaginative gesture. © Yorkshire Naturalists’ Union — 2001 Single copies may be made of single articles in this journal provided that due acknow- ledgement is made and the copies are for non-profit making educational or private use. Copying of more than one article or multiple copying of a single article is forbidden unless special permission has been obtained from the Yorkshire Naturalists’ Union. Permission is granted for the use of brief quotations in published work provided that acknowledgement of the source is clearly stated, but the use of substantial sections of text and any illustrative matter requires the express permission of the Yorkshire Naturalists’ Union. Registered Office (for all enquiries, correspondence and payments): Mr John A. Newbould, Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS (tel: 01305-837384; email: newbould@aol.com) The Naturalist is issued free to individual members of the Yorkshire Naturalists’ Union and to Affiliated Societies. Institutions and Subscribers £24.00 Registered Charity No. 224018 113 THE COMPOSITION AND DISTRIBUTION OF MIXED HEDGES IN EAST YORKSHIRE* D. J. BOATMAN 38 Eppleworth Road, Cottingham, East Yorkshire HU 16 5YF Introduction The publication of a book on hedges by Pollard et al. (1974), stimulated a countrywide interest in the subject amongst both biologists and local historians. Much of the book is devoted to the research done in East Anglia by the authors themselves and they point out that early in their studies they noticed that “hedges tend to be either fairly pure, usually hawthorn or elm, or else rather mixed”. After postulating that “the older the hedge the more shrub species it would have” they go on to develop their well-known hypothesis that a hedge can be dated from the average number of species of shrub it contains in a 30-yard section. Whether or not one accepts this hypothesis, the value of a hedge biologically and from the viewpoint of wildlife conservation increases, other factors being equal, as the number of shrub species increases. In a region of intensive agriculture such as East Yorkshire, hedges are a particularly valuable resource for wildlife conservation but, so far as is known, the only work that has been published on them is that by Boatman (1975, 1980). The paper published in 1980 is based upon a survey of 117 “mixed” hedges, i.e. those containing 4 or more species of shrub, in East Yorkshire (VC61). In this survey, however, only presence was noted and no restrictions were placed on the length of any hedge. Since, from the viewpoint of wildlife conservation, the value of a mixed hedge is likely to vary with the amounts of the various species it contains, it was decided, in the mid-1990s, to undertake another survey during which the species would be assessed quantitatively. Of the 117 hedges included in the earlier survey only 75 were incorporated into the current one. Of the rest, 13 had been uprooted, 6 to make way for the bypass on the south side of Beverley, and 1 1 were considered to be too short. The remainder were discarded because they were composed mostly of a single species or were indistinct from an associated area of scrub. In order to make the current survey as complete as possible, many routes and tracks which had not previously been visited were explored. This resulted in the discovery of another 95 suitable hedges so the current survey is based on a total of 170 hedges. Methods As in the 1980 study, a hedge was defined as one side of a field and only those hedges containing four or more woody species were surveyed. The criterion used for estimating the amounts of woody species in a hedge was frequency. This was determined by recording the species present in each of twelve contiguous lengths of 10 metres (10 paces). This number of lengths was selected so that an average value could be obtained for the number of species present in 30 m lengths, approximately the length recommended by Pollard et al. (1974), for estimating the age of a hedge and therefore widely recorded. Ideally, the survey would have included herbaceous as well as woody species but in practice these would have been very difficult to record. More than one visit to each hedge would have been necessary and anyway many of the hedges were on the far side of a deep ditch. Once grasses, Cleavers ( Galium aparine ) etc. had become well established, it would have been difficult to get close enough to make a sufficiently detailed examination of the hedge bottom. Individuals of the genera Rosa, Salix and Ulmus are notoriously difficult to identify * Developed from a poster originally presented at the YNU’s Conference on “Recording and Monitoring Yorkshire’s Natural Environment”, Harrogate, 26 February 2000. Naturalist 126 (2001) 1 14 Composition and Distribution of Mixed Hedges in East Yorkshire accurately. Either more than one visit is desirable to acquire material in an appropriate condition for identification or material from mature stems (usually not available in hedges) is required. So far as Salix is concerned, features characteristic of Salix cinerea appeared to be conspicuous in most of the individuals encountered, the roses appeared to belong either to the section Caninae or the species Rosa arvensis and the elms appeared to be referable either to Ulmus procera or to U. glabra. More accurate identification of hybrids or microspecies was not attempted. The frequency data were analysed using TWINSPAN. Since the programme cannot handle data with quantitative values in excess of 6, the frequency values were grouped in pairs for the analysis. For wildlife conservation the value of a hedge is influenced by its physical structure and that of various associated features such as ditches and grassy strips as well as its composition. A scheme whereby various features of importance for wildlife conservation can be scored and the scores summed to give an overall figure for the conservation value of the hedge has been devised by Clement and Toft (pers . comm.) and this scoring system was applied to the hedges in the current survey. Overall figures are not presented, however, only information on some of the features. Results Types of location There are three main geomorphological areas in East Yorkshire: the postglacial lake bed of the Plain of York, the chalk Wolds and the boulder clay covered Plain of Holderness. Between the west-facing scarp of the Wolds and the Plain of York is a band of Jurassic rocks which, especially in the north, forms low hills. The distribution of the hedges according to these features was as follows: Plain of York: 58 hedges Jurassic beds: 15 hedges Chalk Wolds: 14 hedges Plain of Holderness: 83 hedges A map showing the number of hedges found in each 4x4 km grid square is presented as Fig. 1. The great majority of the hedges recorded were alongside unclassified roads (Table 1 ) but several were found alongside green lanes. After a roadside hedge had been recorded, the other boundaries of the field were often superficially examined from the road, sometimes through binoculars when the field was large, but these boundaries rarely consisted of a species-rich hedge. Of the 13 “field boundary” hedges recorded for Holderness in Table 1, most were alongside public footpaths. TABLE 1. Location of the mixed hedges recorded in East Yorkshire. Location Plain of York Wolds + Jurassic Holderness “A” class road 0 3 2 “B” class road 5 1 0 Unclassified road 50 19 58 Field boundary (not alongside road) 1 5 13 Species frequency and distribution In Boatman (1980) a list is given of the species that occurred in more than 2% of the total number of hedges recorded in the first survey. All were present in the hedges included in Composition and Distribution of Mixed Hedges in East Yorkshire 1 15 FIGURE 1. Distribution of the mixed hedges recorded in East Yorkshire. Each number represents the number of hedges recorded in a 4x4 km square. the current survey. Additional species, with the number of hedges in which they were recorded, were Euonymous europaeus (4), Fagus sylvatica (2), Populus x canadensis (3), Prunus avium (3), Prunus institia (11), Prunus padus (1), Ribes rubrum (1), Ribes uva- crispa (4) and Salix viminalis (1). Boatman also presents estimates of the percentage of hedges in which each of the species occurred in the Vale of York, on the Wolds and in Holderness. In the second survey the percentages differed to some extent from those obtained in the first so, in Table 2, the results of the two surveys are compared for those species which were represented in 10% or more of the hedges visited in the first survey and which, in the second, differed from the first by ±10% or more for one of the three regions. In Holderness the percentage occurrence for all seven species was within ±10% of that obtained in the first survey but in the Plain of York the percentage occurrence in the second survey differed from the first by more than ±10% for five of the species. For most of these the values obtained in the second survey were more similar to those obtained in Holderness, probably because 47% of the hedges were located on clayey soils compared with 35% for the first. Although most of the species shown in Table 2 are widely distributed in at least one of the regions, only two, Hawthorn ( Crataegus monogyna) and Blackthorn ( Prunus spinosa ), commonly occur with a high frequency. In Fig. 2 the frequency ranges of the 10 most common species are shown for Holderness and the Plain of York separately. Here the pairs of blocks above zero represent the proportion of hedges from which the species was absent and it can be seen that, where they are present, most of the species are generally represented by frequencies within the range 1 to 4. 116 Composition and Distribution of Mixed Hedges in East Yorkshire TABLE 2. Percentage occurrence of selected species (see text) in surveys undertaken in the 1970s (1) and in the 1990s (2), the number of survey 1 hedges used and discarded for Survey 2 and the total number of hedges in each survey. Plain of York Wolds and Jurassic Holderness Species Survey Survey Survey 1 2 1 2 1 2 Acer campestre 17 28 35 52 80 82 Alnus glutinosa 40 28 0 0 0 0 Corylus avellana 40 53 75 55 78 84 Fraxinus excelsior 70 76 50 65 76 69 Malus sp. 47 45 5 17 18 14 Prunus spinosa 50 76 80 86 95 99 Sambucus nigra 23 33 85 90 79 70 Number of hedges Survey 1 hedges included 21 17 37 Survey 1 hedges discarded 9 3 30 Total hedges in each survey 30 58 20 29 67 84 As shown in Fig. 3, TWINSPAN separated the 170 hedges into two major groups as follows: Group I - Field Maple ( Acer campestre ) and/or Elder ( Sambucus nigra ) present (140 hedges) Group II - Field Maple and Elder absent (30 hedges) Subsequent divisions of Group I were based principally on the frequencies of one or more indicator species but it so happens that the group of 24 hedges on the right hand side cannot be separated from that of 29 hedges on the left hand side, and from other groups, by the frequencies of less than 3 species. These two groups have therefore been amalgamated. Thus the groups adopted are as follows: Group 1 A - Elder common - abundant (frequency 3+) - 38 hedges Group IB - Hazel ( Corylus avellana) common - abundant (frequency 5+) - 49 hedges Group 1C - Frequency of Hazel and Elder low - 53 hedges Group II A - Alder ( Alnus glutinosa ) present ± Common Sallow ( Salix cinerea ) 13 hedges Group IIB - Common Sallow present ± Birch ( Betula pubescens ) - 17 hedges Within Group IB is a sub-group (Group IB 1) of 12 hedges with a relatively high frequency (3+) of Dogwood ( Cornus sanguinea). The distribution of some of these groups is shown in Figs 4 and 5. All of the Group II hedges were in the Plain of York, either just east of the River Derwent which, in East Yorkshire, runs mostly between Eastings 69 and 71, or in the neighbourhood of the Pocklington Canal, which extends east-west between northings 44 and 46 (Fig. 4). Most of the Group IB hedges were in Holderness between the eastern side of the Wolds and a line extending through Cottingham, Beverley and Leconfield (about 4 km north of Beverley) though there were a few in the Wolds and in the Plain of York amongst the Group II hedges (Fig. 5). The Group IB1 hedges were almost all in Holderness, mostly in the northern part of the range of the Group IB hedges (Fig. 4). Composition and Distribution of Mixed Hedges in East Yorkshire 117 % Hedges 100 Acer campestre 0 100 Quercus robur 0 100 Malus sp. ii IL j=l 1-4 5-8 9-12 Frequency % Hedges 100 Prunus spinosa 0 100 Fraxinus excelsior 0 l fLrJrl U1 Iran 100 Sambucus nigra 0 1-4 5-8 9-12 Frequency FIGURE 2. Frequency distribution of each of the ten species most commonly recorded in the mixed hedges of East Yorkshire. Open columns represent the % frequency with which a species occurred in 0, 1-4, 5-8 and 9-12 hedge sections in the Plain of York and filled columns in Holderness. 118 Composition and Distribution of Mixed Hedges in East Yorkshire FIGURE 3. Dendrogram showing the separation of hedges into 5 groups by TWINSPAN (the two unlabelled groups were amalgamated). The indicator species for each dichotomy are shown except for the amalgamated groups. Composition and Distribution of Mixed Hedges in East Yorkshire 119 FIGURE 4. Distribution of hedges comprising Groups IIA (black numbers), IIB (open numbers) and IA1 (shaded numbers, one in the Wolds the rest in Holderness). The remaining hedges, i.e. those in Groups IA and IC, were widely scattered in East Yorkshire, some from each group occurring in each of the geomorphological regions. As can be seen in Fig 2. the distribution of species differed considerably between Holderness and the Plain of York. TWINSPAN also, at the first division, separated species into those that occurred only or with relatively high frequencies in the Plain of York and those that were widespread or were absent from that region. Species which were recorded only in the Plain of York and the number of hedges in which each was found were Alder (13), Alder Buckthorn ( Frangula alnus) (4), Beech ( Fagus sylvatica) (2), Birch (11), Broom ( Cytisus scoparius) (5), Goat Willow ( Salix caprea) (4) and Mountain Ash ( Sorbus aucuparia ) (6). Although Field Maple is not represented in Group II hedges, it was recorded in 14 of those that were found in the Plain of York. The soils of this region vary from sands to clays and alluvial soils and Maple was recorded on all types. Nine of them, however, were on clays or alluvium. In Fig. 6 the distribution of the mean number of species per 30 m of hedge is shown for the Plain of York and Holderness separately. In each region the greatest proportion of hedges contain 5-6 species per 30 m and the percentage occurrence declines progressively above and below this value. Structural Features Structural features were recorded for only 123 of the 170 hedges that were surveyed. In Table 3 the features that most clearly represent the structure of a hedge are presented. 120 Composition and Distribution of Mixed Hedges in East Yorkshire The distribution of hedges comprising Group IB (black numbers) and of those containing Holly (open numbers). “Dendroid” hedges are those without branches on the lower part while “rectangular” hedges have branches almost to ground level. Some of the hedges, generally those with Blackthorn, had spread on to the roadside verge and these are recorded as “with outgrowths”. Presumably the value of the hedges for wildlife conservation increases from left to right and top to bottom in the table. TABLE 3. Landscape features of 123 hedges in East Yorkshire. Hedge form Height (m) <1 m 1-2 m 2-4 m >4 m Dendroid in section 6 23 11 4 Rectangular in section 2 31 22 3 With outgrowths 0 3 11 7 Totals 8 57 44 14 Other features which might be particularly important for wildlife are the presence of mature trees and the presence of a ditch. One or more mature trees were present in 36 of the hedges and a ditch was alongside 60 hedges. Most of the hedges surveyed were complete. Gaps extending for 10% to 30% of the total length of the hedges, i.e. the side of the field, were present at only 6 sites while gaps Composition and Distribution of Mixed Hedges in East Yorkshire 121 % Hedges Species/30m of hedge FIGURE 6. Percent frequency distribution of hedges with means of 2-9 species per 30 m in the Plain of York and in Holderness. 122 Composition and Distribution of Mixed Hedges in East Yorkshire extending for more than 30% occurred at only 4 sites. The gaps were ignored when recording the species present, i.e 120 m of complete hedge were recorded. Discussion It is now well established that there is an association between species-rich hedges and ancient countryside features such as old field boundaries, old trackways, former woodlands etc. (Hoskins, 1955; Pollard, et al., 1974; Rackham, 1986; Muir & Muir, 1987) and this lends weight to Pollard, Hooper and Moore’s hypothesis that it is possible to estimate the age of a hedge from the number of species it contains in a unit length. The hypothesis is supported by a number of authors, notably Rackham (1976, 1986) but, in their comprehensive review and analysis of the subject, Muir and Muir (1987) point out that “The theory of hedgerow-dating assumes an initial single-species planting” but that “evidence exists in abundance that old hedgerows were commonly planted as mixed hedges of several species”. This, they understandably claim, invalidates hedgerow-dating as a practical technique. No attempt has been made to conduct extensive documentary research into the relationship between the East Yorkshire hedges and ancient countryside features but, from information and maps published in the Victoria County History, Allison (1976) and Neave (1991) and from O.S. maps, it has been possible to relate 93 of the East Yorkshire hedges to ancient features. Sixteen are associated with former or existing woodlands, 8 are along or within the boundaries of mediaeval parks, 14 bound green lanes or old trackways, 15 form the boundaries of ancient enclosures and 32 are parts of parish boundaries. However, 4 are on land which, before 1755, was an open field and 12 are on land which was common pasture until, at the earliest, the beginning of the 17th century and probably until the middle of the 18th century. If hedges were formerly planted as mixed hedges, it is likely, as suggested by Muir and Muir (1987), that species growing locally were used. Thus the hedges existing today might provide information about the vegetation growing in the locality at the time they were created. The accuracy of such information, however, depends upon the extent to which the current hedges resemble those originally planted. As shown in Figure 2, the only species that are widely distributed within the great majority of the East Yorkshire hedges are Hawthorn and Blackthorn but it seems unlikely that these species would have been used mainly and so consistently in the original plantings with only occasional representatives of other species. In 1963 a long-term experiment was established at Monks Wood Experimental Station to investigate the extent to which the composition of single species and mixed hedges changed with time. Altogether 44 hedges each about 27 m long were planted and they have recently been re-examined by Downing and Sparks (1999). In many of the hedges Downing and Sparks were able to distinguish the original stems from those that had grown more recently and they claim that the most invasive species have been Hawthorn and Blackthorn. They also claim that “survivorship” of the original stems has been greatest for Hawthorn and Maple though their data show that it has been almost as high for Hazel. Unfortunately, however, these hedges have been severely neglected since they were established and had grown to a height of between 6 and 9 m. It must also be borne in mind that the hedges were growing in close proximity so there was a variety of seed available nearby for cross invasion. In 1971 Pollard determined the species present in each 5 m section of a 1105 m length of Judith’s hedge which is thought to date back to the 11th century. In 1998 the hedge was re- surveyed by Garbutt and Sparks (1999) who claim that there were sufficient permanent markers for them to be able to repeat Pollard’s procedure. Garbutt and Sparks report that since Pollard’s survey part of the hedge had been neglected and had grown to a height of 4-5 m. Along this part there had been a decline in the mean number of species per section whereas in the remainder, which had been regularly flailed, the number of species had remained more or less constant. Composition and Distribution of Mixed Hedges in East Yorkshire 123 Thus there is evidence that the composition of a hedge can change over a short time-span relative to those hedges discussed here, at least when it is neglected and there are seed sources of various species closeby. There is very little information about the management of hedges in old documents dealing with agricultural practices in East Yorkshire though Latham (1794) mentions that the “vile custom of heading them at the height of three, four or even five feet prevails in many parts”. Perhaps the best indication that the composition of the East Yorkshire hedges bears a relationship to that at the time of planting is the very localized distribution of some of the groups extracted by TWINSPAN but it is possible that the frequency of at least some of the species has changed. It can be envisaged, for example, that some individuals died, leaving gaps, and that the gaps were subsequently filled with Hawthorn or Blackthorn. In this context there is a striking resemblance between the distribution of Groups IB and II hedges and the distribution of woodland at the time of the Domesday Survey as presented by Darby and Maxwell (1962). Here, in a map of East Yorkshire, the majority of the Holderness woodlands, including all of the larger ones, are shown occupying a zone between the Wolds and a line from Cottingham through Beverley to Leconfield while those of the Plain of York, though more widely scattered, occur predominantly in the western half of the region. The Groups IB and II hedges might therefore provide information on the type of woodland that existed at the time of the Domesday Survey. This is particularly important for East Yorkshire because, although much is now known about the distribution of the various types of woodland in Britain as a consequence of the National Vegetation Survey, not a single East Yorkshire woodland was investigated for the survey (Rodwell, 1991). As can be seen in Fig. 2 very few hedges in Holderness lacked Acer campestre and about 75% of them contained Fraxinus excelsior. So far as is known to the author, there is only one area of woodland in Holderness which might be old and that is Burton Bushes on the western side of Beverley Westwoods. Harris (1971) was unable to prove from documentary evidence that this 6 ha. of woodland was ancient in the sense that it had been in existence since 1600 A.D. but it does contain several herbaceous species considered by Peterken (1974) to be indicative of ancient woodland, including Dog’s Mercury ( Mercurialis perennis ). According to an estimate made by Boatman (1971) 10% of the trees are F. excelsior and 8% are A. campestre. The most common tree is Pedunculate Oak ( Quercus robur ) with 56% frequency. On the evidence of this woodland fragment and the species- rich hedges, it can be postulated that the ancient woodlands of Holderness consisted of one or more facies of NVC type W8 Fraxinus excelsior-Acer campestre-Mercurialis perennis woodland. The most common shrub in Burton Bushes is Holly ( Ilex aquifolium), a species which, according to Rackham (1976), is relatively resistant to grazing. Until recently, Burton Bushes was open to the surrounding pastures and therefore grazed. In Fig. 5 it can be seen that the distribution of hedges containing Holly is similar to that of those with a high frequency of Hazel so this suggests that some of the old woodlands of Holderness were wood pasture. The hedges in Sub-group IB1 are characterized by a relatively high frequency of Dogwood and are mostly in the neighbourhood of Leconfield (Fig. 4). In the past, two large deer parks existed in this area, one just south of Leconfield, on an estate belonging to the Percys and the other just north of Leconfield owned by the Hotham family. Some of the hedges lie precisely on the boundaries of these parks as shown by Neave (1991) and the presence of Dogwood at relatively high frequency would conform with a landscape which, at the time the hedges were created, was more open than that of the woodlands. The Group II hedges on the low-lying land near the River Derwent and the Pocklington Canal in the Plain of York are very different from those in Holderness. Field Maple is absent and Alder and Common Sallow commonly occur. Both of these species are common in North Cliffe Wood which, although it is rather remote from the areas mentioned above, is also low-lying. The Alder and Sallow stands of this woodland have not been surveyed using NVC procedures but it is likely that they would conform to one of the two types of 124 Composition and Distribution of Mixed Hedges in East Yorkshire Sallow woodland and one or more of the three types of Alder woodland described in the National Vegetation Classification. On the evidence of the hedges it seems likely that similar woodlands formerly existed in the neighbourhood of the Derwent and the Pocklington Canal. Unlike the groups considered above, the hedges comprising groups IA and IC were scattered throughout East Yorkshire. Group IA is characterised by a high frequency of Elder, a species which very readily invades existing hedges (Muir & Muir, 1987). It is also a species which cannot be tolerated where a hedge is required to be stock-proof. It is possible, therefore, that the hedges in this group are ones that have suffered long periods of neglect and in this context it is worth noting that, at the time of the survey, 65% were more than 2 m high compared with 48% for the total surveyed for physical features. Very few mixed hedges could be found on the Wolds. It has been claimed that prehistoric man in East Yorkshire lived mainly on the Wolds and cleared the area for cultivation but, according to Allison (1976), more recent archaeological work has suggested that the situation was less clear-cut. Nevertheless, according to Darby and Maxwell (1962) there is no evidence that any woodland existed on the Wolds at the time of the Domesday Survey and it is known that, from the 15th century on, most of the area was open sheepwalk. It is to be expected, therefore, that the Wolds would have been virtually hedgeless until the period of Parliamentary enclosure when much of the land was enclosed for cultivation. One parish boundary hedge on the Sledmere estate, however, is of particular interest as it consists mostly of Spindle ( Euonymous europaeus ) and Hawthorn with some Field Maple, Purging Buckthorn ( Rhamnus cathartica) and Barberry ( Berberis vulgaris). In the earlier survey of East Yorkshire hedges Spindle was found only in two Wolds hedges (Boatman, 1980) but since then it has been found in two Holderness hedges on the boundary of the former medieval park at Scorborough near Leconfield. Finally, consideration should be given to the long-term future of the hedges surveyed in East Yorkshire. It has been stated that a considerable number have been allowed to grow tall, a practice which, according to Garbutt and Sparks’ survey of Judith’s hedge, is likely to have led to a loss of species. Furthermore, eight hedges were less than 1 m in height and a further 57 less than 2 m in height (Table 3). Many of these hedges, especially in the Plain of York, were at least partially smothered by Cleavers, probably a consequence of the spraying of the hedge bottoms, or by tall herbaceous vegetation, especially on the roadside where nowadays only one flail-width of verge nearest the road is regularly cut. Unless action is taken to control the herbs it is likely that the regularly-smothered shrubs will eventually die. Apart from this, coppicing and laying of hedges is almost non-existent in East Yorkshire now and in the absence of these regenerative practices and under the close-cutting management that is so often practiced, there can be no certainty that all of the species that make up a mixed hedge will survive. The implication is, therefore, that a steady decline in the already severely depleted biodiversity of the East Yorkshire countryside can be expected. Acknowledgements The author wishes to thank Dr Ray Goulder for conducting the TWINSPAN analysis on the hedgerow data and Dr Jan Crowther for help and advice concerning the location of documents dealing with former agricultural practices in East Yorkshire. Note A copy of the database has been supplied to the local authority and it is intended that another will be lodged at the Beverley Reference Library. References Allison, K. J. (1976) The East Riding of Yorkshire Landscape. Hodder and Stoughton, London. Boatman, D. J. (1971) Burton Bushes: an ecological view. Bulletin of the Hull Natural Book Reviews 125 History Society 3.2: 8-14. Boatman, D. J. (1975) Hedges and roadside scrub of Walkington parish. Naturalist 100: 91-95. Boatman, D. J. (1980) Mixed hedges of the former East Riding of Yorkshire. Naturalist 105: 41-44. Darby, H. C. and Maxwell. I. S. (1962) Domesday Geography of Northern England. Cambridge University Press, Cambridge. Downing, S. L. and Sparks, T. H. (1999) The Monks Wood mixed hedge experiment. Aspects of Applied Biology 54: 223-228. Garbutt, R. A. and Sparks, T. H. (1999) Changes in composition of a species-rich ancient hedgerow. Aspects of Applied Biology 54: 229-234. Harris, A. (1971) Burton Bushes: the historical evidence. Bulletin of the Hull Natural History Society 3.2: 3-8. Hoskins, W. G. (1955) The Making of the English Landscape. Hodder and Stoughton, London. Latham, I. (1974) A General Review of the Agriculture of the East Riding of Yorkshire. Board of Agriculture, London. Muir, R. and Muir, N. (1987) Hedgerows: Their History and Wildlife. Michael Joseph, London. Neave, S. (1991) Medieval Parks of East Yorkshire. Hutton Press, Beverley. Peterken, G. L. (1974) A method for assessing woodland flora for conservation using indicator species. Biological Conservation 6: 239-245. Pollard, E., Hooper, M. D. and Moore, N. W. (1974) Hedges. Collins, Glasgow. Rackham O. (1976) Trees and Woodland in the British Landscape. J. Dent, London. Rackham O. (1986) The History of the Countryside. Dent, London. Rodwell, J. S. (1991) British Plant Communities Volume 1: Woodlands and Scrub. Cambridge University Press, Cambridge. BOOK REVIEWS Last-instar Larvae and Pupae of the Simuliidae of Britain and Ireland by Jon Bass. 104 pp., 24 figs. Lreshwater Biological Association, Ambleside LA22 OLP 1998. £14.00, inch postage. The simuliid flies, otherwise known as blackflies or buffalo-gnats, are very important insects from a medical standpoint in many parts of the world. In Britain the adults sometimes occur in plague proportions, when they can become a major nuisance owing to their persistence and bloodsucking habits. The larvae and pupae are often extremely abundant in less polluted streams and rivers, and will be familiar to all students of freshwater biology. Many treatises exist on all aspects of simuliid morphology, taxonomy and biology and research in these continues to progress rapidly, but for the identification of final-instar larvae and pupae of British species, this will now be regarded as the standard work. The book under review consists mainly of a copiously illustrated key to the final- instar larvae and pupae of the 32 British species, together with brief ecological notes and a synopsis of the known distribution of each. The nomenclature of the species concerned has been updated to accord with current knowledge and usage, enabling today’s students of the family to use existing literature on the identification of adults of British species. The book ends with a valuable list of references. PS 126 Book Reviews A Key to the Freshwater Triclads of Britain and Ireland with notes on their ecology by T. B. Reynoldson and J. O. Young. Pp. 72, including numerous line drawings and 1 colour plate. Freshwater Biological Association, Ambleside, LA22 OLP. 2000. £14.00 inch postage. This is a welcome addition to the annals of freshwater biology, coming at a time when emphasis is being put on the necessity of monitoring the state of freshwater as a whole. The text is clear and unambiguous and the identification of species made even more useful by the addition of a key to preserved material, something omitted from earlier editions. There is a well designed chapter on the general biology and distribution of the species as well as an extended bibliographical list. Rarely does one get such complete coverage between a single pair of covers. The price may prove a deterrent to students and those with only a superficial interest in triclads, which is unfortunate. For the serious worker it is still a good investment, being more than twice the size of earlier editions and including far more information than is apparent at first glance. DTR Caring for Planet Earth: stories and prayers for children by Ian M. Fraser and illustrated by Miles Forde. Pp. vi + 74. Saint Andrew Press, Edinburgh. 2000. £5.99 paperback. Intended for use in Sunday schools, assemblies and with youth groups, this is a collection of 25 stories, each with a linked prayer. Every story ends with a question about caring for planet earth, but these links with the environment are often contrived. The stories themselves are quaintly old-fashioned. A tale about an engine called Huffity-puff, which has difficulty climbing a mountain, ends by asking how in caring for the planet we can discover “the steepest hills to climb”. A frog arriving late for the toys’ party because he got stuck in a snowdrift prompts the author to pose the question “What can we do to care for planet earth and to free people who are in some way trapped?” Twins called Alice and Andrew have earnest discussions about whether they are making the best use of the small patch of garden given over to them. Out of touch with the reading tastes of today’s children, the author writes about humanised vegetables, a wheelie bin called Willie and a bus named Bessie, even a bamboo tree which has thoughts and feelings - all the type of characters that are anathema to modern editors. A few of the stories for older children fare slightly better: an eighteen year old girl who is made aware of the existence of sweat shops in the Far East through her Sri Lankan pen friend vows to buy only clothes produced under proper working conditions and Kofi, a boy blind and deaf from birth, has serious questions to ask about his new life when his sight is restored. There must, though, be less tenuous ways of making young people aware of environmental issues. PB Lichens by William Purvis. Pp. 112, inch numerous colour illustrations. Life Series, The Natural History Museum, London. 2000. £9.95 paperback. This superbly produced work does justice not only to the beauty of these fascinating plants but also to their uniqueness. Stunning photography is supported by an informative text that effectively portrays their biodiversity, ecological role, ability to survive in extreme environments and value as biological monitors; other sections deal with their structure, growth, dispersal, classification and economic uses, and some practical projects are appended. Thoroughly recommended as an introductory text for schools and natural historians in general. MRDS 127 RECOVERY BY AQUATIC VEGETATION IN A WOLDS HEADSTREAM SUBSEQUENT TO THE 1988-1992 DROUGHT* R. GOULDER Department of Biological Sciences, University of Hull, Hull HU6 7RX Abstract During the 1988-1992 drought the flow of a winterbourne in the Yorkshire Wolds became irregular, culminating in dryness from summer 1991 through summer 1992. Aquatic plants were reduced in species richness and abundance but had partially recovered by summer 1993. Their pre-drought luxuriance was re-established through 1994-1999. Introduction Global warming and potential change in UK weather patterns might lead to greater frequency of hot dry summers (Marsh, 1997; Mayes, 1998) with accompanying drought conditions. The study of vegetation during and following past drought potentially throws light on the response of vegetation to future drought events. The 1988-1992 drought in Southern and Eastern England (Marsh et al., 1994) diminished the recharge of aquifers. This, often coupled with abstraction by pumping from wells, led to low flows and wider occurrence of dry stream beds in chalk river systems. These conditions are associated with loss and impoverishment of aquatic flora (Giles et al., 1991; Hill & Langford, 1992). In the Yorkshire Wolds the drought was evidenced by rainfall between August 1988- February 1992 of 70-75% of long-term average, by aquifer recharge over the four winters 1988/89-1991/92 of 30-70% of the pre-1988 average, and by river run-off between September 1990-August 1992 of 50-60% of long-term average (Marsh et al, 1994). Mill Beck, a Wolds stream near to Market Weighton, was much influenced by the drought. The spring-fed beck, part of the River Foulness drainage system, rises from beneath the Upper Cretaceous Chalk at National Grid Reference SE 899 426. Through the 1980s Mill Beck was a winterbourne, with maximum discharge during January-March and with flow continuing till about September. During spring and summer unshaded reaches, flowing through damp pasture, supported luxuriant emergent aquatic vegetation, largely dominated by Apium nodiflorum (Fool’s Water-cress), Rorippa nasturtium- aquaticum (Water-cress), Veronica anagallis-aquatica (Water-speedwell) and Glyceria fluitans (Floating Sweet- grass) (Goulder, 1984; Rimes & Goulder, 1986a, 1986b, 1987). During 1989-1992, however, the flow of Mill Beck became irregular, culminating in a dry stream bed from summer 1991 to beyond August 1992 (Goulder, 1992). Water levels in the chalk aquifer are exemplified by data from Southwood Farm (SE 922 441), 2.8 km north-east of the Mill Beck source. As was typical for the 1980s, lowest water levels during 1985-1987 were recorded in the final quarter of each year (Fig. 1), a time when Mill Beck was largely dry. Water levels then increased substantially to highest levels in the subsequent first quarter. This coincided with maximum stream flow. During the drought, however, the early-year increase was much less in 1989 and 1990, and failed completely in 1992. Hence the dry stream bed from summer 1991 to the end of 1992, a state which may have been exaggerated by continuous pumping through the drought years from Springwells borehole, about 100 m south of the Mill Beck source. The monthly average abstraction rate from January 1988 to December 1992 was 2.16 x 103 m3 d 1 (range 0.97-2.87 x 103 m3 d ') compared to the similar 2.26 (0.36-2.82) x 103 m3 d~' between January 1984 and December 1987. Surviving aquatic vegetation in the dry bed of Mill Beck during summer 1992 was described by Goulder (1992). The bed of the stream from 0-50 m downstream of the source * Developed from a poster originally presented at the YNU’s Conference on “Recording and Monitoring Yorkshire’s Natural Environment”, Harrogate, 26 February 2000. Naturalist 126 (2001) 128 Vegetation Subsequent to Drought Year (1985-1999) FIGURE 1 Mean quarterly water level in the aquifer at Southwood Farm, 1985-1999 (m above ordnance datum), prepared from records supplied by the Environment Agency. had become a grassy sward and only six of the 13 species of aquatic macrophyte recorded in 1983 survived (Table 1). Of these, the grass Agrostis stolonifera (Creeping Bent) had much increased in abundance, otherwise the survivors were stands of perennial emergent species, together not occupying more than about 15% of the dry stream bed, most notably Glyceria maxima (Reed Sweet-grass), Iris pseudacorus (Yellow Iris) and Mentha aquatica (Water Mint). Seedlings of hawthorn ( Crataegus monogyna) and dog rose ( Rosa canina) were establishing themselves while the formerly dominant hydrophytes A. nodiflorum, R. nasturtium- aquaticum and V. anagallis-aquatica were not recorded. Similarly, the stream bed from 225-360 m downstream of the source was a grassy sward, with abundant thistles ( Cirsium arvense ) and stinging nettles ( Urtica dioica ). Seven species of aquatic macrophyte were recorded compared to 12 in 1984 (Table 2). The formerly dominant A. nodiflorum and R. nasturtium- aquaticum, along with Berula erecta (Lesser Water-parsnip) and Veronica beccabunga (Brooklime), were reduced to a few plants in a damper, more shaded, area towards the upstream end of the reach. Following the end of drought, Mill Beck flowed again during 1993. In that summer, some of the missing hydrophytes reappeared (Goulder, 1993). Nine species were recorded in the 0-50 m length compared to six in 1992 (Table 1); while 12 species were recorded in the 225-360 m reach compared to seven in 1992 (Table 2). However, the vegetation had not fully recovered. Approximately 80% of the steam bed was occupied by inundated non-flowering grasses, thoroughly entangled by filamentous algae. Many terrestrial plants remained conspicuous, especially thistles and stinging nettles in the 225-360 m reach. The aim of the present contribution is to describe the summer aquatic vegetation in Mill Beck over the longer-term, post-drought, period up to 1999. Materials and Methods A yearly visit was made to Mill Beck, during June-July 1994 to 1999, and aquatic macrophytes were recorded in the lengths of stream 0-50 m and 225-360 m downstream of the source. The checklist used was of aquatic plants in England and Wales (Palmer & Newbold, 1983); nomenclature follows Stace (1997). Presence or absence was recorded Vegetation Subsequent to Drought 129 TABLE 1 Aquatic macrophytes recorded 0-50 m downstream of the Mill Beck source. 1983 1992 1993 1994 1995 1996 1997 1998 1999 Agrostis stolonifera + + + + _ 2 3 1 1 Apium nodiflorum + - + + + 3 3 3 3 Berula erecta + Caltha palustris + + - + + 1 1 1 2 Eleocharis palustris - - - + - - 1 - - Glyceria fluitans + - - + + - 1 1 1 Glyceria maxima + + + + + 3 3 3 3 Iris pseudacorus + + + + + 3 3 3 3 Mentha aquatica + + + + + 2 3 2 2 Myosotis scorpioides + + + + + 2 3 2 2 Ranunculus flammula + Rorippa nasturtium-aquaticum + - + + + 3 3 3 3 Veronica anagallis-aquatica + - + + + 2 3 1 1 Veronica beccabunga + - + + - 2 - 1 - Number of species 13 6 9 12 9 10 11 11 10 (+) = Present, (-) = not recorded, 1 = 1994 data are from Goulder (1993). <0.1% cover, 2 = 0.1 -5% cover, 3 = >5% cover. Pre- TABLE 2 Aquatic macrophytes recorded 225-360 m downstream of the Mill Beck source. 1984 1992 1993 1994 1995 1996 1997 1998 1999 Agrostis stolonifera + + + + + _ 1 _ 1 Apium nodiflorum + + + + + 3 3 3 3 Berula erecta + + + + - 2 2 2 1 Callitriche sp. - - + + - - - - - Caltha palustris + + + + + 1 1 1 1 Eleocharis palustris - + + + 1 3 1 1 Equisetumfluviatile ! Glyceria fluitans + - - + + 3 3 3 3 Mentha aquatica - - + + + 2 - 2 1 Myosotis scorpioides + + + + + 3 3 2 2 Ranunculus aquatilis + - + - - - - - '4 - - Ranunculus flammula + Rorippa nasturtium-aquaticum + + + + + 3 3 3 3 Veronica anagallis-aquatica + - + + - 3 3 1 2 Veronica beccabunga + + + + + 2 2 3 3 Number of species 12 7 12 12 9 10 10 10 11 (+) = Present, (-) = not recorded, 1 = <0.1% cover, 2 = 0.1-5% cover, 3 = >5% cover. Pre- 1994 data are from Goulder (1993). 130 Vegetation Subsequent to Drought during 1994-1995. From 1996-1999 an abundance score was used (Holmes, 1983); i.e. 1 = <0.1% of stream bed covered, 2 = 0.1-5% cover, and 3 = >5% cover. Results and Discussion Water flowed from the Mill Beck source during June-July in every year from 1994-1999. Substantial flow was observed in 1994, 1995, 1998 and 1999 whereas the flow was low in 1996 and modest in 1997. These lesser flows followed the 1995 drought and continuing rainfall deficiency in 1996 (Marsh, 1996). During the low flow years, especially 1996, first-quarter water levels were low in the aquifer at Southwood Farm (Fig. f). That flow continued was perhaps due to cessation of pumping at Springwells borehole from April 1995 onwards. The stream from 0-50 m downstream of the source was occupied by an 80-100% cover of luxuriant aquatic vegetation in each of the years 1994-1999, with the exception of 1996. The vegetation was dominated by Apium nodiflorum and, usually profusely flowering Rorippa nasturtium- aquaticum, emergent to 50 cm or more. The adjacent pasture, and the stream vegetation, were not grazed, unlike in the early 1980s when there was grazing by cattle. In the low flow year of 1996, the macrophyte vegetation, largely R. nasturtium- aquaticum, was restricted to about 50% cover, while the shallow water of the gaps between, mostly <10 cm deep, was extensively colonized by filamentous algae attached to the stream bed. The stream from 225-360 m downstream of the source supported a dense, nearly complete, cover of emergent aquatic vegetation in every summer from 1994-1999. Again A. nodiflorum and floriferous R. nasturtium-aquaticum were most conspicuous, encouraged by absence of grazing from 1996 onwards. By June 1994, the second summer with post-drought water flow, the number of aquatic plant species, in both reaches of stream, was similar to that in the pre-drought early 1980s. That is, for 0-50 m, 12 species compared to 13 in 1983 (Table 1); and for 225-360 m, 12 species in both 1994 and 1984 (Table 2). In following years the number of species, the species composition, and their relative abundance (from 1996) remained fairly constant (Tables 1, 2). Some of the, minor, variation may reflect plants being overlooked during once-a-year visits. Probably genuine losses, however, are Ranunculus flammula (Lesser spearwort), scarce in both lengths in 1983-1984 and not recorded since, and Ranunculus aquatilis (Common Water-crowfoot) a few plants of which were found in the 225-360 m reach in 1984 and 1993 but not since. Goulder (1992) suggested that the Mill Beck vegetation represented an aquatic vegetation type, characteristic of intermittent calcareous streams, not widespread in the Yorkshire Wolds, and potentially threatened by drought and abstraction. It is pleasing that the present study showed rapid recovery after drought. Recovery of chalk streams from even longer periods of dryness has been reported. In the River Ver, Hertfordshire, Ranunculus peltatus (Pond Water-crowfoot) was re-established within three years of cessation of groundwater abstraction and resumption of flow after many years of dryness (Mainstone, 1999). Drought tolerance by headstream plant communities is relevant to the ongoing debate over UK climate change (Marsh, 1997; Mayes, 1998). If droughts are to become more frequent it is good news that these communities are sufficiently robust to survive. Acknowledgements I am grateful to the Environment Agency for records of aquifer water levels and to Yorkshire Water for information on abstraction. References Giles, N., Phillips, V. E. and Barnard, S. (1991) The Current Crisis: Ecological Effects of Low Flows on Chalk Streams. Royal Society for Nature Conservation/Wildlife Trusts Partnership, Lincoln. Book Review 131 Goulder, R. (1984) Downstream increase in the abundance and heterotrophic activity of suspended bacteria in an intermittent calcareous headstream. Freshwater Biology 14: 611-619. Goulder, R. (1992) Loss of aquatic vegetation in a Wolds winterbourne during perennial dryness. Naturalist 117: 127-131. Goulder, R. (1993) Aquatic vegetation and resumption of flow after drought in a Wolds winterbourne. Naturalist 118: 85-87. Hill, C. and Langford, T. (1992) Dying of Thirst: A Response to the Problem of Our Vanishing Wetlands. Royal Society for Nature Conservation/Wildlife Trusts Partnership, Lincoln. Holmes, N. (1983) Focus on Nature Conservation No. 4: Typing British Rivers According to Their Flora. Nature Conservancy Council, Shrewsbury. Mainstone, C. P. (1999) Chalk Rivers: Nature Conservation and Management. English Nature/Environment Agency, Peterborough. Marsh, T. J. (1996) The 1995 drought - a water resources review in the context of the recent hydrological instability. In T. J. Marsh (ed). Hydrological Data United Kingdom: 1995 Yearbook, pp. 25-33. Institute of Hydrology, Wallingford. Marsh, T. J. (1997) The UK drought and climate change. Weather 52: 354-355. Marsh, T. J., Monkhouse, R. A., Arnell, N. W., Lees, M. L. and Reynard, N. S. (1994) Hydrological Data UK: The 1988-92 Drought. Institute of Hydrology, Wallingford. Mayes, J. (1998) United Kingdom summer weather over 50 years - continuity or change? Weather 53: 2-11. Palmer, M. and Newbold, C. (1983) Focus on Nature Conservation No. 1: Wetland and Riparian Plants in Great Britain. Nature Conservancy Council, Shrewsbury. Rimes, C. A. and Goulder, R. (1986a) Quantitative observations on the ability of epiphytic bacteria to contribute to the populations of suspended bacteria in two dissimilar headstreams. Freshwater Biology 16: 301-311. Rimes, C. A. and Goulder, R. (1986b) Temporal variation in density of epiphytic bacteria on submerged vegetation in a calcareous stream. Letters in Applied Microbiology 3: 17- 21. Rimes, C. A. and Goulder, R. (1987) Relations between suspended bacteria, epiphytic bacteria and submerged vegetation over the spring growing season in a calcareous headstream. Freshwater Biology 17: 291-305. Stace, C. (1997) New Flora of the British Isles. 2nd ed. Cambridge University Press, Cambridge. BOOK REVIEW Provisional Atlas of British Hoverflies (Diptera, Syrphidae) by Stuart G. Ball and Roger K. A. Morris. 166 pp. with b/w maps and tables. Biological Records Centre, CEH, Monks Wood, Abbots Ripton, Huntingdon. Joint Nature Conservation Committee. 2000. [price not stated on cover], Britain has long been in the forefront of global dipterological research and this extremely comprehensive book on British hoverflies admirably perpetuates this tradition. This indeed provides a notable landmark, for precisely a century has passed since G. H. Verrall published his magnum opus on British Hoverflies (. British Flies , Vol. VIII, published by Gurney & Jackson, London), when the handful of contemporary British dipterists could have been forgiven for believing that the ultimate treatise on these fascinating insects had been produced. Indeed, it is a testimony to the genius of Verrall that his work remains of 132 Book Review value to British students of the family today. However things have moved on apace since Verrall, especially since the publication in 1951 of C. N. Colyer and C. O. Hammond’s Flies of the British Isles (Wayside & Woodland series, Warne, London), which precipitated an explosion of interest in flies that has continued to mushroom worldwide ever since. Indeed, during the past 50 years dipterology has been one of the major growth areas in entomological research on a global scale. Owing to their intrinsic beauty and obtrusiveness, hoverflies have long been one of the most popular families of flies, so vast strides in our knowledge and understanding of these insects have also been made during this same period. Inevitably, with the vast growth in the number of researchers, many hoverflies have been added to the British list since 1950, so that whilst Verrall knew of 194 British species, the number has now risen to 266. The majority of these additions have resulted from “splitting” as species concepts have changed, based mainly on detailed examination of male genitalic characters and fuelled by the kudos derived from describing new species. The “downside” of this activity of course is that in some of the more critical hoverfly genera, such as Sphaerophoria , the females cannot be identified with confidence. Research has progressed on numerous aspects such as ecology, larval and adult morphology and biology, and, thanks to the many recording schemes undertaken through the Biological Records Centre at Monkswood, in phenology and national distribution. During the past 30 years many distributional atlases have been produced through BRC and, amongst these, Ball and Morris’s work on British hoverflies stands as a model for emulation. The coverage is phenomenal, based as it is on an analysis of over 30,000 individual records from all parts of Britain. Although many hoverflies are regarded today as endangered or vulnerable, only one species ( Myolepfa potens) is highlighted as having become extinct in Britain since Verrall’s day, whilst several extremely spectacular ones have evidently colonised Britain during the same period (e.g. Callicera spinolae, Eriozona syrphoides, Volucella zonaria etc.). Apart from the distribution maps, graphs showing the flight periods, a brief description of the biology and national status is given for each species. A map is provided to show the 10 kilometre squares from which hoverfly records have been received by BRC, and another indicates the intensity of recording per square. This will direct recorders to the poorly recorded areas of Britain and hopefully lead to more uniform coverage. Expectedly, the best-recorded squares are in and around the major conurbations and traditional collecting areas, with the main under-recorded areas being in various parts of Scotland, although surprising gaps even exist in southern England. Collectors in Lancashire and Yorkshire will perhaps be alerted to the presence of four squares in these counties from which no hoverflies have been recorded and will react accordingly. For those involved in site conservation issues, an important chapter deals with habitat indicator species, whilst the current national status of each species is also provided. The nomenclature follows the most recent Checklist of British Diptera, edited by R J. Chandler and published in 2000 by the Royal Entomological Society of London. This book represents a major milestone in British hoverfly studies and will occupy a prominent place on the shelves of all those with at least a passing interest in these flies. As stated, the book will also prove invaluable in site conservation programmes. There is a comprehensive list of references to all of the literature of relevance to the British hoverfly fauna. Finally the book provides an invaluable basis for the assessment of any future faunal changes, thus fuelling the hotly contested debate on “global warming”. PS 133 MAPPING DRYOPTERIS SUBMONTANA (RIGID BUCKLER FERN) TO THE METRE: EVIDENCE AND IMPLICATIONS FOR FRACTAL DISTRIBUTIONS* STEPHEN HARTLEY and WILLIAM E. KUNIN School of Biology, University of Leeds, Leeds LS2 9JT Introduction Distributional dot maps are one of the most ubiquitous and useful outputs to emerge from a wide range of recording schemes. As well as suggesting interesting relationships between a species and its habitat requirements, comparisons between maps have been used to assess the relative rarity or commoness of different species. For example, British plants are considered nationally “rare” if they are recorded from fifteen or less 10 km squares (hectads), and “scarce” if known to occur in sixteen to one-hundred 10 km squares. However, the number of occupied squares is not the only property of a distribution map, the relative positions of the occupied squares can also be used to make useful inferences regarding the local frequency of a species. For example, experience tells us that a grid square in the centre of a cluster of occupied squares is likely to harbour many more populations or individuals than a relatively isolated occupied square. Similarly, those species with a restricted but relatively compact distribution of occupied squares are likely to be much more abundant “on the ground” than those species with a widespread but scattered distribution (see Figure 1A and IB). One way of formalising this intuition is through the use of fractal geometry. Fractal objects are characterised by the property of self-similarity: when a small portion of the whole is magnified, it retains certain features which are similar to the whole. Many examples of approximately fractal objects can be found in the natural world, such as cauliflower florets, the branching patterns of trees and fungal mycelia, mountain peaks or the edges of snowflakes (Mandlebrot, 1982; Donnelly et al. 1999). The geographical distribution of a species can also be considered fractal if the density of occurrence at a coarse scale (e.g. a national distribution mapped on a 10 km resolution grid) is similar to the density of occurrence in a small, occupied region viewed at a finer resolution (e.g. a county distribution mapped on a 1 km grid). One mathematical test for a fractal distribution is to plot a graph (on logarithmic axes) of the number of grid-squares occupied against grid resolution: a straight line will result if the distribution is genuinely fractal (Hastings & Sugihara, 1993; Gautestad & Mysterud, 1994). Kunin (1998) provides evidence for the approximately self-similar nature of plant distributions, when viewed at 50 km, 10 km and 2 km (tetrad) resolutions. However, it is not yet known whether the patterns present at these national and regional scales repeat themselves still further at truly local scales (e.g. hectare resolution and below). To test this idea we collected information on the distribution patterns of Dryopteris submontana (the rigid buckler fern) and four other plant species. In this paper the results for D. submontana will be presented in detail, whilst the distributional patterns of Dianthus armeria (the Deptford pink) are used for comparative purposes. Methods National Distributional Patterns National distribution maps were obtained from the Biological Records Centre (BRC), CEH, Monks Wood, at one kilometre resolution. ETsing these, the number of occupied 100 km squares, 10 km squares and 1 km squares was calculated for each species (dimensions refer to the length of one side of the square). * Developed from a poster originally presented at the YNET’s Conference on “Recording and Monitoring Yorkshire’s Natural Environment”, Harrogate, 26 February 2000. Naturalist 126 (2001) 134 Mapping Dryopteris submontana to the Metre Dryopteris submontana Dianthus armed a FIGURE 1A. National distributions of Dryopteris submontana (10 hectads occupied) and Dianthus armeria (19 hectads occupied), as recorded since 1987 (BRC data, Stewart et al., 1998. and Wigginton, 1999). Circled areas are illustrated at a finer resolution in Figure IB. 10 km ■ ■ !■ ■ H b* ■ c<* i ■ ■ 10 km > ■ 7 - lx ■ J 7 Dryopteds submontana Dianthus armeda FIGURE IB. Regional-scale distributions of Dryopteris submontana and Dianthus armeria mapped at 1 km resolution. With the exception of one additional site for D. submontana, all data are courtesy of the BRC. Refer to Figure 1 A for further details and location. Circled areas (D. submontana only) are illustrated at a finer resolution in Figure 2. 135 Mapping Dryopteris submontana to the Metre Field Survey of Local Distributional Patterns To estimate finer-scale occupancy levels, four 1 x 1 kilometre squares were selected at random for detailed field survey during the summer of 1999. At each square kilometre chosen for survey, as far was practically possible, every hectare was searched for an average of twenty person-minutes and the presence or absence of the target species noted, along with a rough estimate of the number of individual plants. This phase of the survey (the coarse-scale field survey) generally took a pair of surveyors two days to complete. The occupied hectares were then ranked according to the estimated abundance of the plant. One hectare from the “high abundance” and one hectare from the “low abundance” half were then selected at random for a “medium-scale” survey at 10 m resolution. To achieve this, a grid of markers was laid out at 10 m intervals within each selected hectare, using Global Positioning Satellite (GPS) equipment that was accurate to ±1 m. Each 10 x 10 m square was then thoroughly searched for three to five minutes (depending upon habitat suitability and complexity) by one person, and once again, population estimates were made for each square. Finally, one “high abundance” and one “low abundance” 10 x 10 m square were selected at random from the medium-scale survey, and the presence or absence of the plant was recorded in a 10 x 10 grid of 1 metre squares. For this, the finest-scale survey, tape- measures and string were used to construct a one-metre resolution grid, and the precise location of each plant within the one metre square was recorded to the nearest centimetre. Obviously, where only one square was found to be occupied at the coarse or medium scale survey, only one square could be surveyed at the respective medium or fine-scale survey. Results Table 1 illustrates the national number of grid squares recorded as occupied (since 1987) for the two example species at three relatively coarse-scale resolutions. Table 2 presents the average occupancy at the three finer resolutions, as determined from our field survey of four square kilometres. Specific examples of the distribution of D. submontana, at each of the three fine-scales, are given in Figure 2. Note, that there is generally a positive correlation between the number of grid cells occupied at successive scales. This tendency for coarse-scale occupancy to reflect local abundance has been well documented in many TABLE 1 . Number of grid squares from which a species has been recorded (nationally). Data courtesy of the Biological Records Centre, Centre for Ecology and Flydrology. Available in summary form in Stewart et al. (1998) and Wigginton (1999). National occupancy since 1987 (BRC data) Species 100 km squares 10 km squares 1 km squares D. submontana 2 10 29 D. armeria 12 19 19 TABLE 2. Average percentage occupancy of squares at three different resolutions. In each case, the larger square was divided into a 10 x 10 grid of smaller squares. Numbers in brackets refer to the number of square kilometres, hectares and 10 m squares surveyed in the field. Average percentage occupancy (from Field Survey) Hectares per 10 m squares 1 m squares per Species 1 km square per hectare 10 m square D. submontana 3.25 (4) 5.83 (6) 4.80(10) D. armeria 1.25 (4) 2.00 (5) 4.00 (7) 136 Mapping Dryopteris submontana to the Metre c) Malham d) High Sleets (1 IL 1 f= FIGURE 2. A sample of the field-survey results for the distribution of D. submontana, as recorded at three different resolutions. Each 10 x 10 grid is a magnification of the area circled in the map directly above it. The square kilometres in the top row correspond to the four sites circled in Figure IB, labelled a-d. species (see Gaston, 1996 for a review), and it is one of the characteristics expected from a fractal-like distribution. Assuming the field survey results from Table 2 are representative for the whole country, we estimate that there are 94 hectares, 549 ten-metre squares or 2363 one-metre squares containing D. submonata in Britain. A log-log linear regression of the BRC data (at 100 km, 10 krn and 1 km resolution), when extrapolated to finer resolutions, predicts 129 hectares, 516 ten-metre squares and 2066 one-metre squares. This level of agreement between the field survey estimates, and the linear (or fractal) extrapolation from national-scale maps, is illustrated in Figure 3. The steep slope for D. submontana (in Figure 3) reflects the fact that each occupied coarse-scale square contains on average about four occupied medium-scale squares, and each occupied medium-scale square contains on average about four occupied fine-scale squares and so-on up the line for every successive pair of scales. By way of contrast, Dianthus armeria is very sparsely distributed at all scales apart from the 1 m resolution grid; each square occupied at one scale contains on average only one or two occupied squares when viewed at ten times finer resolution. The change in slope at fine resolutions is Mapping Dryopteris submontana to the Metre 137 interesting in its own right, but it does signify that the national scale pattern cannot be extrapolated indefinitely. SZ Resolution of map grid FIGURE 3. Logarithmic plot of occupancy against resolution for D. submontana (filled circles) and D. armeria (open squares). Points for 100 km to 1 km occupancy based upon BRC data and points for hectare to 1 m occupancy based upon an extrapolation of field surveys. Solid lines represent a linear regression using the national BRC data, and dashed lines represent the fine-scale predictions made by extrapolating the regression line (i.e. assuming a fractal distribution). Implications British definitions of the rarity and scarcity of vascular plants are currently based upon the number of 10 x 10 kilometre National Grid squares that a species occupies (e.g. Stewart et al., 1998; Wigginton, 1999). Figure 3 demonstrates how our perception of a species relative abundance (as measured by the number of grid squares occupied) can be dramatically altered by varying the resolution of the map (see also Pearman, 1997). From a conservation point of view, it may often be desirable to know the national distribution at a much finer resolution. For example, how many hectares or how many square metres are occupied? Such a mapping task would clearly be impractical for more than a handful of species. However, if most species do show fractal tendencies in their distributions, then coarse-scale knowledge can be extrapolated to make reasonable estimates of fine-scale coverage. We have shown for Dryopteris submontana (and to a lesser extent Dianthus armeria ), that predictions of typical local coverage, made from national distribution maps, broadly 138 Mapping Dryopteris submontana to the Metre agree with our field survey estimates. The key question is, are these species typical or just fortuitous examples? Also, what are the limits of extrapolation? Perhaps the local-scale slope (between 10 m and 1 m in Figure 3) is determined by completely different processes than those that generate national distribution patterns (i.e. the slope between 100 km and 1 km)? This seems to be the case for D. armeria. If so, can we relate the slopes of different sections of the graph to the biology of the particular plant species? For example, “Does the plant reproduce vegetatively?”, “Does it make a difference whether the seeds are wind- dispersed or animal-dispersed?”. By carrying out further work, on more pairs of species, we hope to move closer to answering these and other questions that could make the interpretation of distributional dot maps even more interesting and informative. Acknowledgements Field work was conducted by Hazel Aitken, Kara Clarkson, Duncan Gray, Stephen Hartley and Sarah Mellor, with assistance from Maxine Hartley, Ben Hodgson, Hannah Hodgson, Bill Kunin and Sarah Utley. We would like to thank Phyl Abbott and Phil Wilson for sharing their knowledge regarding the distributions of D. submontana and D. armeria respectively. Also, we are indebted to the many farmers, land owners and reserve wardens who granted us access permission and provided more detailed local information. We are grateful to Johnathon Willot for commenting on an earlier draft of this article. Finally, we thank all the BSBI recorders and the Biological Records Centre, CEH, Monks Wood, without whom the national distribution maps would not be possible. This project is supported by the Natural Environment Research Council, (grant GR3/11916). References Donnelly, D. P, Boddy L. and Wilkins, M. F. (1999) Image analysis - a valuable tool for recording and analysing development of mycelial systems. Mycologist 13: 120-125. Gaston, K. J. (1996) The multiple forms of the interspecific abundance-distribution relationship. Oikos 76: 211-220. Gautestad, A. O. and Mysterud, I. (1994) Fractal analysis of population ranges - methodological problems and challenges. Oikos 69: 154-157. Hastings, H. M. and Sugihara G. (1993) Fractals: A User’s Guide for the Natural Sciences. Oxford University Press, Oxford. Kunin, W. E. (1998) Extrapolating species abundance across spatial scales. Science 281: 1513-1515. Mandelbrot, B. B. (1982) The Fractal Geometry of Nature. Freeman, San Francisco. Pearman, D. (1997) Presidential Address, 1996: Towards a new definition of rare and scarce plants. Watsonia 21: 231-251. Stewart, A., Pearman, D. A. and Preston, C. D., eds. (1994) Scarce Plants in Britain. JNCC, Peterborough. Wigginton, M. J., ed. (1999) British Red Data Books I Vascular Plants. JNCC, Peterborough. 139 NOTES ON THE DISTRIBUTION OF LARGE-LEAVED LIME (TILIA PLATYPHYLLOS SCOP.) ON THE MAGNESIAN LIMESTONE IN SOUTH YORKSHIRE* J. A. NEWBOULD Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS Introduction Large-leaved Lime, a nationally scarce tree (Newlands, 1999), is recorded in Perring and Walters (1962) for the 10 km square SK 58 and in Stewart et al. (1994) as a pre-1970 record. There are no other records from the Magnesian Limestone in South Yorkshire. The status of Large-leaved Lime in Britain has been the subject of studies by a small number of authors; for example, Rackham (1980) considered Lime in eastern England and Pigott (1969) reviewed the status of Lime in the Derbyshire Dales and reported in the same paper that Large-leaved Lime was present in Anston Stones Wood in Rotherham and at Pleasley, on the Magnesian Limestone on the Nottinghamshire and Derbyshire border. Peterken (1981) described Maple-Ash-Lime woodland in Lincolnshire, and Oak-Ash-Lime woodland at Edlington Wood, Doncaster (SK59). Peterken (1980) also visited Roche Abbey and produced an internal report for the Nature Conservancy Council. Langridge (Meade & Langridge, 1987) surveyed a number of woodlands in South Yorkshire for the Nature Conservancy Council. This report did not quantify the presence of Large-leaved Lime, although it reported its presence in seven woods in the Rotherham area alone and ten in total. Pigott (1981) reviewed the status of Large-leaved Lime in Britain without mentioning the South Yorkshire populations. Lees (1888) reported that Tilia grandiflora (= T. platyphyllos ) is an alien, always found in parks, stating “the largest I can recollect is in a park near Firbeck”. This study was undertaken in order to estimate the population and map the location of Large-leaved Lime in woods on the Magnesian Limestone in South Yorkshire, and, where possible, in adjacent counties, to ensure that an accurate representation of the species is provided for the BSBI’s ‘Atlas 2000’ project, and to develop a suitable methodology for the regional study of rare and scarce plants proposed by Wilmore (1997). The study also looked for young trees in order to determine whether the population might be on the increase. Methods The present author reviewed the standard floras, e.g. Clapham et al. (1987) and Stace (1991), before opting to use the instructions and description of Tilia species in Rich and Jermy (1998) to determine the species in the field. Determination in the field was made using only shoots from an exposed portion of the tree’s crown, which had flowers or fruits. Shoots from a shaded portion of a tree or the lower shoots growing from the tree’s base were rejected on the grounds "that identification is difficult or impossible” (Rich & Jermy, 1998). This policy may have left good [Large-leaved Lime] trees uncounted, as many of the Limes are located on the edge of cliffs or crags and the terrain is dangerous for a lone field worker. Approximately four Limes were not determined in Anston Stones Wood (Compartment 13) due to the terrain. In particular, trees should have cordate leaves with triangular teeth having an acute, well-defined apex. The upper surface should be wrinkled, with sparse simple hairs, especially along the main veins, whilst the lower surface should be pale green, with simple hairs forming a fur, often over the whole surface, but especially on the veins. The inflorescence of Large-leaved Lime has one to five flowers hanging below the leaves whilst Small-leaved Lime ( T. cordata ) is quickly identified by an * Developed from a poster originally presented at the YNU’s Conference on “Recording and Monitoring Yorkshire’s Natural Environment", Harrogate, 26 February 2000. Naturalist 126 (2001) 140 Notes on the Distribution of Large-leaved Lime inflorescence held above the bract. The presence of plant galls on the leaves, such as the red nail gall Eriophyes tiliae tiliae, considerably assists a field worker to identify Lime trees in a fully leaved wood. The survey was principally undertaken between the middle of May and the end of July 1999. Ten areas were surveyed during the 1999 season: eight in Rotherham and two in Doncaster. Large-leaved Lime was found in all the Rotherham areas but only confirmed in one Doncaster woodland. The sites were mainly selected on the basis of previous records, but new areas were also inspected. The number of trees and their location are found in Table 1. TABLE 1. Woods surveyed for Large-leaved Lime in 1999. Wood Grid Square No. of trees Comments Rotherham: Roche Abbey: Figure 4 The Nor Woods SK59 3 Quarry Hills SK59 1 Table Rock SK58 8 King’s Wood SSSI SK58 33 1 young tree King’s Wood Lane SK58 13 In the hedges Slade Hill Plantation SK58 2 Maltby Wood Low Castle Lidget Quarter SK59 1 In cleared area Low Shoulder of Mutton Quarter, East of station SK59 5 Pieces Holt SK59 1 Sandbeck Park SK59 2 In parkland Firbeck Hall Anston Stones Wood SK58 Figure 5 1 In plantation Compartment 13 SK58 3 Compartment 15 SK58 2 Compartment 18 SK58 1 In cleared area Compartment 19 SK58 1 By the streamside Compartment' 22 SK58 22 Compartment 24 SK58 3 Compartment 26 SK58 1 Compartment 28 SK58 1 A young tree Compartment 32 SK58 3 Compartment 36 SK58 1 Compartment 39 SK58 1 Old Spring Wood SK58 3 Old Meadow Wood SK58 3 Hawk’s Wood SK58 1 Hooton Cliff Wood SK49 1 A coppice stool Doncaster: Edlington Wood SK59 2 At the north end Sprotborough Gorge SE50 0 Small-leaved Lime only confirmed Results The survey has confirmed the continuing presence of Large-leaved Lime in three 10 km grid squares (Figure 1) and estimated the population on the southern Magnesian Limestone in South Yorkshire. There are, however, only a few young trees, mainly on the edge of Notes on the Distribution of Large-leaved Lime 141 • Species confirmed in the 10 km square during the survey. O Report of species not confirmed in the 10 km square. FIGURE 1. The distribution of Large-leaved Lime in south Yorkshire, north Nottinghamshire and north-east Derbyshire, within the Magnesian Limestone Natural Area (1999). paths or woodland rides. One previously unknown site in the Rotherham area has been found and an additional site, unconfirmed by the author or the Record Centre, has been reported to the Doncaster Biological Records Centre. The number of Large-leaved Lime trees present in each of the woodlands has been recorded, and mapped on 1:10,000 ordinance maps. The one km grid map of the Rotherham area (Ligure 2) shows the distribution of Large-leaved Lime to the east of Rotherham and the south of Doncaster. A number of old records were clearly errors or could not be found, possibly due to forestry operations, e.g. in Anston Stones Wood Compartment 38. The Large-leaved Lime at Sandbeck Park is a large standard tree. Within the Roche Abbey group of woods there is a mixture of standard trees and many quite large and very old coppice stools with three to five large trunks sprouting skywards. The situation is similar at Anston Stones Wood and Maltby Wood. The trees in Old Spring Wood, Old Meadow Wood, Pieces Holt and Edlington Wood are all standards of various sizes. The single tree at Hooton Cliff Wood was felled in 1981 during forestry work, but has subsequently produced coppiced shoots. Additional fieldwork during the summer of 2000 added a new site in Doncaster, viz. Wadworth Wood (SK59). C. A. Howes (pers. comm.) reported one coppiced tree, but the 142 Notes on the Distribution of Large-leaved Lime • Species confirmed in the 1 km square. O Report of species not confirmed in the 1 km square. FIGURE 2. The distribution of Large-leaved Lime in Rotherham and south-east Doncaster by 1 km square. author and Howes failed to re-locate a record of Langridge (Meade & Langridge, 1987) in Engine House Wood, Sprotborough Gorge; the Lime here was all Tilia x europaea. Other trees found in these woods during the survey included Yew ( Taxus baccata ), Oak (' Quercus robur ), Wych Elm ( Ulmus glabra ), Ash ( Fraxinus excelsior ), Common Lime {Tilia x vulgaris), Beech ( Fagus sylvatica ), Guelder Rose {Viburnum opulus ), Field Maple {Acer campestre ), Holly {Ilex aquifolium ), Silver Birch {Betula pendula ), Spurge Laurel {Daphne laureola), Hawthorn {Crataegus monogyna ), Wild Service Tree {Sorbus torminalis ) and, additionally, in the Roche Abbey woods, Norwegian Maple {Acer platanoides). The survey also revealed that Small-leaved Lime {Tilia cordata) is quite scarce in Rotherham, small numbers being recorded in King’s Wood, Roche Abbey, Old Meadow Wood and Anston Stones Wood. Small-leaved Lime has been lost to road Notes on the Distribution of Large-leaved Lime 143 widening on the A63 1 through Maltby Wood. In Doncaster, two trees in Levitt Hagg Wood and three trees in Pot Ridings Wood were noted, whilst the species is common in the northern part of Edlington Wood. Elsewhere on the Magnesian Limestone, Clapham (1969) reported three sites where Large-leaved Lime was found in Derbyshire: Markland Grips, where on 31 May 1999, J. R. Comley and I found a single tree growing out of crags on the south-west side of the wood; Scarcliffe Park Wood, where we found one coppiced tree to the north-east in an area which was mainly plantation, both in SK57; and the Vale of Pleasley, Pigott (1981). There are also a small number of records from Nottinghamshire. We were able to confirm one coppiced tree at Scratta Wood, but were unable to confirm a further record from Cresswell Crags. Two other sites on the Magnesian Limestone in Nottinghamshire may have Large- leaved Lime, but we were unable to visit them to confirm the records. Lavin and Wilmore (1994) do not record the species north of Doncaster, in West Yorkshire. An appeal to the YNU by Newbould (1999) produced no positive sightings, although one member suggested searching woods in the West Tanfield area. Quadrat data have been collected for a representative selection of the sites (summarised in Table 2). Some of this is of limited value in that, for safety reasons, it was not possible to undertake 50 m2 quadrats. The quadrat data from Sandbeck reflect a parkland community, rather than woodland. From the survey, herbs present in the ground flora included Dog’s Mercury ( Mercuralis perennis ), Wood Anenome ( Anenome nemorosa ), Ransoms ( Allium ursinum), Bluebell ( Hyacinthoides non-scripta ), Tufted Hair-grass ( Deschampsia cespitosa ), Bramble ( Rubus fruticosus ), Wild Daffodil ( Narcissus pseudonarcissus ), Ivy ( Hedera helix), Violets ( Viola spp.) and Bearded Couch ( Elymus caninum). Bracken ( Pteridium aquilinum ) is associated with Large-leaved Lime on more acid soils such as in Edlington Wood, the western portion of King’s Wood and the eastern side of Maltby Wood. Additional records were derived from the Recorder database at Rotherham Biological Records Centre; Woolthwaite Bottoms, Loscar Wood, Swinston Hill Plantation and Bondhay Dyke by B. Langridge; Hell Wood by R. J. Hall; Langold Holt by C. P. Harding; Rough Park and Brancliffe Grange area by D. Bailey. C. A. Howes (pers. comm.) reported an additional tree, with a 5 m circumference, at Skelbrook Park, Doncaster in SE51. The author was unable to obtain permission to visit these sites. The results of this baseline survey work have been deposited with the B.S.B.I. ‘Atlas 2000’ project, Rotherham Biological Records Centre, Doncaster Museum and English Nature, Wakefield. Discussion Typically, the Large-leaved Lime was found in national vegetation type W8 woodland (Ash, Field Maple and Dog’s Mercury) (Rodwell, 1991). With the exceptions of Edlington Wood, King’s Wood, Maltby Wood and Old Spring Wood, the Large-leaved Lime was usually found on steep slopes or crags where magnesian limestone out-cropped. The steep slopes are useless for agriculture, grazing would be hard to manage, and forestry operations often difficult to undertake. Surprisingly, when analysing the data, Ash was found in only three of the quadrats. In both Anston Stones Wood, where in compartment 41 Ash saplings are extremely common under the Beech plantation, and also from the late 1960s in the Sycamore plantation alongside King’s Wood Lane, where Ash is almost a weed. Clearly, in areas where Lime- wood has been cleared, it would be necessary to monitor and probably remove Ash if such areas were to be restored to Lime-wood. The presence of Norwegian Maple in three of the four Roche Abbey quadrats could be of concern when, for example, hundreds of 1999 germinants were noted in King’s Wood. In the ground cover, Dog’s Mercury is a constant. The field notes obtained whilst collecting the data show the presence of this species at each site where Lime is recorded. Tufted Hair-grass featured in five of the quadrats; in woodland it occurs in soils of impeded 144 Notes on the Distribution of Large-leaved Lime TABLE 2. Quadrat data from South Yorkshire Lime-woods. Plant Site ~o o o £ Each plant is scored using the % domin scale see below £ Norwood 2 King’s Wood 1 King’s Wood 2 Anston Stones 22 Anston Stones 28 Edlington Wood Sandbeck Frequency Acer platanoides 3 3 3 II Acer pseudoplatanus 4 4 5 5 III Betula pendula 3 4 II Cornus sanguinea 2 I Corylus avellana 2 3 II Crataegus monogyna 1 2 4 1 III Daphne laureola 3 I Fagus sylvatica 4L 3 5 II Fraxinus excelsior 6 3 3 II Ilex aquifolium 3 1 11 Ligustrum vulgare 1 2 2 II Quercus petraea 4 I Quercus robur 4 3 II Sambucus niger 4 I Taxus baccata 4 6 4 II Tilia cordata 4 5 II Tilia platyphyllos 7 5 7 5 4 5 10 V Tilia x vulgaris 7 I Ulmus glabra 4 3 2 1 III Allium ursinum 3 1 4 II Anenome nemorosa 5 5 1 II Anthriscus sylvestris 3 I Arum maculatum 2 3 II Chamaenerion angustifolium 1 I Deschampsia cespitosa 3 3 2 1 2 IV Dryopteris filix-mas 1 I Filipendula ulmaria 1 I Hedera helix 4 1 1 6 III Heracleum sphondylium 1 1 II Holcus mollis 2 I Hyacinthoides non-scripta 4 4 2 2 1 IV Mercurialis perennis 5 5 5 4 4 6 2 4 V Myosotis arvensis 5 1 II Narcissus pseudonarcissus 1 8 II Phyllitis scolopendrium 1 I Potentilla sterilis 1 I Pteridium aquilinum 4 I Ranunculus auricomus 1 I Ranunculus ficaria s.s. 3 I Rosa arvensis 3 3 II Rubus idaeus 2 I Rubus fruticosus 1 1 1 3 4 7 IV 145 Notes on the Distribution of Large-leaved Lime TABLE 2 continued Quadrat data from South Yorkshire Lime-woods. Plant Site Each plant is scored using the domin scale see below '/ 0 1 2 3 4 5 Scale 0 10km FIGURE 5. Distribution of Greylag Geese in the Harrogate area in 1991. peaked in the late 1970s. Whilst some control methods are known to have been carried out (egg-oiling, egg-pricking, shooting and controlled culls), the number of birds has not decreased at a rate that can be explained by these alone. This is shown by the figures in Table 3, where the age ratios of Canadas on estates have remained fairly constant whilst those of Greylags have decreased at these sites. It is likely, therefore, that the population drop has occurred due to a combination of biotic and abiotic factors. One of these may be the massive national population increase in Greylags in some way affecting the Canada Goose population in the HNDS area. Whether this trend is national or regional is part of the reason for this survey, and at the time of writing the results are not available. It is a matter for debate as to whether the Greylag population will follow the same decrease as that of the Canada Goose, or if they will eventually find a population capacity and level out. This may be at the expense of the Canada Goose in the area, but due to differing habits and breeding habitats, it is likely that the two species will remain common in the Harrogate area for the foreseeable future. Acknowledgements The author would like to thank all those who gave up their free time to take part in this survey, especially Joan Owen, and Jill and Simon Warwick. In addition, many thanks to the estates who allowed access to their land for the survey, including Ripley Castle, Swinton Castle, Allerton Park and Studley Royal, and also to Yorkshire Water. 170 A Survey of Feral Geese in the Harrogate Area in the Year 2000 References Blurton-Jones, N. G. (1956) Census of breeding Canada Geese 1953. Bird Study 3: 153-170. Fabricius, E., Bylin, A., Ferno, A. and Radesater, T. (1974) Intra- and interspecific territorialism in mixed colonies of the Canada Goose Branta canadensis and the Greylag Goose Anser anser. Ornis Scand. 5: 25-35. Garnett, M. G. H. (1980) Moorland breeding and moulting of Canada Geese in Yorkshire. Bird Study 27: 219-226. Harrison, J. (1960) Goose conservation. WAGBI Annual Report 1959-1960. 28-29. Harrogate & District Natural History Society Reports 1991-1999. Kirby, J. S., Haines, W. G. and Austin, G. E. (1999) Translocations of Canada Geese Branta canadensis in Great Britain. Ringing & Migration 19: 261-271. Loonen, M. J. J. E., Zijlstra, M. and van Eerden, M. R. (1991) Timing of wing moult in Greylag Geese Anser anser in relation to the availability of their food plants. Ardea 79: 253-260. Mather, J. R. (1986) Birds of Yorkshire. Croom Helm, London. Ogilvie, M. (1969) The status of the Canada Goose in Britain 1967-96. Wildfowl 20: 75-85. Ogilvie, M. (1977) The numbers of Canada Geese in Britain. Wildfowl 28: 27-34. Owen, J. (1991) Introduced Goose Survey. In: Harrogate & District Naturalists’ Society Report 1991: 8-10. Owen, M. and Black, J. M. (1990) Waterfowl Ecology. Blackie, Glasgow and London. Owen, M. and Salmon, D. G. (1988) Feral Greylag Geese Anser anser in Britain and Ireland 1960-86. Bird Study 35: 37-45. Walker, A. F. G. (revisions by Kinnear, A. M. G) (1991) Birds in the Harrogate District. Harrogate and District Natural History Society. Willoughby, F. (1676) Ornithologia. John Ray, London. Wright, R. and Giles, N. (1988) Breeding Success of Canada and Greylag Geese Branta canadensis and Anser anser on Gravel Pits. Bird Study 35: 31-36. APPENDIX 1 List of all sites surveyed in 2000 (including grid reference) Site Grid Ref. Angram Reservoir Scar House Reservoir Gouthwaite Reservoir Roundhill Reservoir Leighton Reservoir Swinton Castle Lakes Marfield Nature Reserve Lumley Moor Reservoir Beaver Dyke and John O’Gaunts Reservoirs Eave stone Lake Scargill Reservoir Westwood Lake (West Tanfield) Sawley Lake Nosterfield Nature Reserve and Nosterfield North Gravel Pit Ripley Castle Lake Studley Royal & Fountains Abbey North Stainley Pond Lightwater Valley Bellflask Quarry Ripon Parks Ripon Racecourse SE040761 SE052770 SE128696 SE149769 SE 160786 SE205795 SE2 18822 SE223707 SE224545 SE226678 SE232535 SE258776 SE263677 SE275795 SE279608 SE280690 SE288767 SE292757 SE298773 SE304749 SE326698 Book Reviews 171 Site Grid Ref. Crimple Valley (Travellers Rest) SE333540 Rudding Park Lake SE337532 Copgrove Lake SE343633 Copt Hewick Hall SE348717 Farnham Gravel Pits SE350596 Barr Lane Quarry (Knaresborough) SE356590 Staveley Fish Ponds SE359625 Staveley Nature Reserve SE360630 Hay-a-Park Gravel Pits SE363582 Hutton Moor Closes Pond SE368723 Spofforth Sewage Farm SE373525 Roecliffe Brick Ponds SE382658 Allerton Park SE415585 Thorpe Underwood Fish Ponds SE457593 Thorpe Underwood Lake SE483608 BOOK REVIEWS Photographic Guide to the Sea and Shore Life of Britain and North-west Europe by Ray Gibson, Benedict Hextall and Alex Rogers. Pp. xiv + 436 with numerous coloured photographic plates, line drawings and maps. Oxford University Press. 2001. £35.00 hardback. £15.95 paperback with plastic cover. This book is a guide to the common animals and plants that can easily be seen with the naked eye without the need of habitat destruction, e.g. digging in sand and mud. A ‘conservation of specimens attitude’ is encouraged with identification in the field, or where this is not possible a sketch or photograph should be taken. A brief introduction is given to rocky, sand and muddy shores. All the species considered have been recorded in the British Isles. There is a short introduction to each taxonomic group with references for further study. There is a caution not to be surprised if a specimen cannot be identified but there is no indication of the number of species within each taxonomic group. There are no keys. Each species is illustrated in colour in its natural habitat with the following information: its English and biological name with taxonomic position, description supported by line drawings where needed, distinctive features, differences from similar species, geographical distribution with a map and ecological habitat. This book will be useful to the beginner and student. MEA Animal Tracks and Signs by Preben Bang and Preben Dahlstrom. Pp. 264 with some 240 colour photographs and over 300 drawings and diagrams. Oxford University Press. 2001. £12.50 hardback. This most practical of pocket guides, containing the masterly illustrations and diagrams by Preben Dahlstrom and very clear text re-edited by Martin Walters, makes it possible to identify a wide range of mammals and birds from the traces they leave behind, whether footprints, feeding debris, hiding places and homes, pellets and droppings. Originally a classic title in the familiar Collins Field Guide series, Animal Tracks and Signs is now issued as one of the new Oxford University Press Pocket Guides. My 1970s copy, now dilapidated and stained from close contact with countless muddy footprints, piles of excrement and pellets, and bearing the forensic evidence of many an engrossing rural excursion, is in need of replacement. Like the welcome return of an old and trusted 172 Book Reviews friend it is heartening to see this title re-issued and improved for the benefit of a new generation of naturalists. Though the art-work is virtually identical to earlier editions, improvements in reprographic technology substantially enhance the illustrations in the OUP version and the photographs have been supplemented by some very dramatic additions. A slight, though intriguing, problem with this guide is its significant reliance on tracks and signs in the snow. Since Bang and Dahlstrom compiled the first edition back in 1974 prolonged snowfall in Britain has become somewhat less frequent - a field sign in its own right, though of a somewhat larger footprint. Even if readers have the old edition in their rucksacks, I would recommend replacing it with this new improved version - but save the old dust jacket featuring Dahlstrom’s fabulous art work, re-using it to mask the drab corporate formality of the OUP covers. CAH The New Encyclopaedia of Mammals edited by David Macdonald. Pp. 960 with over 960 colour photographs and over 700 drawings and diagrams. Oxford University Press. 2001. £35.00 hardback. With a team of eight world class mammalogists as advisory editors supported by 240 academic experts (including Prof. M. J. Delany, the editor of Yorkshire Mammals ) as sectional authors, Professor David Macdonald has compiled a truly encyclopaedic review of the lives, evolution, adaptations, ecology, behaviour and social structures of the 2 sub- classes, 27 orders, 139 families, over 1,100 genera and some 4,600 species of mammals of the world. Since 1984 when the first edition was assembled. Professor Macdonald’s team have collated the veritable deluge of new discoveries resulting from a revolution in new techniques for the study of mammals. This, with the benefit of word processing and access to the internet, not generally available two decades ago, has resulted in the extensive revision and in most cases the entire re-writing of most sections of the encyclopaedia. The work covers all species from the smallest mammal, Kitty’s hog-nosed bat weighing 1.2 g, to the blue whale which weighs 100 million times as much. Interesting changes in the sequence of the mammalian orders have occurred since the earlier edition; for example, the Golden moles and Tenrecs are now placed with the Hyraxes and Elephants in a new super-order, the Afrothernia. Arranged generally according to orders and families, the book introduces at appropriate places a range of case studies and special features which illustrate a wide range of fascinating biological phenomena. These make compelling reading and are given both magazine-style and technical headings such as ‘Mammalian central heating - endothermy and its costs’; ‘Body size, the cost of living and diet - nutrition and metabolism’; ‘Size and the energy crisis - allometry’. With quick access ‘Factfile’ panels and with compelling yet authoritative text well punctuated with punchy subject headings, each of the eighteen chapters serve the needs of pressured students with essay deadlines, yet allows the general reader to wallow in this rich, stimulating and fast developing subject. This truly luxurious volume is remarkable in combining glamorous magazine-style presentation with critical academic rigour. It succeeds in placing the best in reprographic technology, high quality artwork and exciting prose style at the service of a very broad readership. The New Encyclopaedia of Mammals is a standard work ideally designed for any school, college or public library yet at a price which puts it within reach of many a domestic bookshelf - though weighing 4.5 kg, some DIY shelf reinforcement may be necessary! CAH 173 LICHEN FLORA OF COUNTY DURHAM (VC66): SUPPLEMENT 1 D. E. McCUTCHEON Normans Riding Poultry Farm, Winlaton, Blaydon-on-Tyne, Tyne & Wear NE21 6LA Since the publication of the Lichen Section in the Flora and Vegetation of County Durham (Graham, 1988), a systematic survey of VC66 has taken place and a considerable number of new species have been found. A number of species not seen for over 100 years have now been refound in the county. This is therefore a convenient time to produce an update. Where the distribution of species has changed considerably since that known in 1988, this has been commented upon. The vast majority of the records are the results of my own investigations and are not credited. Where joint ventures were involved both recorders are credited. Other recorders are duly credited. G. G. Graham’s herbarium has been examined and several changes have been noted under the appropriate species. Unfamiliar material has been either confirmed or determined by colleagues in the Brilish Lichen Society and this is noted. More recently, leprose crusts have received greater scrutiny from notable taxonomists and the parameters for identification of species have moved considerably. A reasonable attempt has heen made to keep up with the evolving situation and in particular the devolution of Lepraria incana which in the past has been a convenient catch-all. Several new species have been segregated from it, but it still remains the commonest member of the group and its widespread distribution remains valid. This area of study remains a major challenge for future lichenologists. The occurrence of lichenicolous fungi or “parasites” has scarcely been noted and the time is ripe for further investigation, especially as recent discoveries in Northumberland are proving to be so exciting. 84 species are reported as being new to VC66. 14 species not seen for many years now have a modern record. The total of reliably recorded species now stands at 604. Two species of Parmelia collected by W. Mudd have been commented upon but not added to the county list as they may have been collected from VC65. Nomenclature follows Purvis et al. (1993). E = Herbarium, Royal Botanical Garden, Edinburgh. * = new species, including a few previously rejected because of insufficient supporting data. • = modern records of species not seen for a very long time. *Acarospora cervina A. Massal. Ledge on limestone outcrop, Coldberry End, 35/83, 1998, conf. B. J. Coppins. * A. Sinopica (Wahlenb.) Korb. On levelled colliery shale on the backshore, S. of Noses Point, Seaham, 45/44, 1997. A. smaragdula (Wahlenb.) A. Massal. Now known to be frequent in the W. of the county, especially on metalliferous rocks. * Acarospora verruciformis H. Magn. Rare on metal-rich boulders in the vicinity of old lead mines. First record Killhope Burn, 35/84, 1997; Scaud Hill, 35/83, 1997. Agonimia tristicula (Nyl.) Zahlbr. Sparse but widespread. Generally associated with decaying mosses on limestone. Recorded from 8 squares. Amygdalaria pelobotryon (Wahlenb.) Norman. On several acidic stone walls in the Derwent Vallen, 35/94, 1992. Anisomeridium polypori (Ellis & Everh.) M. E. Barr. Widespread. Generally in woodland and often on mature elders. * Arthonia arthonioides (Ach.) A. L. Sm. At base of vertical acidic cliffs, Gibraltar Point, 35/94, 1990, det. B. J. Coppins. A. didyma Korb. On smooth bark of old hazel, Lower Hisehope, 45/04, 1990, G. G. Graham; Westernhopeburn Wood, 35/93, 1994; Slit Wood, also 35/93, 1997. * A. lapidicola (Taylor) Branth & Rostr. On iron tube railings next to concrete post in wooded valley, West Butsfield, 45/14, 1997, conf. B. J. Coppins. Naturalist 126 (2001) 174 Lichen flora of County Durham (VC66): Supplement 1 Anaptychia ciliaris (L.) Korb. Not seen during the present survey and feared to be extinct in the county. Arthrorhaphis grisea Th.Fr. On Baeomyces rufus in recess on bankside, Lady’s Rake Mine area, 35/83, 1997. Aspicilia caesiocinerea (Nyl. ex Malbr.) Arnold. Now known from 9 squares, mostly in the western uplands of the county. * A. grisea Arnold. On dolerite block in wall, Force Garth, 35/82, 1996, det. B. J. Coppins. * A. laevata (Ach.) Arnold. Now known from 35/93, 35/94 & 45/05 on damp rocks in and near streams in woodland. First modern record Nookton Burn Woods, 35/94, 1990. Bacidia arceutina (Ach.) Arnold. Base of ash on dry bank of stream, Crimdon, 45/43, 1997; low on boles in shaded woodland. Bishop Middleham Quarry, 45/33, 1998; on bleached sheep bone on open moorland, Muggleswick Common, 45/04, 1992. B. bagliettoana (A. Massal. & De Not.) Jatta Over mosses in old limestone quarry, Wingate, 45/33, 1997. B. caligans (Nyl.) A.L.Sm. Decaying bark on larch fence post, Nookton Burn, 35/93, 1991; on moss and soil on basic outcrop, Killhope, 35/84, 1997; similar substratum, Coldberry End, 35/83, 1997. * B. chloroticula (Nyl.) A. L. Sm. An early coloniser of the tops of shaded fence posts, gates, benches, etc. Widespread in the E. of the county and recorded from 9 squares. First record from tops of fence posts near Winlaton, 45/16, 1996. * B. naegelii (Hepp) Zahlbr. First record from shaded old Populus by the R.Tees near Whorlton, 45/01, 1988, det. B. J. Coppins. Now known also a 35/93, 35/94 & 45/1 1 . B. rubella (Hoffm.) A. Massal. Very local but not uncommon in the sterile state. Great Wood and Shipley Wood, 45/02, 2000, O. L. Gilbert & D. E. McC. * B. saxenii Erichsen. Cut stump of conifer, Chopwell Wood, 45/15, 1996, det. B. J. Coppins; stone in sheep pasture, Westernhopeburn Farm, 35/93, 1997. B. subfuscula (Nyl.) Th.Fr. Top of sandstone retaining wall below the Tyne Bridge, Gateshead, 45/26, 1997. * B. viridescens (A. Massal.) Norman. On moss and soil in old mineworks. First record at Bolts Burn, 35/94, 1991; also in other similar sites nearby and at Marl Beck, 35/92, 1997. * B. viridifarinosa Coppins & P. James. Thinly spread throughout the county, although sometimes locally abundant. On shaded boles and rocks, etc. in generally moist situations. First record on shaded bole of birch, Nookton Burn Woods, 35/94, 1991 . * Bryoria capillaris (Ach.) Brodo & D. Hawksw. On bole of mature oak near R. Derwent, Gibraltar Point, 35/94, 1992, conf. D. Hawksworth, who commented that the bark was probably acidified. Caloplaca isidiigera Vezda. Sandstone rock on ground next to gate into pasture, Lamb Shield, 45/04, 1997. * C. marina (Wedd.) Zahlbr. ex Du Rietz. Sparse to frequent on sea defences and piers the whole length of the coast. Very rarely on harder ridges of Magnesian Limestone. Generally in the higher area of the splash zone. First record on concrete sea wall at Seaham, 45/45, 1997. The appearance of this, along with other maritime species, is almost certainly a consequence of the major clean up of the Durham coast during the last decade or so. * C. obscurella (Lahm ex Korb.) Th.Fr. First record from the bole of a huge Populus near the R. Tees in woodland, S. of Sledwich Flail, 45/01, 1988, D. E. McC. & G. G. Graham, det. B. J. Coppins 1988; also found in three woods in the Upper Derwent Valley, 45/04 & 45/05, during the 1990s. * C. ochracea (Schaer.) Flagey. Hard part of Magnesian limestone rock at base of cliff, S. of Noses Point, 45/44, 1997. * C. teicholyta (Ach.) J. Steiner. South-facing Magnesian limestone outcrop next to road, Trow Point, 45/36, 1997. C. variabilis (Pers.) Miill.Arg. On hard limestone blocks in walls and natural outcrops in the hills. Now known from 35/83, 35/92, 35/93 & 35/94. Candelariella reflexa (Nyl.) Lettau. Generally on nutrient-rich fairly well-lit trees. Now Lichen flora of County Durham (VC66): Supplement 1 175 known from 35/82, 35/93, 45/01, 45/05, 45/13 & 45/16. * Carbonea vorticosa (Florke) Hertel. Vertical face of siliceous rock in wall. Force Garth, 35/82, 1996. Catillaria atropurpurea (Schaer.) Th.Fr. At base of old ash in Shipley Wood, 45/02, 2000, D. E. McC. & O. L. Gilbert. * C. modesta (Miill.Arg.) Coppins. Shaded vertical hard limestone outcrop in the hills, Upper Teesdale, 35/83, 1997, conf. B. J. Coppins (E). New to England. Cetraria pinastri (Scop.) Gray. This species persisted for three or four years, producing soredia which developed into thirteen tiny thalli. After a break of two years, the site was revisited: the boulder that hosted the specimens was now well colonised by Parmeliopsis ambigua (Wulfen) Nyl., but with no trace of Cetraria pinastri which must now be considered extinct. It was strange that it had been replaced by a species which produces the same coloured soredia. (Could it possibly have been stolen ?) Chaenotheca chrysocephala (Turner ex Ach.) Th.Fr. Now known from 9 woodlands centred on the Derwent Valley. C. trichialis (Ach.) Th.Fr. Now known from 12 sites, generally in deeply fissured bark of oak, but also ash. 35/93, 35/94, 45/04, 45/05, 45/15 & 45/16. * Chaenothecopsis nigra Tibell. On shaded rootstock of massive old oak overhanging rock face near R.Tees, Shipley Wood, 45/02, 2()00, D. E. McC. & O. L. Gilbert, det. B. J. Coppins. * Chrysothrix flavovirens Tpnsb. Dry bark of mature oak on stream bank, near Tunstall Reservoir, 45/04, 1997. Cladonia diversa Asperges. All Durham records for C. coccifera (L.) Willd. are now considered to belong here. C. rangiformis Hoffm. Now known to be widespread in the W. of the county in short turf and sparsely vegetated old spoil heaps, etc. C. scabriuscula (Delise) Nyl. As the previous species, but appears to tolerate rather taller vegetation. Recorded from 13 squares. * C. strepsilis (Ach.) Vain. On wet peat, Bolts Law, 35/94, 1994. * C. sulphurina (Mich.) Fr. On peat in the hills. Castleberry Cleugh, 35/94, 1992; also at Swinhope Head, 35/83, 1997. * Cyphelium inquinans (Sm.) Trevis. On gatepost of cattle grid, near Harwood, 35/83, 1997; on bole of mature ash near stream, Shipley, 45/02, 2000, D. E. McC. & O. L. Gilbert. Cystocoleus ebeneus (Dillwyn) Thwaites. Now known from 15 sites with a westerly distribution. Always in shaded acidic situations generally near rivers or streams. * Dermatocarpon intestiniforme (Korb.) Hasse In seepage track of basic rocks on steep hillside, Widdybank Fell, 35/82, 1985, conf. O. L. Gilbert. * D. luridum (With.) J. R. Laundon. Submerged on dolerite at the edge of the R.Tees, Upper Teesdale, 35/82, 1993, O. L. Gilbert & V. J. Giavarini. * D. meiophyllizum Vain. On dolerite at the edge of the R.Tees, Upper Teesdale, 35/82, 1993, O. L. Gilbert & V. J. Giavarini. Diploschistes scruposus (Schreb.) Norman. Now known from numerous walls in the western uplands. Diplotomma alboatrum (Hoffm.) Flot. On limestone rocks, walls and mortar. Sparsely but widely distributed. Recorded from 10 squares. Enterographa crassa (DC.) Fee. Saxicolous at Castle Eden Dene, 45/43, 1997. * E. hutchinsiae (Leight.) A. Massal. Deeply shaded underhang at base of crag near waterfall, Allensford, 45/05, 1996, det. B. J. Coppins; on rock underhang at top of gorge, Westernhopeburn, 35/93, 1997. E. zonata (Korb.) Kallsten. Now known to be widespread on shaded rocks and walls, etc., very rarely on rootstock. * Epigloea soleiformis Dobbeler. On moss over dolerite block in wall. Force Garth, 35/82, 1996, D. E. McC. & C. J. B. Hitch, det. C. J. B. Hitch; near top of shaded rock amongst bank of boulders, Killhope, 35/84, 1997, D. E. McC. & C. J. B. Hitch, 1997, det. C. J. B. Hitch. 176 Lichen flora of County Durham (VC66): Supplement 1 Fuscidea lygaea (Ach.) V. Wirth & Vezda. On hard acidic rocks, often in walls, in the uplands. Saltersgate, 45/04, Killhope, 35/84 & Westernhopeburn, 35/93, all 1997. F. praeruptorum (Du Reitz & H. Magn.) V. Wirth & Vezda. Widespread in the uplands on vertical faces of acid rocks in walls. * F. recensa (Stirt.) Hertel, V. Wirth & Vezda. Sandstone wall in Derwent Gorge, 45/04, 1977, G. G. Graham & B. J. Coppins; acidic boulder in high moorland with Allantoparmelia alpicola (Th.Fr.) Essl., Dead Stones, 35/73, 1980, G. G. Graham & R. W. Corner, det. D. E. McC. 1997. Hymenelia prevostii (Duby) Kremp. Hard part of Magnesian limestone boulder at base of cliffs on the backshore, S. of Noses Point, 45/44, 1997. * Hyperphyscia adglutinata (Florke) H. Mayrhofer & Poelt. On the bark of dying elm, Stanhope, 35/93, 1989. The elm has now gone, but there are many ash trees present in Weardale and it is possibly still present in that area. Hypocenomyce caradocensis (Leight. ex Nyl.) P. James & G. Schneider. Now known from several sites in the Upper Derwent Valley woodlands, all on old oaks, 35/94, 45/04 & 45/05, 1990s. * Ionaspis heteromorpha (Kremp.) Arnold. On hard limestone rock at top of gorge, Westernhope, 35/93, 1997; on hard smooth limestone, semi-inundated at edge of Rough Syke, 35/83, 1998. Lecania cyrtella (Ach.) Th.Fr. Now known to be widespread and recorded from 8 squares, usually on elder. L. erysibe (Ach.) Mudd. Now known from at least 20 squares, on natural basic substrates and more usually on mortar, gravestones, etc. * L. inundata (Hepp ex Korb.) M. Mayrhofer. On top of carved weakly calcareous sandstone boulder, car park at Winlaton Mill, 45/16, 1996. * L. rabenhorstii (Hepp) Arnold. Around cracks on harder parts of Magnesian limestone cliffs, Souter Point, 45/46, 1997, conf. B. J. Coppins. Lecanora aitema (Ach.) Hepp. Now known to be widespread in the upland areas of the county, often on lignum, but rarely extremely abundant on exposed conifer twigs and branches, along the edges of fire lanes, etc. Often included in L. symmicta, of which most past records almost certainly belong here. * L. caesiosora Poelt. On acidic rocks in walls in the uplands. Recorded from 9 upland squares throughout the 1990s. Most well colonised walls will host this species. Found once in fruit at Hunstanworth churchyard, 35/94. First record on sandstone wall, Westernhope, 35/93, 1988. L. carpinea (L.) Vain. On older trees and now known from a dozen sites in 6 squares, 35/83, 35/93, 35/94, 45/05, 45/13 & 45/23. L. epanora (Ach.) Ach. Sandstone wall, W. of Lanchester, 45/14, 1979, G. G. Graham Since found on several walls and rocks, some moderately polluted, 35/94, 45/14, 45/15 & 45/22. * L. horiza (Ach.) Linds. A single thallus on lignum, Westernhope, 35/93, 1997; twig of Sambucus at edge of wood, Allensford, 45/05, 1997. * L. persimilis (Th.Fr.) Nyl. On rose on roadside verge, Ireshopeburn, 35/83, 1997; on ash sapling on roadside, Westgate, also 35/83, 1997, det. B. J. Coppins, who comments that it is a common species with a preference for ash. Possibly overlooked as L. dispersa. L. pulicaris (Pers.) Ach. Now known to be frequent in the N/W of the county, rare elsewhere. Generally on smooth bark along the valley woodlands, but also on fence posts, etc. Recorded from 12 squares. L. saligna (Schrad.) Zahlbr. Horizontal surface of gate, near Esh Winning, 45/14, 1994, P. A. R. J. Stevenson. L. symmicta (Ach.) Ach. Generally on the nodes of twigs and branches of trees and shrubs. It has been reliably been recorded in the 1990s from 35/84, 35/93, 35/94 & 45/16; see L. aitema. * Lecidea auriculata Th.Fr. Very rare. A specimen of L. confluens from the collection of Lichen flora of County Durham (VC66): Supplement 1 111 G. G. Graham has been redetermined as this species and is consequently the first county record. Acid rock on high moorland, Lintzgarth, 35/94, 1971, G. G. Graham, det. D. E. McC; sandstone windowsill below iron grill, Hunstanworth church, 35/94, 1991; quartz projection on rock on top of wall. Stanhope Common, also 35/94, 1994, conf. O. L. Gilbert. L. diducens Nyl. On acid boulders and stone walls in the hills. Recorded from 9 western squares. * L. lactea Florke ex Schaer. Rare on well-lit acid rocks in the hills especially walls. On rock near top of wall, Westernhopeburn Farm, 35/93, 1988; sandstone rocks in walls, Nookton and Norham Burns, 35/94, 1991, 45/05, 1992 & 35/83, 1998. L. plana (Lahm) Nyl. Now known to be frequent to abundant on acid rocks and walls in hilly areas. Recorded from 12 generally westerly squares. L. lapicida (Ach.) Ach. Very rare. The two records in tile Durham Flora were both redetermined as L. plana , det. D. E. McC. 1997. Only seen on acid rock in wall, Wharnley Burn, 45/04, 1990. * L. pycnocarpa (Korb.) Ohlert. Local in the western hills, on acid rocks usually in walls with a preference for flatter rocks in sheltered, but well-lit moist situations. First record acid stone at top of wall at edge of wood, Thornhope Moor, 45/03, 1987. Forma sorediata Coppins & Friday has been found only once on acidic stone in wall near stream, Ladys Rake Mine (disused), 35/83, 1997. * Lecidella elaeochroma f. soralifera (Erichsen) D. Hawksw. Very rare on mature old trees. Sycamore beside R.Wear, Stanhope Bridge, 35/93, 1994, G .G. Graham; base of well-lit ash on bankside, Rabbit Bank Wood, near Consett, 45/14, 1997; knurl of pollarded old ash on riverbank, Witton Bridge, 45/13, 1998. * Lecidoma demissum (Rustr.) G. Schneider & Hertel. Soil pocket on shaded ledge in acid wall, Nookton Burn, 35/94, 1991. A specimen collected from Widdybank Pasture in 1973 named as Schaereria cinereorufa was redetermined as this species by D. E. McC. 1997. * Lepraria cacuminum (A. Massal) Lohtander. Upper Teasdale, 35/82, 1996, conf. C. J. B. Hitch. On moss on dolerite block in wall. Force Garth, Upper Teasdale, 35/82, 1996, conf. C. J. B. Hitch. L. caesloalba (de Lesd.) J. R. Laundon. On well-lit acidic outcrops and walls, often on moss, with a stronghold in Upper Teesdale. Seen at 7 sites in 35/82, 35/83, 35/84, 35/92 & 45/04, 1996/7. * L. elobata Tpnsb. Near top edge of semi-shaded rock on bankside of boulders, Killhope, 35/84, 1997, det. C. J. B. Hitch & P. M. Earland-Bennett; dangling roots on bankside of old railway line, Saltersgate, 45/04, 1997. * L. lobif leans Nyl. Widespread and frequent on mossy rocks and tree bases in medium shade. Generally as fluffy thumbnail-sized light green thalli. L. nivalis J. R. Laundon. On shaded faces of limestone outcrops in stream valleys. On sheer limestone faces in the gorge of the Stanhope Burn, 35/94, 1994; also in 35/83 & 35/94; 1997. * L. rigidula (Hue) Tpnsb. In shaded acidic rocky underhang amongst boulders on bankside, Killhope, 35/84, 1997, D. E. McC. & C. J. B. Hitch, det. C. J. B. Hitch; similar situations at Nookton Burn Woods, 35/94, 1997 & Hamsterly Forest, 45/02, 1998. * Leproloma diffusum var. chrysodetoides J. R. Laundon. On bare bank in young plantation on undulating quarry floor, Quarry Wood, 45/04, 1997, conf. A. Orange. Leproloma membranaceum (Dicks.) Vain. Rare, but true status unclear as past records should be treat with caution; confused in the past with other leprose species. * L. vououxii (Hue) J. R. Laundon. Widespread on shaded limestone rocks and walls. Recorded from 12 squares from east to west. First noted on shaded vertical rock face (weakly calcareous), Ravens Crag, 45/51, 1996. Lobaria pulmonaria (L.) Hoffm. Healthy colonies on a few trees in Great and Shipley Woods, 45/02, 2000, O. L. Gilbert & D. E. McC. L. virens (With.) J. R. Laundon. Almost certainly extinct as the host tree is believed to have fallen into the river. 178 Lichen flora of County Durham ( VC66 ): Supplement 1 Micarea bauschiana (Korb.) V. Wirth & Vezda. Now known from 7 modern sites, always in shaded underhangs generally near water, 35/84, 35/94, 45/02 & 45/04. M. botryoides (Nyl.) Coppins Always in deep shade in acidic conditions. Now known from numerous sites in 7 squares. M. denigrata (Fr.) Hedl. Now known to be very common on lignum and rarely on stone. * M. erratica (Korb.) Hertel, Rambold & Pietschm. Rare on hard smooth rocks and lignum. Hard stone on open ground near path, Hisehope Burn, 45/04, 1990, D. E. McC. & G. G. Graham; hard pebble on old colliery site, Sacriston, 45/24, 1996; old pallet on waste ground near Newburn Bridge, 45/16, 1996, det. C. J. B. Hitch. * M. lithinella (Nyl.) Hedl. Rare on stones near tracks. On half buried stone at edge of moorland track, 35/94, 1993, conf. B. J. Coppins; sandstone block in corner of field next to farm road, Leadgate, 45/15, 1996; sandstone rock by stream, N. of Tunstall Reservoir, 45/04, 1997. * M. myriocarpa V. Wirth & Vezda ex Coppins. On shaded sandstone rocks, generally in damp valleys. Known from 4 sites in the Upper Derwent Valley and one in Weardale. First record on shaded sides of sandstone boulders by R. Derwent, Allensford, 45/04, 1995, conf. B. J. Coppins. M. peliocarpa (Anzi) Coppins & R. Sant. Shaded back of acidic rock on hillside, Killhope, 35/84, 1997; stump in woodland, near Winlaton, 45/16, 2000; with Trapelia coarctata on siliceous rock on exposed moor, Feldon Carrs, 45/04, 1977, G. G. Graham, det. D. E. McC. 1997. M. prasina Fr. Now known to be very common in woodland. M. sylvicola (Flot.) Vezda & V. Wirth. Sandstone boulder on riverbank, near Muggleswick, 45/05, 1989, conf. B. J. Coppins; acidic rocky underhang, N. of Tunstall Reservoir, 45/04, 1997. * Moelleropsis nebulosa (Hoffm.) Gyeln. Very rare, on edge of small bank in old mine area on the bank of stream. Slit, 35/93, 1997, conf. A. Fryday. * Mycoblastus sanguinarius (F.) Norman. Now known from a cluster of 5 sites in the Upper Derwent catchment and one adjacent square in Weardale. On acidic rocks and trees reasonably well-lit but sheltered to some degree. Trunk of oak in valley woodland, Hisehope Burn, 45/04, 1985; locally frequent on acidic capstones of boundary wall of wood, Thornhope Moor, 45/03, 1987; also occurs in 45/05. * Omphalina ericetorum (Fr.) M. Fange ex H. E. Bigelow. Probably frequent to abundant on wet peat and moribund bryophytes on rocks. First fertile record on wet peat, with much sterile material in the vicinity, Swinhopehead, 35/83, 1987. * O. pseudoandrosacea (Bull.) M. M. Moser. On peat on hilltop near heaps of iron-rich rocks, Scaud Hill, 35/83, 1997. * Opegrapha mougeotii A. Massal. Shaded limestone cliffs, Ryehope Dene, 45/45, 1997; a repeat of an 1825 record. O. ochrocheila Nyi. Shaded base of ash, Westernhopeburn Wood, 35/93, c. 1992; shaded sandstone underhang near stream. Slit Wood, also in 35/93, 1997. O. dolomitica (Arnold) Clauzade & Cl. Roux. Fimestone rocks near old mine site, Swinhope, 35/83, G. G. Graham, 1981, det. D. E. McC. 1997; shaded limestone cliffs in stream gorge, tributary of Harwood Beck, also in 35/83, 1997. O. rupestris Pers. Parasitic on Verrucaria baldensis on harder part of limestone cliff, Hawthorn Hive, 45/44, 1997. O. rufescens Pers. At base of elm on riverbank. Great Wood, 45/02, 1997. O. vulgata (Ach.) Ach. The most frequent member of the genus having been recorded in 10 squares, mostly in the west. Pachyphiale carneola (Ach.) Arnold. Very local but present in quantity on several old ash and oak in Shipley and Great Woods, 45/02, 2000, O. F. Gilbert & D. E. McC. Parmelia caperata (F.) Ach. A single small thallus on sycamore, car park, Harperly Hotel, 45/15, 1997. Lichen flora of County Durham (VC66): Supplement 1 179 P discordans Nyl. Now known to be locally frequent on acid boulders on the high western moorland. Recorded from four upland squares. P. elegantula (Zahlbr.) Szatala. Alder bole on stream bank, Snowhope Close, 35/93, 1997; bole of mossy sycamore, Hardwick Country Park, Sedgefield, 45/32, 1998. * P. exasperata de Not. Hawthorn twig on woodland margin, Rookhope Burn, 35/94, 1994. P. incurva (Pers.) Fr. Locally frequent, generally on walls in the hills. At 10 sites in six western squares. P. laevigata (Sm.) Ach. Without details other than collected in Teesdale by W. Mudd, det. B. J. Coppins, 1991; see P. sinuosa. P. mougeotii Schaer. ex D. Dietr. On siliceous boulder on heather moor, Muggleswick Common, 45/04, 1997; also two sites in 35/94. P. sinuosa (Sm.) Ach. Two specimens of this species collected in Teesdale by W. Mudd with no details were detetermined by B. J. Coppins in 1991 from a collection ex herb. P. B. Mason lodged at Bolton Museum. Peltigera canina (L.) Willd. Infrequent, but found in 9 western sites, generally in short turf. * P. degenii Gyeln. Very rare, but locally frequent in one site. On mossy boles of fallen trees and mossy rocks in river gorge woodland, Derwent Gorge, 45/04, 1991, conf. B. J. Coppins. New to England. A specimen of a Peltigera sp. collected by G. G. Graham & B. J. Coppins 20 years previously had a scrap of this species attached, det. D. E. McC. 1997; it was collected a few hundred yards downstream, showing evidence of colonisation for some considerable time at this site. P. didactyla (With) J. R. Laundon. Now recorded from 14 sites in central and western parts of the county. P. leucophlebia (Nyl.) Gyeln. Limestone outcrop on hilltop, Coldberry End, 35/83, 1997. * P. neckeri Hepp ex Mull.Arg. Locally frequent in short turf on old railway lines and old mine sites in the upland west. First record from short grass at edge of Bolts Burn, Ramshaw, 35/94, 1990; Waskerly Way, 45/04, 1997. Now known from 12 sites in 35/83, 35/84, 35/93, 35/94, 45/03 & 45/04. * P. polydactyla (Necker) Hoffm. Distribution closely follows the previous species. Known from 10 sites in the same areas. First record loose scree and soil on spoil heaps, Jeffries Rake, 35/94, 1 965, G. G. Graham, conf. P. W. James. Pertusaria coccodes (Ach.) Nyl. Ash bark in gorge woodland, Westernhopeburn Wood, 35/93, 1997. P. aspergilla (Ach.) J. R. Laundon. Most frequent in the west on acid rocks and especially walls in the hills. Known from 9 sites in 35/82, 35/83, 35/84, 35/92, 45/14 & 45/15. P. lactea (L.) Arnold. In habitats similar to the preceeding species. Found at 11 sites in 35182, 35/83, 35/92, 35/94 & 45/05. P. leioplaca DC. Now known to be locally frequent on smooth bark in the moist wooded valleys in the western half of the county. P multipuncta (Turner) Nyl. On old rowan in river valley, Beldon Burn, 35/94, 1991. * Petractis clausa (Hoffm.) Kremp. On shaded limestone rock in Castle Eden Dene, 45/43, 1997. * Phaeophyscia sciastra (Ach.) Moberg. On dolerite at the edge of the R.Tees, Upper Teesdale, 35/82, 1993, O. L. Gilbert & V .J. Giavarini. * Phaeopyxis varia Coppins, Rambold & Triebel. Parasitic on Trapeliopsis gelatinosa, Force Garth, 35/82, 1996, det. B. J. Coppins. Physcia aipolia (Ehrh. ex Humb.) Furnr. On dying elm, Crawleyside Bank, Stanhope, 35/93, 1988; nutrient-enriched bark on bank of R.Tees, Neasham, 45/31, 1998; large wayside ash near stream, Shipley, 45/02, 2000, O. L. Gilbert & D. E. McC. Physconia perisidiosa (Erichsen) Moberg On nutrient-rich bark of large ash in picnic area, Allensford, 45/05, 1992; also in 45/04, 1992. Placopsis gelida (L.) Linds. Now known from 1 1 sites in the W. of the county, often on old mine sites. However, all these records probably refer to P. lambii Hertel & V. Wirth (B. J. Coppins, pers. comm.). 180 Lichen flora of County Durham (VC66): Supplement 1 * P. lambii Hertel & V. Wirth. Status unclear owing to confusion with previous species. On dolerite boulders amongst juniper scrub. Force Garth, 35/82, 1996. * Polyblastia cruenta (Korb.) P. James & Swinscow. On dolerite at the edge of the R.Tees, Upper Teesdale, 35/82, 1993, O. L. Gilbert & V. J. Giavarini. Polysporina simplex (Davies) Vezda. Sandstone wall at Baal Hill House, 45/03, 1998, G. G. Graham & D. E. McC.; coarse-grained sandstone rock in wall, Pow Hill, 45/05; 1997; sandstone wall of old workings on hillside, Stanhope Burn, 35/94, 1998. Porina aenea (Wallr.) Zahlbr. Now known to be widespread on smooth bark in sheltered moist woodland. Concentrated mainly in the major river valleys of the west. P. chlorotica (Ach.) Miill.Arg. On shaded rocks with a similar distribution to the previous species, but much less frequent. Porpidia cinereoatra (Ach.) Hertel & Knoph. Most previous records of Huilia albocaerulescens will belong here. On rocks and walls in usually moist situations and normally in the hills. At numerous sites in 8 squares in the west, with an outlier in 45/24. * P. contraponenda (Arnold) Knoph & Hertel. Rare on acid rocks on tops of walls in the uplands. Acidic stones in wall top (sheltered by conifers), Stanhope Burn, 35/94, 1994; on dolerite block in wall. Force Garth, 35/82, 1996. * P. hydrophila (Fr.) Hertel & Schwab. Very rare on top of siliceous boulder on bank of R.Tees, partially shaded by trees. Great Wood, 45/02, 1997. P platycarpoides (Bagl.) Hertel Emergent acidic rock, Horsleyhope Burn, 45/04, 1992, det. B. J. Coppins. Slightly basic sandstone rocks on riverbank. Great Wood, 45/02, 1997. P. soredizodes (Lamy ex Nyl.) J. R. Laundon. Now known from numerous widespread sites. Often found on flat stones in old mine sites, and frequently on gravestones by D. H. Smith. Difficult to separate from P tuberculosa, but usually in small neat patches. Protoblastenia calva (Dicks.) Zahlbr. Rare on pure limestone rocks on hill-top, Coldberry End, 35/83, 1997. * P. incrustans (DC.) J. Steiner. Rare on pure limestone. On limestone rocks, Tarnhole Edge, Teesdale, 35/83, 1993, G. G. Graham & P. S. Graham; on limestone block in wall on road into the hills, Ireshopeburn, also 35/83, 1997. Pseudevernia furfuracea (L.) Zopf. Both varieties occur in the county, but distribution uncertain as the vast majority of records were assumed in the past to be var. ceratea. * Psilolechia leprosa Coppins & Purvis. Widespread in the county under metal run-offs. Found in many churchyards by D. H. Smith; also accompanied by abundant Stereocaulon nanodes on metalliferous clinker on riverbank, Ryton Willows, 45/16, 1997. * Psoroma hypnorum (Vahl) Gray. Very rare in wet underhang on the bank of the Beldon Burn, 35/94,' 1988, D. E. McC. & G. G. Graham, det. G. G. Graham. * Pterygiopsis lacustris P. M. Jprg. &. R. Sant. On dolerite at the edge of the R.Tees, Upper Teesdale, 35/82, 1993, O. L. Gilbert & V. J. Giavarini. * Pyrenocollema halodytes (Nyl.) R. C. Harris. Sparse to abundant below H.W.M. on the harder parts of Magnesian limestone rocks, concrete piers and limpet shells, wherever these habitats occur along the Durham coast, except those areas where dense algal growth associated with sewage and coal dust pollution has not been cleared from the rocks. First seen at Trow Point, 45/36 and abundant on S. wall of pier below H.W.M. , Roker, 45/45 in coastal survey by D. E. McC. 1997. * P sublitorale (Leight.) R. C. Harris. Similar habitat to the previous species, but not yet found on shells. On hard ridges of Magnesian limestone boulders below H.W.M.. Trow Point, 45/36 1992; also 45/45 & 45/53, 1997. * Pyrenula chlorospila Arnold. On base of old ash in gorge, growing adjacent to Biatorina atropurpurea, Shipley Wood, 45/02, 2000, O. L. Gilbert & D. E. McC. Racodium rupestre Pers. Shaded damp acidic rocks, generally in gorges, 35/83, 35/93, 35/94, 45/02, 45/04 & 45/05. Ramalina farinacea (L.) Ach. The distribution of this species has spread more easterly in recent years as efforts to clean up the air have intensified. Now recorded in at least 19 squares. Lichen flora of County Durham (VC66): Supplement 1 181 R. fastigiata (Pers.) Ach. On old ash at edge of pasture, Westernhopeburn Farm, 35/93, 1988; on old sycamore, Horsley House, also in 35/93, 1997. * Rhizocarpon amphibium (Fr.) Th.Fr. On dolerite at the edge of plunge pool below Cauldron Snout, Upper Teesdale, 35/82, 1993, O. L. Gilbert & V. J. Giavarini. Second British record. * R. hochstetteri sensu qtr. Fryday. Very rare on acid rock in pile of stones on hillside of high moorland, Swinhope Head, 35/83, 1997, conf. A. Fryday. * R. distinctum Th.Fr. Very rare on acidic stones in walls in the W. of the county. On acid stone in wall of churchyard, Hunstanworth, 35/94, 1991, conf. O. L. Gilbert; on dolerite block in wall. Force Garth, 35/82, 1996. R. lavatum (Fr.) Hazsl. Acidic rocks in wall near stream in old mining area, Ladys Rake, 35/83, 1997. R. polycarpum (Hepp) Th.Fr. Now known to be frequent on acid rocks in walls in the highest parts of the W. of the county. Rimularia furvella (Nyl. ex Mudd) Hertel & Rambold Now known to be occasional to frequent on well-lit acidic rocks and capstones in walls in the western uplands; very rarely on tops of gates. Recorded from 9 western squares. * Rinodina sophodes (Ach.) A. Massal. Rare, but locally frequent; concentrated in Upper Weardale, but with a far flung outlyer near Sunderland in 45/35. Generally on the nodes of twigs or on smooth bark. First modern record on ash twigs in gorge woodland, Westernhopeburn Wood, 35/93, 1988; also in 35/83 - 6 sites in total. Sarcosagium campestre (Fr.) Poetsch & Schied. On spoil from old ironworks next to track through woodland, West Wood, Consett, 45/05, 1992, det. O. L. Gilbert; on very short moss on consolidated ground of old colliery site, Hobson Industrial Estate, 45/15, 1996. * Schaereria cinereorufa (Schaer.) Th.Fr. Locally frequent on ledges on walls in the western hill areas. Known from 16 sites in 35/73, 35/83, 35/94 & 45/04. First noted on acidic stones in walls (especially on ledges) in the Upper Derwent Valley, 35/94, late 1980s, conf. O. L. Gilbert. The two entries in the Durham Flora listed as this species were erroneous and have been redetermined as Lecidoma demissum and Umbilicaria torrefacta. * S. fuscocinerea var. sorediata (Houmeau & Cl. Roux) Coppins. On acid boulders in boulder field on exposed hillside, N/W of Burnhope Reservoir, 35/83, 1998. * Scoliciosporum pruinosum (P. James) Vezda. Very rare on sycamore in river valleys. On sycamore bole on wooded fringe of R.Derwent, Ravens Crag, 45/04, 1989, det. B. J. Coppins; on sycamore bole on wooded fringe of river Hisehope Burn, also in 45/04 1994. Sphaerophorus glohosus (Huds.) Vain. On top of acid boulders that are big enough to avoid the worst effects of heather burning, 35/93, 35/94 & 45/04. * Staurothele fissa (Taylor) Zwackh. Very rare. On dolerite at the edge of the R.Tees, Upper Teesdale, 35/82, 1993, O. L. Gilbert & V. J. Giavarini; on harder stone in unnamed syke, Ashgill Head, 35/83, 1997. Steinia geophana (Nyl.) Stein. On spoil from old ironworks at edge of woodland track, West Wood, near Consett, 45/05, 1992, conf. O. L. Gilbert. Stereocaulon nanodes Tuck. On old mine spoil, Ramshaw, 35/94, 1990, G. G. Graham & D. E. McC.; on small stone in shaded recess of pile of rocks from old mine workings, Scaud Hill, 35/83, 1997; abundant on metal-rich rocks on the riverbank (anti-erosion), Ryton Willows, 45/16, 1997. * Strangospora pinicola (A. Massal.) Korb. Occasional on lignum and rare on bark; mostly on rails in hedges or at the edges of woods. Seems able to cope with moderate pollution. First record on bole of half-grown sycamore, car park, Allensford, 45/05, 1992; also in 45/15, 45/16 & 45/43 - 6 sites in all, with 5 on lignum. Thelidium minutulum Korb. Now known from 7 sites, usually on shaded rocks or stones, but is probably much overlooked. On shaded limestone rock at side of steps to lower level of cliff, Roker, 45/45, 1997; on emergent hard pebbles/cobbles in R.Wear, Witton Bridge, 45/13, 1998; also in 45/16 & 45/13. 182 Lichen flora of County Durham (VC66): Supplement 1 * T. zwackhii (Hepp) A. Massal. On limestone in sheltered gorge in the hills, Upper Teesdale, 35/83, 1997, conf. O. L. Gilbert. * Tomasellia gelatinosa (Chevall.) Zahlbr. On hazel twigs near stream, Swinhopeburn Wood, 35/93, 1994, G. G. Graham. Toninia aromatica (Turner ex Sm.) A. Massal. Now known from 9 sites on limestone, widely separated: three sites in Upper Teesdale, 35/83 and the others on coastal Magnesian limestone, 45/36, 45/43, 45/44 & 45/46. * Trapelia corticola Coppins & P. James. Rare on semi-shaded boles of oak in ancient woodland, Great Wood, 45/02, 2000, O. L. Gilbert & D. E. McC., det. O. L. Gilbert. An inconspicuous species, possibly overlooked elsewhere. T. obtegens (Th.Fr.) Hertel. On moorland stone on edge of Waskerly Way, 45/04, 1997, D. E. McC. & C. J. B. Hitch, det. C. J. B. Hitch; on block of coarse-grained acid sandstone in wall, Ireshopeburn, 35/83, 1997. * Trapeliopsis gelatinosa (Florke) Coppins & P. James. Rare on decaying roots and bare peat in the uplands. On decaying roots on bankside underhang, Force Garth, 35/82, 1996, conf. B. J. Coppins; on small recess in bankside, Ladys Rake Mine track, 35/83, 1997; on bare peat on heather moor, near quarry at Doctors Gate, 45/03, 1997. * T. glaucolepidea (Nyl.) G. Schneider. Very rare on bare peat at edge of track in mature lichen heath, Waskerley Way, 45/04, 1997. * T. pseudogranulosa Coppins & P. James. Occasional on roots on banks and mossy tree bases in woodland. Scattered in the western half of the county. First record from decaying Calluna roots on bankside near Beldon Burn, Castleberry Cleugh, 35/94, 1991. * Umbilicaria polyrrhiza (L.) Fr. Very rare on moorland boulder that has managed to escape the worst effects of heather burning. First seen c.1988 on top of large acid boulder on Muggleswick Common, 35/94. Still there in 1997 as three small thalli on top edge of same boulder in a sea of burnt heather and denuded boulders. Refind of a species long thought extinct in the county. U. torrefacta (Lightf.) Schrader. On large boulder on track to quarry, Harthope Hill, 35/83, 1989, G. G. Graham, D. E. McC., B. Humphreys & S. Wharton; on acidic boulder on exposed grassy hillside, near Mathews Stone, 35/83, 1998. Usnea filipendula Stirt. Now known from 5 sites in the Upper Derwent Valley woodlands. The first record was in fact from the Burnhope Burn Wood, near Edmundbyers, 45/05, 1985 and not the Bollihope Burn as previously reported; also in 35/94 & 45/04. * U. wasmuthii Rasanen. Very rare on oak in valley woodland, Nookton Burn, 35/94, 1995, conf. B. J. Coppins. Verrucaria aethiobola Wahlenb. Rare, but status uncertain due to confusion with V. praetermissa. On cobbles on the edge of the R.Tees, Great Wood, 45/02, 1989. The Widdybank record should be safe as the species is known to be frequent above Cow Green. Other records are probably V. praetermissa. * V. amphibia R. Clemente. Sparse on harder parts of Magnesian limestone and concrete piers at the H.W.M. At base of cliff, Blackhall Rocks, on concrete sea-wall at Seaham Harbour and on concrete and hard boulders at the base of piers at South Shields, 45/36 45/43 & 45/44, 1997. * V. caerulea DC. Rare on shaded limestone cliffs and boulders in gorges at 4 sites near the coast and one in Weardale. First record from shaded harder part of Magnesian limestone cliff, Hawthorn Dene, 45/44, 1997; also Castle Eden Dene, 45/43, Easington Dene, 45/44, Tunstall Hills, 45/35 & Westernhopeburn Wood, 35/93, all 1997. * V. funckii (Spreng.) Zahlbr. Rare on hard pebbles and bedrock in upland streams. On dolerite at the edge of the R.Tees, Upper Teesdale, 35/82, 1993, O. L. Gilbert & V. J. Giavarni; on hard pebbles in upland stream, Swinhope Burn, 35/83, 1997; on hard pebbles at edge of R.Tees, Great Wood, 45/02, 1997; on hard shelves in stream. Slit Wood, 35/93, 1997. V. margacea (Wahlenb.) Wahlenb. On semi-inundated sandstone pebbles, Swinhope Burn, 35/83, 1997; on emergent acidic boulder in upland stream, S. of Edmundbyers, 45/04, Book Reviews 183 1997; on semi-submerged acid boulder in upland stream, Wellhope Beck, 35/84, 1997. V. maura Wahlenb. Sparse but in several sites down the coast where the Magnesian limestone rocks are hard enough and cleared of the debris of pollution. Found in 7 sites in 1997. First seen on harder vertical faces of rough cliffs at Trow Point, 45/36; extending to several metres up cliff face above FI.W.M. at Blackhall Rocks, 45/43. • V. murina Leight. Very rare on limestone boulder, adjacent to undescribed Acarospora sp., on shaded stream bank, Westernhopeburn Wood, 35/93, 1995, conf. O. L. Gilbert. * V praetermissa (Trevis.) Anzi. Widespread and frequent in unpolluted streams. First record uncertain due to confusion with V. aethiohola. V. striatula Wahlenb. Sparsely scattered down the coast and found in 10 sites in similar situations to other maritime Verrucaria species, 1997. References Graham, G. G. (1988) The Flora and Vegetation of County Durham. Durham Flora Committee & Durham County Conservation Trust, Durham. Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. W. and Moore, D. M., ed. (1992) The Lichen Flora of Great Britain and Ireland. Natural History Museum, London. Purvis, O. W., Coppins, B. J. and James, P. W. (1993) Checklist of lichens of Great Britain and Ireland. British Lichen Society Bulletin 72 (Supplement): 1-75. Gilbert, O. L. & Giavarini, V. J. (1997) The lichen vegetation of acid watercourses in England. Lichenologist 29: 347-367. BOOK REVIEWS Lichens. An Illustrated Guide to the British and Irish Species by Frank S. Dobson. Pp. 431, inch numerous line drawings, maps and colour plates. Richmond Publishing, Slough. 2000. £45.00 hardback, £30.00 paperback. This fourth edition of Britain’s most popular guide to the identification of its lichens is a considerable improvement on its predecessors in terms of its scope and presentation. In all, about 700 species (approx, half the British flora) are treated, the texts to the majority of which are supported by coloured plates and distribution maps. Keys to determine all lichens covered to specific level are provided and where necessary line drawings of diagnostic microscopic features are illustrated. The work is complemented by useful introductory matter, including information on microscopical analyses, chemical tests, photobionts and air pollution zones, and a glossary. There should be no excuse now for the inability of the general natural historian or ecologist to identify such organisms, at least the commoner ones, which form key components of many ecosytems. MRDS The Flora of Dorset by Humphry Bowen. Pp. viii + 373, inch numerous line drawing, maps & colour plates. Pisces Publications. 2000. £45.00 hardback + £5.00 postage & packing from Naturebureau, 36 Kingfisher Court, Hambridge Road, Newbury, Berks. RG14 5SJ. Although the author, a friend and colleague of this reviewer, saw this volume through to print, his recent untimely death precludes his enjoyment of the many complimentary reviews of this splendid work. The scope of the Flora is ambitious in its review of the vascular plants, bryophytes, lichens, fungi and algae, but the author’s undoubted knowledge of all these groups provides a balanced coverage, lacking in some Floras where non-vascular plants are often treated as an afterthought. Although the author had been interested in the Dorset flora since 1934, his main work was undertaken after he retired to his holiday home there in 1988; during the last decade, for example, he assembled 220,000 flowering-plant records. However, previous work on the county’s flora is not forgotten, and the review (by David Allen) of some earlier workers is first-rate. 184 Book Reviews In all, over 2000 species of vascular plants (more than 900 of which have distribution maps at the 2 km x 2 km level), 324 mosses, 101 liverworts, 652 lichens, 1496 fungi and c.400 algae are treated within the main body of the text. Alien plants are also given full coverage, including details of some ornamental plants in cultivation at selected localities. Introductory chapters by several authors, on physical features, vegetation and effects of man (including archaeobotany by Pat Hinton) are also provided. This volume is a fitting tribute to a distinguished chemist who through meticulous recording proved himself to be one of Britain’s most versatile botanists. MRDS The Flora of the Bristol Region by Ian P. Green, Rupert J. Biggins, Clare Kitchen and Mark A. R. Kitchen, and edited by Sarah L. Myles. Pp. vi + 276, inch numerous line drawings, maps and colour plates. Pisces Publications. 2000. £27.50 hardback + £4.00 postage & packing from Naturebureau, 36 Kingfisher Court, Hambridge Road, Newbury, Berks. RG14 5SJ. This Flora, the culmination of a 15-year project, details the occurrence of more than 1600 vascular plants, 1008 of them on a 1 km x 1 km basis, within the area formerly known as the County of Avon. Due to the detailed nature of the maps it is possible to gauge the pattern, ecology and status of the species, but template maps of a range of topographical features, etc. would have been useful for comparative purposes; however, useful introductory chapters on the Physical Character and the Habitats of the Region are provided. Of particular interest is the chapter on 20 Sites of Botanical Interest, including the famous Avon Gorge, which are analysed in detail. Other chapters are devoted to the History of the Avon Flora Project and Botanists in the Bristol Region, as well as Lists of Native and Non-native Species not recorded in the area investigated in recent years, and a Gazetteer. This work is a model of its kind in that the comprehensiveness of the botanical survey will provide an invaluable benchmark against which to monitor any future changes. MRDS The Aurelian Legacy. British Butterflies and their Collectors by Michael A. Salmon, with additional material by Peter Marren and Basil Harley. Pp. 432, inch 162 b/w and 42 colour plates. Harley Books, Great Horkesley, Colchester. 2000. £30.00 hardback. This magnificently produced book is a fitting tribute to dedicated lepidopterists past and present. The author is clearly one of a very long line of enthusiasts, some more appropriately labelled as eccentrics, stretching back over 300 years. By the early 18th century entomology had becomes a social, even fashionable, pursuit. One such group, meeting at the Swan Tavern in London at that time, was dubbed “The Aurelians” (aurelius referring to the gilt-decorated chrysalids of certain nymphalid moths) - hence the title of this book. Collecting reached its zenith in the 19th century, when the esoteric activities of Victorian entomologists were often little more than trophy hunting. Although environmental changes also impacted dramatically on our butterfly fauna, such over- zealous collecting undoubtedly led to the decline, indeed disappearance, of many species. The main part of this work is devoted to succinct but fascinating biographies of 101 notable lepidopterists. Other sections deal with the history of butterfly collecting in Britain, the techniques and equipment used in their collection, and the development of knowledge of 35 butterfly species recorded at some time in Britain, such as the Large Blue, the Arran Brown and the Scarce Copper. Overall, the volume is a visual delight while the text is both informative and a joy to read. Thoroughly recommended, not only to entomologists but also to those with wider natural history interests. MRDS THE COLLYRINAE {HYMENOPTERA, ICHNE UM ON I DAE ) OF YORKSHIRE. 185 W. A. ELY Rotherham Biological Records Centre, Norfolk House, Walker Place, Rotherham S65 IAS This small subfamily has two British species whose adults are quite distinctive in appearance (Fig. 1) and which can be identified by reference to Fitton (1984). Kerrich (1936) was the first to recognise that two species of Collyria occurred in Britain and Fitton’s key introduces a new character for separating them. The “commoner” one is parasitic on the larvae of stem-mining sawflies (Hymenoptera: Cephidae), especially the Wheat-stem Sawfly (Cephus pygmaeus), and the other is likely to have a similar host spectrum, though that has not yet been proven. Not surprisingly, the adult ichneumon has a similar slender shape to that of its host. FIGURE 1. Collyria trichophthalma (Thomson, 1877) left wings omitted. All the Yorkshire records of Collyria have previously been reported as C. coxata (Villers) (Hincks, 1946; Hincks, 1951 and Hincks, 1953) and were collected between 1936 and 1947 (Figs. 2 and 3). Most of the insects on which these records are based have been seen by G. J. Kerrich, J. F. Perkins or myself in the Manchester Museum, Leeds Museum or Don Smith collections and all are correctly identified; there remain a couple of records for which I have not been able to trace any specimens. Fitton states that this insect is Naturalist 126 (2001) 186 The Collyrinae (Hymenoptera, Ichneumonidae) of Yorkshire Yorkshire Recorded distribution of Collyria coxata (Villers, 1789) (O) and Collyria trichophthalma (Thomson, 1877) (•) in Yorkshire. “common and has been collected as far north as North Yorkshire”. However, no specimens of Collyria coxata have been seen from Yorkshire since 1947. Fitton records the flight period as June and July, which is true for the Yorkshire records. The second species is Collyria trichophthalma (Thomson), which Fitton states “is rare ... in southern England ... as far north as Cambridgeshire”. However, I have seen Yorkshire specimens collected from fifteen 10 km squares in 4 vice counties between 1944 and 1991 in the collections of the Yorkshire Museum and Doncaster Museum and Art Gallery as well as freshly-collected material (e.g. Ely, 1986) (Figs. 2 and 3). Fitton records the flight period as May and June but there are Yorkshire records from July and the very first are from August 1944 and April 1945! Both were collected from Malton Road, York, by J. H. Elliott. ■ B r " I T— I 1 T 1 -1- 1 1 1930 1940 1950 1960 1970 1980 1990 2000 FIGURE 3. Recorded occurrence of Collyria coxata (Villers, 1789) (bottom line) and Collyria trichophthalma (Thomson, 1877) (upper line) in Yorkshire. The Collyrinae (Hymenoptera, Ichneumonidae) of Yorkshire 187 The Yorkshire records of these two insects fall into two patterns, a temporal one and a spatial one. Although there is an overlap, the records of C. coxata generally predate those of C. trichophthalma and the latter has been the only Collyria recorded in Yorkshire for half a century. This implies that a change in status has occurred and we may have observed the final stages of the replacement of C. coxata by C. trichophthalma. This change may not have occurred in the south of England, though it would be interesting to know if Fitton’s review, which is based on museum specimens and is necessarily historical, reflects the current position there. (I have collected C. trichophthalma in the south of England but have never taken C. coxata .) The cause of this change in status is unclear and the picture is made more complex than normal because these are parasitic insects. The change may be the result of an effect on its host or on itself or on both together. The second pattern is again not absolutely clear-cut but most records of C. coxata are coastal or estuarine while none of the C. trichophthalma records can be so described. The YNU has been as active in the coastal fringe of Yorkshire since 1950 as it was before and the flight periods of both species coincide with the months of the Union’s Vice County field meetings, so collector bias seems to be an unlikely explanation for this difference. It may reflect a differential host ditribution, with the effects of the coast exerting an influence on the host instead of, or as well as, on the ichneumon. There are no records for either species from the uplands, which is what would be expected of insects which are at or close to the northern limits of their British ranges. There are other anomalies in the distribution of the two Collyria species and I would like to draw attention to two inexplicable gaps. Two of the most diligent collectors of insects in Yorkshire were William Fordham, operating in the Allerthorpe area from 1920 to 1935, and John Wood, , who added so much to our knowledge of the entomofauna of Keighley between 1928 and 1950. Neither naturalist studied the Parasitica but both collected them extensively and passed specimens on to others, yet there are no records of Collyria from either of their principal collecting grounds. It is possible that Keighley is too far into the uplands (and Wood did collect C. coxata on one visit further east) but Fordham lived in the area which has provided many examples of C. trichophthalma recently. His failure to find Collyria is particularly interesting. Acknowledgements I am grateful to all those collectors who have passed specimens to me or deposited them in a museum collection: Austen Brackenbury, Harry Britten jnr, J. H. Elliott, Simon Hayhow, Douglas Hincks, Alan Eazenby, Joyce Payne, Peter Skidmore, Don Smith, Derek Whiteley and John Wood. I also wish to express my gratitude to Colin Johnson, Peter Skidmore, Martin Limbert, Paul Howard, Stuart Ogilvie, Adrian Norris and Mike Fitton for providing access to the entomology collections at Manchester Museum, Doncaster Museum and Art Gallery, Yorkshire Museum, Leeds Museum and the Natural History Museum. References Ely, W. A. (1986) Entomology, in Yorkshire Naturalists’ Union Excursions in 1985: Thrybergh (VC63), 6 & 7 July. Naturalist 111: 106-107. Fitton, M. G. (1984) A Review of the British Collyrinae, Eucerotinae, Stilbopinae and Neorhacodinae (Hymenoptera: Ichneumonidae) Ent. Gaz 35: 185-195. Hincks, W. D. (1946) Entomology, in Yorkshire Naturalists’ Union Excursions in 1946: Spurn, June 8th- 10th. Naturalist 71: 159-160. Hincks, W. D. (1951) Hymenoptera, in The Yorkshire Naturalists’ Union: Eighty-ninth Annual Report. Naturalist 76: 26-29. Hincks, W. D. (1953) X. Hymenoptera excluding Aculeata, in The Entomology of Spurn Peninsula. Naturalist 78: 129-139. Kerrich, G. J. (1936) Systematic note on two species of Collyria Schiodte (Hym., Ichneumonidae). J. Soc. Br. Ent. I: 124-125. 188 BOOK REVIEWS The Changing Wildlife of Great Britain and Ireland edited by David L. Hawksworth. Pp. xvi + 454, incl. numerous tables, graphs & maps. Taylor & Francis, London. 2001. £150.00 hardback. This volume is a milestone in our understanding of the changes to and current status of our country’s fauna and flora. Like its predecessor, The Changing Flora and Fauna of Britain published in 1974, it provides a reliable baseline for conservationists since it documents losses and declines, and additions and increases, to a wide range of organisms over the past 25 years. Each of the following groups has received specialist treatment: Flowering plants, Ferns and allied plants, Mosses, liverworts and hornworts. Larger fungi, Microscopic fungi, Lichens, Terrestrial and freshwater eukaryotic algae, Cyanobacteria, Diatoms, Viruses, Protozoa, Freshwater invertebrates, Nematodes, Mites and ticks, Flies, True bugs, leaf- and plant-hoppers, and their allies. Butterflies and moths, Grasshoppers, crickets and allied insects, Dragonflies and damselflies, Land slugs and snails, Birds, Mammals, and Fishes - in all, a remarkable survey that provides a fairly comprehensive database. These 23 chapters are supported by three valuable contributions on ‘Fifty years of statutory nature conservaion in Great Britain’, ‘Tracking future trends: the Biodiversity Information Network’ and ‘Prospects for the next 25 years’, the latter making the point that “monitoring the status of the perhaps 100,000 species in great Britain is unlikely to become realistic in the foreseeable future” and that “overall, the future looks bleaker than the past” - clearly the prognosis is one of on-going concern. This book is an absolute must for the serious naturalist, but at this exorbitant price such readers will only be able to consult library copies. MRDS The Correspondence of Charles Darwin: Volume 12, 1864 edited by Frederick Burkhardt and others. Pp. xl + 694, incl. several line drawings, plus 13 pp. of b/w plates. Cambridge University Press. 2001. £55.00 hardback. The latest volume in this monumental series, reviewed extensively in The Naturalist over the past 16 years, charts a single but highly productive year: despite his continued ill- health, Darwin reveals through his correspondence the impact of his theory of evolution on the scientific world and the public in general. His researches in support of his theory and in other biological fields are also evidenced from his letters; of particular interest are those relating to his work on cross-pollination and climbing plants, the latter resulting in the publication of his treatise On the movement and habits of climbing plants the following year. In 1 864 he had also commenced work on what was to be one of his most influential publications in support of his theories, namely The variation of animals and plants under domestication (published in 1868). This volume contains important correspondence with Jean Louis Agassiz, George Bentham, Hermann Criiger, Hugh Falconer, Asa Gray, Joseph Hooker, Thomas Huxley, William Jenner, John Lubbock, Daniel Oliver, John Scott, Richard Spruce and Alfred Russel Wallace and many other notables, as well as considerable information relating to Kew, the Linnean Society and The Royal Society (including the award of its highest honour, the Copley Medal, to Darwin in 1864, detailed in Appendix IV). As would be expected now of this definitive work, the letters are supported by a very considerable textual apparatus, as well as bibliographic and biographic material, and a comprehensive index. MRDS 189 FUNGUS-GNATS FOUND NEW TO YORKSHIRE IN 2000 JOHN COLD WELL 16 Railway Cottages, Dodworth, Barnsley S75 3JJ The five families of Nematocerous Diptera popularly referred to as fungus-gnats are essentially woodland insects, which, as the name implies, develop in various types of fungi. Some, like Keroplatus testaccus , are characteristic of old forest or parkland around rotting logs whilst a few may be more readily found by sweeping tree foliage. The majority, however, are denizens of damp, shady woodland floors and streamsides with peaks of abundance in early summer and especially autumn. A number of fungus-gnats are reported for the county for the first time as a result of recording carried out by the author in the Barnsley area and of specimens collected by Roy Crossley in various parts of Yorkshire during the year 2000. The following list consists of species ‘reported’ for the first time, although it is probable that some will already have been found in the county by visiting dipterists in the past and the author would be keen to hear of any such additions. Keroplatus testaceus Dalman Worsbrough Country Park 1 $ swept from a rotting log in willow carr, 3.9.2000. A large yellowish gnat found in southern England with odd records from Cumbria and Scotland. Neoplatyura modesta Winnertz Skipwith Common. 1 9 swept by R. Crossley, 22.8.2000. A predominantly southern species although known from Cheshire and Lancashire. Macrocera fastuosa Loew Chafer Wood, 7.8.2000. A Red Data Book 1 species, this cf , found by R. Crossley, constitutes only the third British record. Boletina dispecta Dziedzicki Houghton Common, South, Yorkshire. 1 cf swept from bracken in shade under Quercus, 8.9.2000. A scarce though widespread fly nationally. Boletina edwardsi Chandler Sieve Dale Fen. lcf 11.9.2000 by R. Crossley. Very local in the north though proving to be generally widespread. Boletina rejecta Edwards Clough Wood, Gunthwaite. lcf , 9.9.2000. Both this species and the very similar B. dispecta are proving to be more frequent than once thought (Chandler, pers. comm.). Boletina trispinosa Edwards Found at Nabs Wood, Silkstone and by the wooded edge of Ingbirchworth Reservoir. This species will probably prove to be fairly common. Cordyla fasciata Meigen A local gnat, found at Houghton Common and Nabs Wood. Presumably overlooked by previous recorders in Yorkshire. Synplasta ingeniosa (Kidd) Clough Wood, Gunthwaite. 2cf cf , 29.8.2000. Found in broad-leaved woodland in southern England, the larvae probably developing in soft fungi. Scarce, but beginning to be found more widely. Anaclileia dispar (Winnertz) Fox Clough, Langsett. lcf, 30.5.2000. A small, dark sciarid-like gnat which could overlooked. Scarce nationally. Naturalist 126 (2001) 190 Book Review Neuratelia nigricornis Edwards Uncommon, with scattered records throughout Great Britain, this species was found near the Little Don Valley in mid-July amd also in a similarly shady streamside moorland valley habitat at Broadhead Clough a few days later by R. Crossley. Phthinia mira (Ostroverkhova) Clough Wood. 1 cT , 20.8.2000. The one record of P. humeralis in Yorkshire (excluding one from Malham Tarn based on an unreliable find in 1980) may well have been this species as the humeralis group was not clarified until Chandler (1987). Sciophila fenestella Curtis Single cfcT found in willow carr by Elsecar Reservoir in July and by streamside vegetation near Wortley, South Yorkshire in September. Mainly southern but present in north-west England and may well have a wider distribution than currently believed. Two further species have been recorded during 2000, which, although stated as having occurred in ‘Yorks’ in Hutson et al. (1980), only appear in the YNU index on that account and without locality details in the first case and only otherwise on the aforementioned Malham list in the second. Bolitophila glabrata Loew 1 cf , Dearne Valley Park 18.10.2000. Rather uncommon in Great Britain. My corny a fimbriata (Meigen) lcf, Gypsy Marsh, near Wombwell, 16.8.2000. Widespread nationally. Acknowledgement I would like to thank Peter Chandler for checking a number of specimens and also Roy Crossley for collecting and making available material for study, knowing that I had become interested in this group of flies. References Chandler, P. J. (1987) Notes on British fungus gnats of the smaller Families and sub-Families (Diptera, Mycetophiloidea). Proc. Trans. Br. Ent. Nat. Hist. Soc. 20: 105-118. Hutson, A. M., Ackland, D. M. and Kidd, L. N. (1980) Mycetophilidae (Bolitophilinae, Ditomyiinae, Diadocidiinae, Keroplatinae, Sciophilinae and Manotinae). Handb. Ident. Brit. Insects 9(3): 1-111. BOOK REVIEW Riverside Journey: A Portrait of the Derwent by Ashley Bryant. Pp. 90, half paintings, half text. The Lutterworth Press, Cambridge, 2001. £30.00 hardback. Riverside Journey is a portrait of the author/artist’s three year journey down the Derbyshire Derwent from its source to where it joins the Trent. The volume contains over 80 paintings and sketches in a variety of media which chart the river in all its moods over these three years. The accompanying text details the author’s memories, for example of walks in his home county as a child, and information about the activities of naturalists, historians and others in this beautiful county. Bryant captures the mood of the river in his paintings, which vary from evocations of woodland and moor to the fun of raft races, riverside gardens and walks. The majesty of mills and stately homes contrasts with more tranquil scenes in winter or at night. The book is a pleasure to the eye, of interest to the natural historian and of especial pleasure to those who have lived in or visited the areas of Derbyshire which are covered. JLA 191 A TAWNY OWL SEEN TAKING A FYLING BAT IN BRADFORD DIANE GREGORY West Yorkshire Bat Group, 16 The Drive, Alwoodley, Leeds LS17 7QA The role of owls as the major natural predator of bats has been discussed for many years. The literature was reviewed by Speakman (1991). Evidence from owl pellets has left no doubt that owls eat bats. Speakman (1991) concluded that bats account for less than 1% of the prey items taken by owls in the British Isles, but even so, this may be as much as 10% of annual bat mortality. Julian and Altringham (1994) analysed a total of 741 owl pellets, collected on a monthly basis, from a tawny owl’s territory in a small wood adjacent to a sewage works on the outskirts of Leeds. They found that bats were regularly consumed, probably by a single owl, and reported that they had encountered no other case where a British owl had predated upon bats to so marked an extent. On the evening of Monday, 23 July 2001, in the garden of a house north-west of Four Lane Ends, Bradford, a tawny owl was observed using a tree as an observation post and intently watching Pipistrellus pipistrellus bats emerging from their roost. The house owners commented that the owl and its mate were regular visitors to the garden at dusk. Bats were both emerging from the roost and feeding in the surrounding garden. The owl launched itself, manoeuvred above a flying bat and in a movement that involved a turn through 180°, took the bat and flew off. The sequence of events was too fast to enable accurate observations of how the bat was taken. Within a relatively short time the owl was back, and again watched the bats as they continued to emerge. John Altringham (pers. comm.) has seen a similar occurrence at Malham. On this occasion the bat was taken during the dawn swarm, before re-entering the roost. Despite the fact that only a single observation was made, it is probable that this particular owl takes regular meals of pipistrelle bats from this Bradford roost. There is little doubt that the numbers of bats from the roost will be severely depleted. It seems that where owls encounter bat roosts within their territories, they learn to exploit bats as a food source, both at dusk and dawn, the periods of maximum bat density. It is likely that such encounters are purely adventitious, and that having recognised the availability of an unusual food supply, owls adopt strategies that enable them to utilise it. Acknowledgements Thanks are due to the Soni family who allowed me to make this observation, to John Drewett who commented upon the unusual nature of this event, and to Gordon Richardson who encouraged me to describe it. References Julian, S. and Altringham, J. D. (1994) Bat predation by a Tawny Owl. Naturalist 119: 49-56. Speakman, J. R. (1991) The impact of predation by birds on bat populations in the British Isles. Mammal Rev. 21: 123-142. BOOK REVIEWS Snakes by Peter Stafford Pp. 112 with 114 colour photographs and 12 drawings and diagrams. The Natural History Museum (Life Series), London. 2000. £9.95 softback. This book by the editor of the Bulletin of the British Herpetological Society highlights the variety and complexity of an often misunderstood group of vertebrates which worldwide includes almost 3,000 living species. The evolution, biology and natural history of snakes is fascinatingly and comprehensively dealt with and the quality of the numerous colour photographs is justification enough to acquire this book. Naturalist 126 (2001) 192 A Tawny Owl seen taking a Flying Bat in Bradford The Natural History Museum’s Life Series form an authoritative, attractive and highly accessible introduction to a range of natural history subjects written by some of the world’s leading specialists. A standard and particularly useful feature is a good glossary, up to date bibliography, contact with relevant specialist societies and of great value is the range of addresses for internet resources. CAH Deep Ocean by Tony Rice. Pp. 96 with over 80 colour photographs and 14 drawings and diagrams. The Natural History Museum (Life Series), London. 2000. £9.95 softback. A mere 150 years ago the geography of deep ocean areas was a total mystery, biologists judging it inconceivable that they could support life. From 1872 to 1876 tantalising glimpses into the ocean deeps were revealed by the celebrated oceanographic expeditions of HMS Challenger. However, in the past couple of decades a range of new technologies have shone an enquiring light into the abyss of ‘inner space’ producing a veritable deluge of new discoveries. As with the opening up of the world’s continents to botanists and zoologists during the 18th century, it is our privilege to live at a time when technological developments, in this case deep sea submersibles, cameras, videos, satellite imaging and computers, are not only revealing a 21st century ‘cabinet of curiosities’ but pointing to new pathways of evolution and revealing totally new ecosystems fuelled by geological sulphides and chemosynthetic bacteria rather than sunlight and photosynthetic plants. We are shown the ability of countless vertebrate and invertebrate organisms to signal, deter, attract and lure by a dazzling array of bio-luminescent techniques. We see scavengers at work seven miles down on the abyssal plain. There are giant worms, monster clams, heat seeking shrimps and creatures surviving temperature gradients ranging from freezing point to that of molten lead. Whether by coincidence or good planning, the issuing of Deep Ocean in the Natural History Museum’s ‘Life’ series coincides with the screening of the BBC TVs highly successful marine wildlife spectacular The Blue Planet. In covering the same subject areas as the film, even illustrating the same points with the same examples and images, one could be excused for believing this book provided the blueprint for the film’s fabulous and revealing deep sea episode. The author, Dr Tony Rice is a respected authority in the world of oceanography, having worked as a marine biologist at the Natural History Museum and at the NERC Institute of Oceanographic Sciences where for 26 years he led the deep-sea benthic team. Certainly Dr Rice’s fascinating and beautifully illustrated book forms an excellent companion volume to the BBC’s film and would form an ideal gift for any aspiring oceanographer or marine biologist. CAH Earth and You. Tales of the Environment by Charles Officer and Jake Page. Pp. viii + 254, inch line drawings & b/w plates. University Press of New England, Hanover. 2000. $25.00 hardback. A thoughtful appraisal of our planet’s environmental health which, while it addresses such major problems as global warming, population growth, resource depletion, species decimation and chemical pollution, is nevertheless optimistic in its approach. The authors show how more imaginative approaches to solving these problems could be pursued through national and international initiatives, moral and religious leadership, and research and development. A refreshingly positive treatment of some very important and sensitive issues, rather than the pessimistic or indeed apocalyptic overview of all too many environmentalists and conservationists. MRDS 193 Entomological Report: Diptera (Tipuloidea and Empidoidea) ENTOMOLOGICAL REPORT: DIPTERA (TIPULOIDEA AND EMPIDOIDEA) ROY CROSSLEY Introduction An outstanding feature of this Report are the additions to the County’s diptera list made by Andrew Godfrey as a result of his splendid work in rearing specimens from exudations and the debris of rot-holes sampled from Horse-Chestnut (. Aesculus hippocastanum) and Sycamore {Acer pseudoplatanus). Of particular note are the Systenus species which are rarely encountered except by rearing them; in twenty years of collecting I have taken only one specimen with the sweep-net. Mr Godfrey’s work suggests that these enigmatic flies are probably more widespread and numerous in Yorkshire than had hitherto been thought. In addition to Mr Godfrey the following dipterists have submitted records, the more significant of which are included in this Report: J. D. Coldwell, W. A. Ely, and A. E. Stubbs, to all of whom I express my thanks; unattributed recent records are those of the writer. New County records are indicated by t, and Vice-County records by * . National rarity classifications which follow, where appropriate, immediately after the species names, are those provisionally recommended by Falk (1991) for Tipuloidea, and by Falk and Crossley {in prep.) for Empidoidea. The systematic order of the list, and nomenclature, follow Chandler (1998). I am grateful, as ever, to Mr J. H. Cole and Mr A. E. Stubbs for much help and advice. Tipulidae *Tipula (s.g. Pterelachisus) pseudovariipennis Czizek Nb. (61) Barmby Moor, 1$ 29/5/99 {leg. & det. R. C., teste J. H. Cole). The only previously reported occurrence of this species in Yorkshire is from High Batts, Ripon (64), April 1999 {Nat. 125 p. 137). Pediciidae Dicranota (s.g. Rhaphidolabis) exclusa (Walker) (63) Broadhead Clough, 15/5/00. The first Yorkshire record was from Malham Tarn (64), 1956, and then Midhope Reservoir (63), 1999 {Nat. 125 p. 137). Limoniidae *Gonomyia (s.g. Prolipophleps) abbreviata Lw. RDB3 (62) Forge Valley Woods NNR, lcf 26/7/99. Apart from a 1927 report from Austwick (64), the only previous Yorkshire records are from the Doncaster area (63) in the mid-1980s (Conisborough, Sandall Beat and Pot Ridings Wood). Hoplolabis (s.g. Parilisia) vicina (Tonnoir) (62) Seivedale Fen, 1 1/9/00. There is a record from Mulgrave Woods (62), in 1937 by the late F. W. Edwards of the British Museum (Nat. Hist.) (now the Natural History Museum), where the specimen(s) was deposited. The only other Yorkshire record is from Birk Crag Wood, Harrogate (64), in 1996 {Nat. 123 p. 121). Ormosia (s.g. Ormosia s.s.) pseudosimilis (Lundstrom) (*62) Coulton Mill Fen, 5/7/00; (63) Broadhead Clough, 24/8/00. There are a further ten widely scattered records from the west and south of Yorkshire, half of which are fifty or more years old. Scleroprocta sororcula (Zett.) Nb. (62) Chafer Wood, 22/5/00. (63) Broadhead Clough, 12/6/98 {leg. & det. R. C., teste J. H. Cole). The only previous Yorkshire records are Bilsdale (62), 1921; Colsterdale (65), 1981; Little Don Valley (63), 1997. Eloeophila trimaculata (Zett.) Nb. (63) Broadhead Clough, 15/5/00. Apparently scarce in Yorkshire, with only four previous widely scattered records, all post- 1976 {Nat. 124 p. 84). Naturalist 126 (2001) 194 Entomological Report: Diptera (Tipuloidea and Empidoidea) Hexatoma (s.g. Hexatoma s.s.) fuscipennis (Curt.) (65) Great Langton, 17/7/97 A.G., swept ex river shingle. This appears to be a very localised species in Yorkshire with only one post-1953 record (Nosterfield gravel pit [65], 1998) (Nat. 124 p. 84). *Limnophila (s.g. Limnophila s.s.) pictipennis Mg. RBD2 (65) Pond near Wath (Ripon), -/9/00, A.E.S. The only previous Yorkshire record is from Levisham (62) in 1895. t Pilaria decolor (Zett.) (62) Chafer Wood, 11/7/00. Treated as a variety of P. discicollis (Mg.) by Edwards (1938), P. decolor has only recently been added to the British List (Stubbs, 1997). P discicollis is common and widespread, and it is probable that examples of P decolor have been incorrectly named as discicollis in the past. Atypophthalmus inustus (Mg.) Nb. (*62) Chafer Wood, 11/7/00. (63) Worsborough Country Park, 23/7/00 J.D.C. There are only two previous Yorkshire records for this species: Rushy Moor Wood (63) 1982, and Acklam (61) 1999. It occurs in damp woodland, where the larvae breed in fungi (Falk, 1991). *Dicranomyia (s.g. Dicranomyia s.s.) sera (Walker) (61) Welwick, 15/8/00. A coastal marsh species; the only other Yorkshire record is from Blacktoft Sands (63), 1976. *D. (s.g. Idiopyga) nigristigma Nielsen (62) Ashberry, 1 UGhP s t l> 1 I rJ IS i z • 1 a ,/ *S EDBE -A’ RGH V C6J K 1 \ VC 61 •4 ■» I jj> X •v — ^ < • r CARE O tc o IGH % / ( e l\_ < V -ol • T' ■v-V c t , | t r'" A V ■ y rC6 (I / > \ 1 \ /' ) i BR tDFO ID eed: i '*w’v HULL V if c & r t < '"'--A c vj 1 \ > L ) /' VC 63 c * -- _ Vice Cc bounda )unty rv ... V NJr’ T y c ; A Boundaries < -- areas lost tc of o or gaine d fro '“j hi " = LD ( U-J rJ V „ other Counti 1 EL es UT 1 d 1 / 0 10 20 30 Km 0 10 20 Miles FIGURE 1 Tetrad records of B. selene Watsonian Yorkshire. Small dots represent 1 record, larger dots represent 2-9 records, the largest dots represent 10+ records. In this study, a site refers to where the imago was recorded close to its food plants. In an exceptional case, the Gisburn Forest site, this designation was based on the sighting of a single adult (Appendix 1). It is very easy to overlook the species at any one site as it may exist in very small numbers, which are on the wing for a short period and only flies in sunshine; a scarce commodity in West Yorkshire. Under-recording in areas remote from Status of the Small Pearl-bordered Fritillary in Western Yorkshire 5 population centres is also a common bias of recording, a problem discussed in detail by Dennis et al. (1999). The Life Cycle A description of the species and its life cycle can be found in Emmett and Heath (1989), South (1962) and Thomas and Lewington (1991) but the following summarises the situation. Eggs Eggs are of a rounded conical shape, with 18-20 ribs. Once thought to be discharged in flight in the vicinity of its foodplant, Viola spp. (Emmett & Heath 1989), it is, however, more likely that they are deposited directly on to the violets or vegetation near them (Asher et al. 2001). They are sticky and adhere to any material they encounter. Their colour initially is greenish, then yellowish and shortly before hatching, greyish, the blackish head capsule of the larvae being visible near the concave tip. Hatching takes place within a fortnight. Larva On emerging, the caterpillar eats most of its eggshell. Initially it is of a pale olive colour with brownish warts from each of which a long and jointed bristle emerges. Its head is shining black with a notch on the crown. Small larvae feed until they have moulted to the fourth instar when they enter diapause around the end of September. By this time many of the violet leaves are yellowing and senescent. According to Emmett and Heath (1989), caterpillars overwinter beneath leaves or in ground detritus, but this probably only applies to woodland sites. Sites for diapause of moorland mire populations are not known but are probably within Juncus or grass tussocks. Diapause lasts from October to March. With rising temperatures the larvae commence feeding in April. The velvety full-grown fifth instar larva is about 2 cm long, blackish with a brownish line along the middle of the back and a pinkish stripe which runs along the lower sides of the body just above the legs. Its spines are ochreous, tinged with pink and bear pointed black bristles. The upper spines are stouter than the rest and the single pair on the first segment is more than twice the length of the others and passes forward over the caterpillar’s head, giving the appearance of a pair of horns. The second and third segments each have four spines which are rather finer than those on the rest of its body where they are arranged in six rows. Larvae appear to be less reliant on solar heat for their well-being than B. euphrosyne and do not bask in the sun as do many fritillary larvae. Indeed they are secretive, hiding in the shadier parts of the vegetation and only emerging for feeding forays. Pupation takes place around the end of May in Yorkshire, but is much earlier in milder parts of England and Wales. Pupa The chrysalis is suspended from a silk pad by its cremaster, low down on a plant stem or in the detritus. It is brown, with the wing cases being more ochreous and marked with black near the edges. On the thorax there is a black V-mark and metallic silver spots on each side, one silvery spot on the head and other metallic spots near the thorax. It remains at this stage for about three weeks. Imago In the west of Yorkshire emergence usually starts around the second week of June but can be up to two weeks later in some years. Adults are seen on the wing for about a month and have been sighted between the second week in June and the last week of July (47 days), a very similar period to the 41 days on the North York Moor sites (Clough 1998, 1999b, 2000b; Parkinson 1997; Sutton & Beaumont 1989). The maximum adult life span is unknown but is unlikely to be more than about two weeks. In VC64 the times when peak numbers of adults have been observed vary from year to year between mid-June and mid- July. Behaviour of the sexes is markedly different. Males actively patrol the breeding sites whereas females usually make short flights or remain at rest low down in the herbage. 6 Status of the Small Pearl-bordered Fritillary in Western Yorkshire When approached, they often initially reject the attentions of the male by moving deeper into the vegetation. These rejection rituals are described more fully by Emmett and Heath (1989). Butterflies are torpid on cool days but when temperatures are low and there is strong sun they may spend considerable periods dorsally basking. If the air temperature is lower than c.17°C they remain at rest in the herbage for long periods and are difficult to locate. Their habit of roosting with closed wings on the flowers of Juncus spp. has often been noted (Walsh 1956; Emmett & Heath 1989). It is probable that this energy-conserving strategy is fundamental to their survival during inclement weather. Walsh (1956) considered their underside pattern to be cryptic, closely resembing the Juncus flower- heads. However, Skinner (1984) mentions another member of the Juncaceae, Wood-rush, (. Luzula sp.), as a nectar source for the moorland moth, Haworth’s Minor ( Celaena haworthii ) and Mounsey (2001) records seeing moths collecting in numbers on Juncus effusus, finding the flower-heads had ‘quite a sweet taste’. Is it possible that Juncus itself could be a nectar source? In this study adults were rarely seen on known nectar flowers. Although Cirsium palustre was observed as the preferred nectar plant, a single individual was seen on Erica cinerea, while all Brian Shorrock’s (pers. comm.) sightings of the butterfly at nectar were on Marsh Thistle. It is also reported that Cardamine pratensis , Lychnis flos-cuculi and, on sites with improved pasture adjacent. Ranunculus spp., are also nectar sources (Emmett & Heath 1989), but this has not been observed in West Yorkshire. Foodplants In the south of England the larval foodplant is usually the Common Violet ( Viola riviniana ), but in northern counties it is now usually the Marsh Violet ( V. palustris ). On the West Yorkshire sites it is almost entirely V. palustris, which is readily distinguished from V. riviniana by its kidney-shaped (less heart-shaped) leaves. The habit of V. palustris plants can be very variable. The leaves grow throughout the season, their widths varying from 1.5 mm to over 6 cm. On some sites they may be heavily overgrown by sedges and grasses by late summer but appear to be able to cope with this and grow through the upright vegetation, their habit becoming very extended. V. riviniana is present in small numbers on the drier areas of several sites (Tables 1 & 2). On the West Yorkshire sites, violet plants occupy discrete areas on the margins of the wettest mire habitat, usually within rush pasture. Most of these areas are of very limited extent (mean site size 167 m2) ranging up to a maximum of 400 m2. On Newby Moor the foodplant sites occupy less than 0.15% of the total area of the SSSI (Tables 1 & 2). Whilst mainly occurring towards the wetter parts of the habitat, they occur in the zones shared with sedges and Juncus. The violets often grow on the sides of tussocks in areas dominated by Sphagnum or Juncus. Cotton grasses ( Eriophorum spp.), are present on the wettest (more waterlogged) parts of most of the sites but not in conjunction with the Marsh Violets. The presence of Marsh Thistle often indicates suitable locations for the less obvious violets, which usually form only a small percentage; less than 5%, of the local vegetation cover. The marginal positions of the foodplant can render it vulnerable to grazing, especially in drier years. On many sites the violets are often flooded in late winter but may be quite dry by midsummer. On some of the driest locations such as Austwick Moss (Sub- site C), the plants are very small and the leaves yellow later in the season; only in marginal areas shaded by birches and sallows do they appear healthy and this may be where oviposition is occurring. Botanical aspects of the sites Sites for the butterfly are typically on mires, rush pasture or marshy grassland on shallow clay soils with a wet peaty surface and impeded drainage. These stagno-humic gley soils are mainly derived from glacial drift, chiefly boulder clay. Sites can be divided into two main types by their vegetation characteristics: 1) Moorland mire sites: Most of these vegetation communities are some variant of Molinia caerulea - Potentilla erecta Mire (NVC type M25 mire; Rodwell 1991). Molinia 7 Status of the Small Pearl-bordered Fritillary in Western Yorkshire may be dominant around the edges but the violets are found in conjunction with Juncus acutiflorus and J. effusus. Sub-shrubs such as Calluna vulgaris, or on wetter ground Erica tetralix , are also present away from the wettest areas where Eriophorum spp. may dominate. These types of site lack any major trees but small sallows and stunted birches may provide shelter for adult butterflies as on Cocket Moss and Swarth Moor. No trees are present on Long Preston Moor or on any of the Newby Moor sites although hedge-line willows and sallows are present on the southern margin of Hardacre Moss. 2) Rush pasture sites: Mainly NVC M23 Juncus effusus! acutiflorus - Galium palustre rush pasture, these are meadows and pastures of moist mineral and peaty soils with flushing or impeded drainage. This community can be indicative of base enrichment These sites have more trees in their vicinity and have Marsh Violet patches amidst boggy rushy meadows dominated by Juncus effusus and J. acutiflorus, typically with Galium palustre and Potentilla erecta which is strongly preferential for the M23a J. acutiflorus Sub- community. Rushes are characteristically dominant with abundant Molinia. J. effusus also tends to be tussocky and these structures enable other species such as V. palustris to survive. The J. acutiflorus Sub-community is virtually continuous with Molinia - Potentilla grassland which is found around the margins of some of the the mires. The central site at Lawkland Moss and the Wharfe Wood sites typify this type. The latter sites are surrounded by copses and small stands of birch and hazel, with some hedgerow sallow and hawthorn. As Rodwell (1991) states ‘fen meadows and rush pasture are difficult to define but are generally the product of some sort of agricultural treatment’. The vegetation on the Newby Moor sites is typical of many moorland mire sites on acidic rocks or impervious substrates. Many sites represent the latest serai stages of late glacial lakes. Sniddle Moss (Newby Moor SSSI) is documented from cores to have passed from a lake with a wide variety of aquatics including stoneworts, through several terestrialization stages of fen and fen carr. This included the establishment of dense Alnus glutinosa and Betula pendula and B. pubescens woodland, from c.7,300 BP (Oybak 1993; Oybak & Bartley 1994, 1997). Austwick Moss, Cocket Moss, Locker Tarn and Swarth Moor also represent the wetter residual parts of much more extensive areas of M25 mire vegetation whose extent has been greatly reduced by combinations of serai progression, drainage, grazing and agricultural ‘improvement’. The dryer parts of these sites are dominated by Molinia caerulea, with Deschampsia flexuosa, D. cespitosa , Anthoxanthum odoratum Holcus lanatus, Festuca ovina, Nardus stricta, and herbs such as the P. erecta and Galium saxatile. These areas may merge into wet heath vegetation dominated by heathers ( Calluna . vulgaris and Erica spp.), and Vaccinium myrtillus. The Marsh Violet food plant is found in the damper areas typically between tussocks of Juncus effusus and J. acutiflorus along with small amounts of J. inflexus, J. articulatus and J. conglomeratus . Various sedges (e.g. Carex spicata, C. flacca, C. panicea and Luzula multiflora) are also typically present as are various species of Sphagnum moss (especially S. recurvum ). Forbes such as Cirsium palustre, Lychnis flos-cuculi, Cardamine pratensis, Galium palustris and Ranunculus spp. are usually present near to the violets. Erica cinerea is found on wetter heath often with hummocks of the mosses Polytrichum commune and Calliergon cuspidatum. In the very wettest parts of many sites is an anoxic mire vegetation dominated by Eriophorum vaginatum and E. angustifolium and mosses such as Sphagnum recurvum, S. magellanicum and S. papillosum, which merges into ‘blanket bog’ vegetation. Wetland plants such as Menyanthes trifoliata, Geum rivale and Narthecium ossifragum occupy nutrient-rich and less acidic mire locations. Alongside the sluggish drains or minor streams which meander through several of the sites (e.g. Swarth Moor, Cocket Moss and most of the Newby Moor sites) is a well defined strip (2-3 m wide) of M27 mire ( Filipendula ulmaria - Angelica sylvestris tall herb fen) where F. ulmaria is found in profusion as are clumps of Mentha aquatica. The classical site of Austwick Moss is characterised by a mosaic of lowland wet heath and residual raised mire bordered by dense wet birch scrub/woodland. The violets extend into the denser shade of the trees in several places. The main site at Sub-site A is on the TABLE 1. Characteristics of B. selene sites on the Newby Moor SSSI. Status of the Small Pearl-bordered Fritillary in Western Yorkshire Key Trees on breeding site 0 = No trees, 1 = few isolated trees or bushes, 2 = Significant trees Trees adjacent site 0 = No trees, 1 = few isolated trees or bushes, 2 = Significant trees Site Wetness 0 = dry, 1 = damp, 2 = wet, 3 = very wet ? - suitable site species not confirmed present * = Viola palustris + V. riviniana ** = Viola riviniana TABLE 2. Characteristics of other B. selene sites. Status of the Small Pearl-bordered Fritillary in Western Yorkshire 9 Key Trees on site 0 = No trees, 1 = few isolated trees or bushes, 2 = Significant trees Trees adjacent to site 0 = No trees, 1 = few isolated trees or bushes, 2 = Significant trees Site Wetness 0 = dry, 1 = damp, 2 = wet, 3 = Very wet ? = suitable site species not confirmed present * = Viola palustris + V. riviniana ** = Viola rivineana 10 Status of the Small Pearl-bordered Fritillary in Western Yorkshire wettest area with abundant V. palustris. Austwick Moss is unusual in being the only site with large numbers of V. riviniana especially at the eastern end (Sub-site B) where there is virtually no V. palustris. This may reflect the drier nature of these locations. There are some indications that Sub-site C is drying out and is evolving into herb rich grassland; certainly the small sub-colony has appeared less abundant in recent years (Stuart Ralph & Brian Shorrock, pers. comm.). The whole of Austwick Moss is fenced off and managed for game, and is gradually being overwhelmed by the spread of birch scrub. Boloria selene is present in a railway cutting near Otterburn (SD878571) on scattered V. riviniana in neutral herb-rich grassland. These banks host a relict population derived from that which was probably present on nearby rush pastures adjacent to the small beck flowing north from a larger wet area bordering Pitts Hill. These areas of rush pasture are now useless as a fritillary habitat. North of the railway line almost all the rush pasture is drained and converted to improved pasture. The extensive (3-4 ha) marshy area to the south (SD875566), is now almost devoid of marsh violet and most other forbes and has been converted by eutrophication and heavy grazing to species-poor wet rush pasture dominated by rank Deschampsia cespitosa and J uncus spp. The residual butterfly population is now restricted to the railway embankments. Unless serai progression is halted by scrub cutting, it is doomed to extinction on this site. The site in Gisburn Forest (VC64, now in Lancashire) is a NVC type M23 J uncus ejfusus/ acutiflorus - Galium palustre rush-pasture. It was planted with conifers in the early 1950s by the Forestry Commission, harvested in 1983, and replanted in 1987. Many conifers planted in the wetter parts now occupied by the Marsh Violets did not prosper and the tallest were only 2 m in height. However, the lack of grazing allowed scrub birch and some sallow to invade the area and the J uncus is very rank. In the south of England 35% of B. selene colonies were found in young conifer plantations but the future for butterflies in such locations is limited, unless active management is undertaken to conserve the ground flora (Warren & Key 1991). The altitude of the sites where the species has been confirmed ranges from 130 m to 257 m (mean 171 m). The height range of the sites on Newby Moor is 145 m to 165 m (mean 158 m). Local Agriculture, Conservation and Management Grazing Regimes Grazing on all of the butterfly’s breeding sites is light in comparison to the enclosed improved pastures of the region. Sites can be divided according to intensity of grazing regimes into three main groups: 1) Sites grazed by farm stock, mainly sheep but with a few cattle (Long Preston Moor, Newby Moor, Swarth Moor, Lawkland Moss Sub-site A, Wharfe Wood). Farmers do not now allow cattle to range freely and sheep are loath to penetrate onto the wetter parts of mire sites; consequently grazing pressure is often very small in the wetter areas. The Newby Moss SSSI is subject to common grazing rights that extend beyond the boundaries of the SSSI. These rights appear to be more traditional than formally enshrined and the commoners have habitually temporarily enclosed certain areas, now usually with electric fences, which allows intensive grazing. This often follows the flail mowing of rush pastures. 2) Fenced-off sites maintained for forestry or game cover. These are subject to very low levels of grazing mainly by the Brown Hare, Roe and Muntjac Deer (Austwick Moss, Lawkland Moss Sub-sites B, C, and D, Gisburn Forest). 3) Fenced-off sites with virtually no grazing (Cocket Moss, Otterburn Railway Embankment). Other Agricultural Operations Rush cutting is allowed on parts of the Newby Moor SSSI, with mown areas on the west 11 Status of the Small Pearl-bordered Fritillary in Western Yorkshire side of Crook Beck, the south-west of Tewitt Hall Farm and at Newby Mire. Cutting, originally by hand, was intended to reduce the vigour of Juncus and encourage a more open vegetation but is now undertaken annually by close flail mowing to produce big bale silage. The effects of this change needs a more complete assessment of its impact on the flora. Fortunately it can only be done in late summer when the land is at its driest. It usually takes place in mid to late August and generally presents little further danger to the butterfly populations, as there are now few violets in those areas. The single exception could be Newby Mire (Site 11), which is fairly dry but breeding success utilising the sparse violets of that location is questionable. The presence of Boloria selene here probably depends on colonists from the adjacent Gill Sike sites. The tendency of the mowing to encroach onto the adjacent Molinia grassland has been noted by English Nature and will be discouraged, as will any encroachments on to other known breeding sites. Streams have been deepened on many of these sites, e.g. the outflow streams of both Sniddle Moss and Hardacre Moss (Gill Sike). This has increased the drainage and probably reduced the extent of the central mires. Close to Austwick and Lawkland Mosses, Fen Beck, the main area drain on the southern boundary, was subject to extensive deepening by the drainage authority in 1984, as was its tributary, the stream near the western boundary of Austwick Moss. This has definitely damaged the SSSI. On the western side of Austwick Moss three parallel drainage ditches were cut across the SSSI to these streams; this was not a permitted operation and was never rectified, despite protests by local conservationists. Damming of the ditches to reduce the drainage of the Moss could help conserve this area of rare lowland mire, which is currently in a major phase of decline. Drainage ditches have also been cut on the Long Preston Moor and Gisburn Forest sites. Management No sites have been specifically managed to conserve or encourage Boloria selene but certain incidental aspects of management probably benefit the species. A tussocky vegetation structure appears important but light grazing is essential to prevent Juncus and grasses from becoming tall and rank and to produce a more open structure suitable for the violets. Not all species of Juncus are equally suitable for the violets. In several mires J. articulatus, which does not form tussocks, occurs in dense patches which are attractive to sheep but become wind blown later in the season and rot away during winter. This is detrimental to the violets though they are able to grow up through dense standing vegetation of grass or Juncus tussocks. Tall rank Juncus growth promoted by eutrophication shades out other plants and may be flattened by wind when it forms a dense wet mat. Floral changes observed at Austwick Moss (Sub-site C) may be symptomatic of insufficient grazing. The Moss was regularly grazed by cattle within living memory and the extensive birch cover has only developed in the last four decades, overwhelming over half of the area of original designated SSSI. The fencing off of Cockett Moss took place about seven years ago and this may be similarly detrimental in the long term unless controlled grazing is introduced in future years. Similarly Sub-sites C and D on Lawkland Moss had not been grazed for some years and this was starting to affect the quality of the vegetation but some scrub cutting has recently been undertaken. Excess grazing can also destroy sites and it is unfortunate that the disused railway line bisecting the north part of Newby Moor was omitted from the SSSI designation. Its previously rich flora with abundant common butterflies has now been totally destroyed by intensive grazing over the last twenty years. (B. & E. Shorrock pers. comm.). This was probably an important nectar site for the adjacent Small Pearl-bordered Fritillary populations. The ban on fertiliser applications on all the SSSIs has maintained floral diversity overall, but illegal applications in the past have seriously reduced floral diversity on parts of Newby Moor especially the mowed areas west of Tewit Hall Farm (SD71 1700) and near the roadside east of Sniddle Moss. Elsewhere on the moor, fertiliser applications have totally destroyed the floral value of about 2 ha west of Hardacre Farm (SD709689), 12 Status of the Small Pearl-bordered Fritillary in Western Yorkshire northeast of Newby Mire at SD724692 and adjacent to Crook Beck (SD723697 & SD723693). The western margin of Sub-site A at Lawkland Moss also appears to have received fertiliser input, probably in drainage from adjacent pasture, as Juncus here is unusually dense and rank and infested with nettles. This has exterminated the violets on at least 20% of the Sub-site. The illegal application of slurry adjacent to roadways in the Tewit Hall Farm area of Newby Moor still continues periodically and is difficult to stop unless the perpetrators are caught in the act. The area of Crook Beck received a heavy application of farm slurry as recently as November 2000. Periodic rush cutting can have beneficial effects. If used as reclamation management it can progressively reduce the fertility of eutrophic areas by removing nutrients with the biomass and could eventually allow the fertiliser damaged areas to recover some of their original floral diversity. It promotes a more open vegetation where smaller plants prosper and more nectar plants are available. Occasional cutting may have enhanced the floral diversity of parts of Newby Moor adjacent to Site 2 (north of the disused railway line). However, annual cutting will eventually destroy the rush pasture habitat. The survival of B. selene at the Gisburn Forest site for over twenty years, despite over- planting of the mire by conifers, is probably the fortuitous result of the site being adjacent to the original compartment boundary and access roadway. The butterfly probably survived on V. riviniana which is still sparsely present in that adjacent area. After timber harvesting the re-vegetation of the mire with the re-establishment of V. palustris allowed it to spread back onto the mire. When it became aware of the species, Forest Enterprise took a robust management attitude and all the conifers and scrub birch and sallow were removed from the site in Autumn 1999 and the possibility of selective Juncus cutting on parts of the site, aimed at reducing the ranker vegetation, is being considered for future years. The Butterfly populations: Most animals and plants are distributed across the landscape in a non-random manner, the sub-populations being linked by dispersing propagules (seeds or spores in plants or individual animals). Sets of spatially dispersed populations (metapopulations) interact within a dynamic environment where habitats and sub-populations are gained or lost at variable rates. The result of alternate extinctions and recolonisations in habitat patches is a shifting distribution pattern over the years (Hanski & Gilpin 1991, 1997). Fragmentation describes the process whereby a species’ habitat is dissected into smaller units separated by strips of unsuitable land. How this affects survival depends on the extinction rates of sub- populations and the colonising ability of the species from adjacent sub-populations (Opdam 1990; Hanski 1991). B. selene may have a long history on some of the West Yorkshire sites. It almost certainly became restricted to mire habitats following the general and massive deforestation of 2000 BP (Oyback & Bartley 1997; Warren & Key 1991) and the consequent loss of V. riviniana sites. Many of these mire sites were associated with the lowland flood plain of the River Ribble, but most of those have now been lost due to agricultural improvement and drainage and are now confined to a very few slightly higher, more remote sites. The Newby Moor populations make up a metapopulation which is possibly linked by occasional migration with the seven populations on Austwick and Lawkland Mosses and also perhaps those of Wharfe Wood. It is suggested that this type of breeding population may be typical of B. selene populations in northern Britain (Barnett & Warren 1995). On some sites it is probable that the population is totally isolated (Cocket Moss, Gisburn Forest, Long Preston Moor, Otterburn), while others may fall into a spectrum between the extremes (Swarth Moor & Studfold Moss) where occasional vagrants from Wharfe Wood may influence the gene pool. How much migration occurs between the main breeding locations of the Newby Moor/Austwick area and other sites is unknown and would need a detailed mark release and recapture study to elucidate it. Evidence concerning mobility is contradictory. Thomas and Snazell (1989) showed that woodland populations of B. selene in Dorset were extremely sedentary. Movement of marked individuals to suitable, newly Status of the Small Pearl-bordered Fritillary in Western Yorkshire 13 coppiced areas was very small and vacant sites were not colonised except when the area was immediately adjacent to the original population. Barnett and Warren (1995) suggested that the northern populations may be more mobile but no supporting data have been produced. In West Yorkshire very few butterflies have been seen more than 100 m from suitable habitat patches. This, however, might be related to the priority given by observers to visiting the breeding sites. On emergence, females are very sedentary but this may alter later after mating. Females of all animal species are the colonisers and their dispersive ability determines survival in a changing fragmented habitat. Genetic isolation will occur if the dispersal is poor and the resultant loss of genetic variability may lower the fitness of the individuals in a sub-population (Opdam 1990). Dempster (1991) demonstrated morphological changes (ratio of thoracic width to length) in isolated populations of endangered butterfly species. He considered that the changes could be a measure of a reduction in mobility and that isolation may select for less mobile forms. How observed differences of behaviour between the sexes relate to colonisation ability and genetic isolation of the northern B. selene populations is unknown. Hanski (1985) warns that human activities and other factors often decrease the density of habitat patches and such changes may cause the regional extinction of specialist species before the last habitat patches have been destroyed. It could be that we are seeing this sort of erosion of the habitats of the Small Pearl-bordered Fritillary in West Yorkshire. Summary The Small Pearl-bordered Fritillary, Boloria selene, has become rare in south, central and eastern England. In Yorkshire it has been confined to wet grassland and marshy upland areas in Watsonian vice counties 62 and 64. Its status in West Yorkshire has been unverified since the mid-1980s records published in Sutton and Beaumont (1989). In the present study at least 18 breeding locations for B. selene have been identified in rush pasture and mire habitats in VC64. Other sites where the species may be present have been identified. This probably represents an improved identification of the habitat locations rather than any range expansion. The vegetation, management and other characteristics of the sites are discussed and the life-cycle of the insect is summarised. Local populations are primarily using Marsh Violet ( Viola palustris ) as the larval food plant on at least 32 smaller sub-sites within the main locations. The populations on Newby Moor SSSI represent the greatest concentration of locations within the vice county and the species can be locally common. Although many sites in VC64 are SSSIs, erosion of the habitats by agricultural operations continues. However, few local populations in VC64 are considered to be at immediate risk. Habitat management of most sites is sub-optimal or non-existent and this is affecting the long-term persistence of the species; we may be observing its final decline in West Yorkshire. It would be fairly simple and not overly expensive to instal management which would improve the survival prospects of this species in VC64. The remote locations of some of the sites suggest that parts of the population could be genetically isolated. Some basic aspects of the life-cycle and population dynamics such as population densities, the location of larval over-wintering sites and the degree of mobility of the imago, are inadequately known and need further study. Acknowledgements Thanks are due to Brian and Elizabeth Shorrock, Stuart Ralph and Peter Summers for generously sharing data and information about locations. Without their help this article would be less complete. I would also like to acknowledge the help given by Butterfly Conservation in confirming some of the distribution data used to produce Figure 1. Paul Evans of English Nature was most helpful in providing local information about some of the SSSIs mentioned. Edward Metcalfe of Richard Turner and Sons (Bentham) provided the maps for fieldwork. I would also like to thank Stephen Sutton for his encouragement in all things lepidopteran. 14 Status of the Small Pearl-bordered Fritillary in Western Yorkshire References Asher, J., Warren, M., Fox, R., Harding, R, Jeffcoate, G. and Jeffcoate, S. (2001) The Millenium Atlas of Butterflies in Britain and Ireland. Oxford University Press, Oxford. Barnett, L. K. and Warren, M. S. (1995) Species Action Plan, Small Pearl-bordered Fritillary, Boloria selene. Butterfly Conservation, Wareham. Butterfly Conservation (1999) Butterfly Conservation North East Regional Action Plan. Butterfly Conservation, Colchester. Clough, S. (1998) 1601 Pearl-bordered Fritillary. In: Yorkshire Lepidoptera Report 1997. (H. Frost, ed.). Argus 33: 36. Clough, S. (1999a) 1600 Small Pearl-bordered Fritillary. In: Yorkshire Lepidoptera Report. (H. Frost, ed.). Argus 35: 46. Clough, S. (1999b) 1601 Pearl-bordered Fritillary. In: Yorkshire Lepidoptera Report. (H. Frost, ed.). Argus 35: 46. Clough, S. (2000a) 1600 Small Pearl-bordered Fritillary. In: Yorkshire Lepidoptera Report. (H. Frost, ed.). Argus 37: 46. Clough, S. (2000b) 1601 Pearl-bordered Fritillary. In: Yorkshire Lepidoptera Report. (H. Frost, ed.). Argus 37: 46-47. Clough, S. and Robinson, P. (1998) 1600 Small Pearl-bordered Fritillary, Boloria selene. In: Yorkshire Lepidoptera Report 1997. (H. Frost, ed.). Argus 33: 36. Dempster, P. D. (1991). Fragmentation, isolation and mobility in insect populations In: The Conservation of Insects and their Habitats (Proc. 15th Symposium of the Royal Entomological Society of London, 14-15 September 1989). (N. M. Collins & J. A. Thomas, eds): 143-153. Academic Press, London. Dennis, R. L. H., Sparks, T. H. and Hardy, P. B. (1999) Bias in butterfly distribution maps: the effects of sampling effort. Journal of Insect Conservation 3: 33-42. Duddington, J. and Johnson, R. (1983) The Butterflies and Larger Moths of Lincolnshire and South Humberside. Lincolnshire Naturalists’ Union, Lincoln. Emmett, A. M. and Heath, J. (eds) (1989) The Moths and Butterflies of Great Britain and Ireland , 7(1). Harley Books, Colchester. Emmett, A. M. and Heath J. (eds) (1991) The Moths and Butterflies of Great Britain and Ireland , 7 (2). Harley Books, Colchester. Frost, H and Robinson, P. (1997) 1600 Small Pearl-bordered Fritillary, Boloria selene-, In: Yorkshire, Lepidoptera Report 1996, (H. Frost ed.). Argus. 31: 37. Hanski, I. A. (1985) Single-species spatial dynamics may contribute to long-term rarity and commonness. Ecology 66: 335-343. Hanski, I. A. (1991) Single-species metapopulation dynamics: concepts, models and observations. Biol. J. Linn. Soc. 42: 17-38. Hanski, I. A. and Gilpin, M. E. (1991) Metapopulation dynamics: brief history and conceptual domain. Biol. J. Linn. Soc. 42: 3-16. Hanski, I. A. and Gilpin, M. E. (eds) (1997) Metapopulation Biology: Ecology, Genetics and Evolution. Academic Press, London. Mounsey, J. (2001) What is going on in Sedbergh?, Butterfly Conservation North West (Cumbria) Branch Newsletter, No. 3 (September 2001), 12. Opdam, P. (1990) Dispersal in fragmented populations: the key to survival. In: Species Dispersal in Agricultural Habitats (R. G. H. Bunce & D. C. Howard, eds): 3-17. Natural Environment Research Council (ITE)/Belhaven Press, London. Oybak, E. (1993) Palaeoecological studies of selected mires in the Craven District of West Yorkshire with special reference to the late Devensian period and the Ulmus decline. Unpubl. PhD thesis, University of Leeds. Oybak, E. and Bartley D. D. (1994) Some sub-fossil stone wort records from two late glacial and post glacial sites in the Ingleborough region. Bulletin YNU 21: 6-7. Oybak, E. and Bartley D. D. (1997) Lateglacial and postglacial moss records from a valley bog, Sniddle Moss, in the Ingleborough region. Naturalist 122: 53-56. Status of the Small Pearl-bordered Fritillary in Western Yorkshire 15 Parkinson, D. (1997) 1601 Pearl-bordered Fritillary. In: Yorkshire Lepidoptera Report. (H. Frost, ed.). Argus 31: 38. Porritt, G. T. (1883) List of Yorkshire Lepidoptera. Ent. Trans. Nat. Un. 2: 1-158. Porritt, G. T. (1904) List of Yorkshire Lepidoptera (Supplement). Ent. Trans. Yorks. Nat. Un., Ser D 2, Part 30, i-xvi, 193-269. Rodwell, J. S. (1991) British Plant Communities . Volume 2, Mires and Heaths. Cambridge University Press, Cambridge. Skelton, M. J. and Heath, J. (1975) Lepidoptera Rhopalocera, Butterflies; Provisional Distribution Maps. Biological Records Centre, Monkswood. Skinner, B. (1984) Colour Identification Guide to Moths of the British Isles. Viking, London. South, R. (1962) The Butterflies of the British Isles. 6th ed. Wayside and Woodland Series, Warne, London. Sutton, S. L. S. and Beaumont, H. B. (eds) (1989) The Butterflies and Moths of Yorkshire. Yorkshire Naturalists’ Union, Doncaster Museum, Doncaster. Thomas, J. A. (1991) Rare species conservation: case studies of European butterflies. In The Scientific Management of Temperate Communities for Conservation (I. F. Spellerberg, F. B. Goldsmith & M. G. Morris, eds): 149-197. Blackwell Scientific Publications, Oxford. Thomas, J. and Lewington, R. (1991) The Butterflies of Britain and Ireland , Dorling Kindersley, London. Thomas J. A. and Snazell, R. (1989). Declining Fritillaries: the next challenge in the conservation of Britain’s butterflies. Annual Report of the Institute of Terrestrial Ecology, 1988, 54-46. Walsh, G. B. (1956) Order Lepidoptera (Moths and Butterflies). In The Natural History of the Scarborough District, (G. B. Walsh & F. C. Rimmington, eds): Vol. 2 Zoology, 146- 195 Scarborough Warren, M. S. and Key, R. S. (1991) Woodlands: past, present and potential for insects. In: The Conservation of Insects and their Habitats. (N. M Collins & J. A. Thomas, eds): 155-211. Academic Press, London. YNU Lepidoptera Committee (1967) The Lepidoptera of Yorkshire. Butterflies. Naturalist 92: 55-62. APPENDIX 1 Recent Records of B. selene in VC64 Yorkshire (Site and Sub-site Records arranged by 10 km grid squares) SD 96 Local Population not confirmed Chapel House Wood, Wharfedale SD978654 30.6.78 1 female wanderer? (PS) SD 86 3 Local Populations Studfold Moss SD808699 27.6.00 10+ Swarth Moor SD806693 25.6.98 20+ (BS) 24.6.99 2 mating (PS) 25.6.99 4+ (BS) 27.6.00 10+ (BS) SD805692 6.7.99 1 patrolling Long Preston Moor SD825605 27.6.00 10+ (ES/BS) SD 85 1 Local Population Railway cutting near Otterburn SD878571 8.7.95 3+ (BS) SD 77 2+ Local Populations all on parts of Newby Moor SSSI Un-named Mire SD712701 18.6.78 + (ES) Un-named stream valley A SD709701 5.7.99 1 patrolling (North of Railway line) B SD710703 5.7.99 3 patrolling 16 Status of the Small Pearl-bordered Fritillary in Western Yorkshire APPENDIX 1 (Continued) Un-named stream valley Crook Beck nr A65 Crook Beck nr A65 C SD711702 SD711703 SD715701 SD7 15702 5.7.99 17.6.96 15.7.96 1997 7.7.99 1 10 50+ 1 1 patrolling (BS) (PS) (BS) N. on marsh thistle SD 76 14+ Local Populations? all but three on Newby Moor SSSI Cocket Moss (SE Side) A SD787619 26.6.95 30+ (PS) SD787617 1995? Several (BS) Cocket Moss (N End) B SD786620 26.6.99 3 patrolling Austwick Moss A SD760664 21.6.94 18+ (BS) 7.6.97 30 (BS) 16.6.97 50+ (BS) 6.7.97 8 (BS) 3.7.98 10 (BS) B SD759667 1999 + (SR) C SD764669 25.6.98 5+ Lawkland Moss A SD768664 7.6.97 8+ (BS) B SD769664 + (SR) C SD769666 25.6.98 1 very small colony D SD769667 + (SR) Un-named Mires Newby Moor SD708699 5.7.99 1 patrolling (South of Railway line) SD7 10698 5.7.99 1 patrolling Sniddle Moss (East) A SD707696 5.7.99 2 patrolling 27.6.00 4+ patrolling SD708696 5.7.99 1 patrolling Sniddle Moss (West) B SD709696 24.6.99 1 patrolling SD708697 27.6.00 1 patrolling Sniddle Moss Valley C SD7 10696 5.7.99 1 patrolling 13.7.99 1 female disturbed from veg. SD711696 24.6.99 2 patrolling 27.6.00 4+ 3 patrolling. 1 Female SD7 12696 15.7.96 30+ (PS) 24.6.99 1 patrolling 5.7.99 1 roosting on Juncus New Butt SD705693 5.7.99 1 wanderer? Hardacre Moss A SD712688 25.6.99 3 2 patrolling 1 Female SD712688 5.7.99 3 patrolling Mire nr Butt Hill A SD706689 24.6.99 1 patrolling SD706690 24.6.99 1 patrolling B SD708689 17.7.96 4 (BS) Mire E of Banks Head SD7 17692 6.7.99 3+ patrolling 7.7.99 2 patrolling SD7 17692 13.7.99 1 male patrolling 1 female disturbed from veg. 26.6.00 4 patrolling Gill Syke, Green Close A SD7 19692 6.7.99 3+ patrolling 26.6.00 5+ patrolling/ basking Notes on some Yorkshire Diptera, 2001 17 APPENDIX 1 (Continued) Gill Syke, Green Close B SD720690 6.7.99 1 patrolling C SD720691 7.7.99 6+ patrolling 13.7.99 4+ 1 3 patrolling N. on bell heather 24.7.99 3+ patrolling ( 1 very worn) SD722690 7.7.99 2 patrolling 19.7.99 4+ patrolling 26.6.00 3+ patrolling Gregory Hole SD724688 7.7.99 6+ 1 5 patrolling N. on marsh thistle 26.6.00 5+ 5 patrolling SD725688 15.7.96 12+ (PS) 26.6.00 2 patrolling Newby Mire SD721693 7.7.99 1 patrolling Nr Conisber SD743671 16.7.99 1 female wanderer (PS) Wharfe Wood A SD781688 27.6.95 4 (BS) B SD786688 26.6.00 5+ patrolling C SD789688 26.6.00 5+ N. on marsh thistle SD 75 1 Local Population Gisburn Forest SD742556 19.7.96 1 (BS) Key to Record Sources & abreviations: PE = Paul Evans, SR = Stuart Ralph, BS = Brian Shorrock, ES = Elizabeth Shorrock, PS = Peter Summers. N = at nectar. NOTES ON SOME YORKSHIRE DIPTERA, 2001 ROY CROSSLEY 1 The Cloisters, Wilberfoss, York YQ41 5RF Stratiomyidae Odontomyia hydroleon (L.), a striking green and black medium-sized species, is one of Britain’s rarest flies (RDB1), thought to be extinct in these islands until a specimen was taken in south-west Wales in 1986; subsequently it was found at a site with calcareous seepages in the North York Moors National Park (Stubbs & Drake, 2001). This is the only known English locality, where it has been monitored for a number of recent years. On the hot sunny morning of 16 July, for a period of an hour before mid-day, I observed a number of males hovering at varying heights up to about 18 inches above the ground. There were possibly six specimens involved, and the resemblance to hoverfly behaviour was remarkable, the flies being capable of moving rapidly sideways as well as remaining in a stationary position. The abdomen of the males is predominantly black and this, coupled with pale yellowish pubescence on the thoracic margins, gave the impression of a large Cheilosia species. Occasionally individuals would settle for a while on the leaves of sedges or other low-growing vegetation. No chasing was observed, but the behaviour seemed to be territorial, and this does not appear to have been reported before. Two females were found by sweeping low vegetation, which is the usual way of locating specimens. Naturalist 127 (2002) 18 Notes on some Yorkshire Diptera, 2001 Syrphidae Cheilosia albitarsis Mg. is a widespread spring species, usually found in association with buttercups. It is distinguished by having, on the front legs, yellowish mid-tarsal segments which contrast with the otherwise all-black legs. For some time it has been known that a closely similar species is present in Britain, and this has now been determined as Cheilosia ranunculi Doczkal (Doczkal 2000). On 1 1 May numbers of what I took to be C. albitarsis were present at an extensive colony of Lesser Celandine ( Ranunculus ficaria L.) in Park Wood, Elland, and I collected half a dozen for subsequent examination. All proved to be C. ranunculi ; subsequently I sought the species in a number of widely scattered Yorkshire localities, but found only one further example. This was at Askham Bog on 26 May when one was present in company with numbers of C. albitarsis at Creeping Buttercups (. Ranunculus repens L.). The two species are only separable in the males, and an examination of collections will probably reveal C. ranunculi mixed in with C. albitarsis, although this has proved not to be the case with my own collection. Xanthandrus comtus (Harris) appears to be predominantly a southern species (Falk 1991), for which there are few Yorkshire records. It was gratifying to find a single male example flying sluggishly amongst fringing vegetation at a pond in the Arboretum at Castle Howard on 17 August. Because of the highly episodic and random occurrences it is possible that this is a migrant species. SCIOMYZIDAE In my paper ‘Notes on the marsh flies (Diptera: Sciomyzidae) of Yorkshire’ ( Naturalist 122 [1997] pp. 93-97), I noted that the only record for Salticella fasciata (Mg.) was ‘Spurn 1928’, by C. A. Cheetham, the precise location not being given. I had overlooked subsequent records from the Warren area in 1950, several specimens having been found there during the Spurn survey undertaken by members of the Y.N.U. Entomological Section (Hincks 1953). It is gratifying to report that I swept a single female from roadside vegetation near ‘Chalk Bank’ on 4 June, thus confirming the continued presence of this nationally rare species at Spurn. S. fasciata was the subject of biosystematic studies during the 1960s based on a coastal population at Tenby, South Wales (Knutson, Stephenson & Berg 1970). A single female Ditaeniella (= Pherbellia) grisescens (Mg.) was swept on 9 August by the side of a shallow pool in a field north of Spurn Warren. This constitutes the first documented record for this Nationally Notable species in Yorkshire. The majority of British records are from coastal situations (Falk 1991). Tetanura pallidiventris Fall, is a distinctive species, the female ovipositor being flattened like a shoe-horn. It is reported here for the first time in vice-county 63 on the basis of specimens found in Park Wood, Elland. Although most of the Yorkshire sites are calcareous woodland, and in my experience the fly is often associated with extensive stands of Mercurialis perennis L. (Dog’s Mercury), this is clearly not always the case, for the Elland site is on the heavy clays of the Lower Coal Measures. In the larval stages the fly is a parasitoid of a number of terrestrial molluscs, so habitat is probably not a limiting factor of distribution. References Doczkal, D. (2000) Description of Cheilosia ranunculi sp.n. from Europe, a sibling species of C. albitarsis Meigen (Diptera, Syrphidae) Volucella 5: 63-78. Stuttgart. Falk, S. (1991) A Review of the Scarce and Threatened Flies of Great Britain ( Part 1). Research and Survey in Nature Conservation No. 39. Nature Conservancy Council, Peterborough. Hincks, W. D. (1953) XII Diptera. In: The Entomology of Spurn Peninsula. Naturalist 78:164, Hull. Knutson, L. V., Stephenson, J. W. and Berg, C. O. (1970) Biosystematic studies of salticella fasciata (Meigen), a snail-killing fly (Diptera: Sciomyzidae). Trans Roy. Ent. Soc. 122, pt. 3: 81-100, 3 plates. Stubbs, A. E. and Drake, M. (2001) British Soldierflies and their Allies. British Entomological and Natural History Society, Reading. 19 EFFECTS OF THE REINSTATEMENT OF COPPICE MANAGEMENT ON THE VASCULAR GROUND FLORA AT EAVES WOOD SSSI, SILVERDALE, LANCASHIRE JOHN JONES Department of Environmental Sciences, University of Hertfordshire, College Lane, Hatfield, Hertfordshire ALIO 9AB* Abstract This paper assesses the effects of the reinstatement of coppice management on the vascular ground flora at Eaves Wood SSSI, Silverdale, Lancashire. The study was carried out during May 2000 on a selection of neglected and actively coppiced areas within Eaves Wood. The vegetation of the various areas was compared and likely causes of differences were examined. The results showed that the greater ground flora cover, species richness and number of species per quadrat in certain areas appeared to coincide with the reinstatement of coppice management. Although numerous possible influences on the vegetation were recognised, it was concluded that the growth of coppice stems and the amount of leaf litter were likely to have been important influences. It was also found that the ground flora species composition has most probably been influenced by coppice reinstatement, many species having a tendency to occur in coppiced areas. Furthermore, the flowering vigour of certain species was found to be increased in recently coppiced areas. The results did, however, suggest that a small number of shade tolerant species might have benefited from non-intervention. Introduction Despite the fact that coppicing was originally employed as a timber production technique, it is now widely regarded as being an important form of management for the conservation of flora and associated fauna in certain woodlands (Kirby, 1993). However, any form of habitat management is likely to favour some species and harm others (Goldsmith, 1992). The widespread reinstatement of coppice management is largely the result of concern among conservation managers that ground flora, particularly that of open woodland areas, would be diminished without canopy management (Barkham, 1992a), especially in isolated woodlands (Peterken, 1992). By increasing the rate of gap formation, coppicing can aid the conservation of woodland glade species in small woodland fragments, where the minimum dynamic areas necessary for the survival of some species may be greater than would be provided if the woodland remained unmanaged (Kirby, 1996). However, many woodland species, especially those which thrive on shade and moisture, may be harmed by coppice reinstatement (Hambler & Speight, 1995). Because of poor dispersal powers and short lived seed banks, such species are particularly vulnerable to management methods that reduce their dominance in the vegetation (Brown & Oosterhuis, 1981). Certain species may also be smothered by tall vegetation resulting from increased light levels and the exclusion of grazers during the early stages of a coppice rotation (Barkham, 1992b). Others may be unable to tolerate the exaggerated microclimate fluctuations associated with coppicing (Hambler & Speight, 1995). Because of a poor understanding of the requirements of many species and of some of the influences of coppicing, there is widespread reliance on the premise that the species of a site with a history of coppicing are best conserved by coppice reinstatement. Validation of this assumption, through vegetation surveys following coppicing, is desirable in order to ensure the conservation of diversity and of important species on individual sites. * Current address: Hangman’s Cottage, 3 A St. Mary’s Square, Horncastle, Lincolnshire LN9 5HJ Naturalist 127 (2002) 20 Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora This paper assesses the effects of reinstated coppice management on the vascular ground flora at Eaves Wood SSSI. A selection of coppice coupes, i.e. areas of woodland cut in a particular year of a coppice rotation, including neglected and restored coupes with varying ages of regrowth, are compared in terms of ground flora cover, species richness, number of species per quadrat, diversity and evenness of species distribution, species composition and flowering vigour. Possible reasons for differences in their vegetation are then examined. Eaves Wood SSSI is owned by the National Trust and is situated 1 kilometre northeast of Silverdale in Lancashire (grid ref. SD466763). The site lies within the Arnside and Silverdale Area of Outstanding Natural Beauty, which encompasses numerous other woodlands and areas of ecological importance. Eaves Wood is a 50 ha. woodland site, which mainly occupies the southern aspect of a low carboniferous limestone hill and is a designated SSSI on account of its limestone woodland and grassland flora. The site contains a mosaic of habitats, including broad-leaved, mixed and coniferous woodland, scrub, limestone pavement, calcareous grassland and heath. The dominant habitat is broad-leaved semi-natural woodland, including areas of coppice with standards. The coppice stools are dominated by Corylus avellana, but with abundant Fraxinus excelsior, and the standard trees largely consist of Quercus spp. and Fraxinus excelsior. Much of Eaves Wood was coppiced until the early 20th century, but was neglected from the mid- 1900s until coppice management was reinstated in the winter of 1991/92 (National Trust c. 1993; Alexander et al., 1984). Methods Selection of study areas Two neglected coppice coupes were selected, which, according to ring counts, appeared not to have been coppiced since around 1950 and 1960 respectively. Five restored coppice coupes were also selected, which were coppiced in the winters of 1993/94, 1995/96, 1996/97, 1997/98 and 1998/99. Study areas were selected on the grounds of similarity of site conditions (Table 1) and proximity to one another, in order to minimise differing influences on the vegetation. TABLE 1. Coppice coupe descriptions. 1998/99 1997/98 1996/97 1995/96 1993/94 c. 1960 c. 1950 Restoration Restored Restored Restored Restored Restored Neglected Neglected Area (M2) 375 810 1,160 810 2,250 960 2,300 Geology Limestone Limestone Limestone Limestone Limestone Limestone Limestone Aspect (approx.) Southwest South South South South Southwest South Altitude (approx.) 60 m 65-70 m 65 m 65-70 m 65-70 m 60-65 m 55-65 m Deer fencing Fenced Fenced Unfenced Unfenced Unfenced Unfenced Unfenced Dominant Corylus Corylus Corylus Corylus Corylus Corylus Corylus species avellana avellana avellana avellana avellana avellana avellana Sampling Sampling was carried out during May 2000, when a relatively complete range of vascular ground flora species would have been visible (Kirby, 1988). Data were collected on vascular ground flora and other ground cover features (Table 2), from 30 quadrats of 1 m2 within each coupe. The outer 3 metres of the coupes were not surveyed, in order to reduce Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora 21 the likelihood of complications resulting from edge effects. Data were collected on coppice stools and standard trees (Table 2), from up to five quadrats of 10 x 10 metres within each coupe. All quadrats were distributed on a stratified random basis (Kirby, 1988). TABLE 2. Data collected from each coupe. Coupe Feature Data Collected Coppice stools No. of individuals of each species within each quadrat. Maximum height of regrowth for each species. Maximum number of stems for each species. Girth range of stems for each species. Standard trees No. of individuals of each species within each quadrat. The range of diameters at breast height (DBH) for each species, measured at 1.3 m above ground level (Kirby, 1988). Vascular ground flora The percentage cover of each species (visually estimated to the nearest 5%). Presence/absence of flowers and fruit within the quadrat for each species. Other ground cover features e.g. leaf litter Percentage cover (visually estimated to the nearest 5%). Data analysis The vascular ground flora of the various coupes was compared in terms of overall mean percentage cover, the mean number of ground flora species per quadrat and the mean percentage cover of individual species (Table 3). The significance of differences was assessed using analysis of variance (ANOVA). The coupes were also compared in terms of species richness and the frequency of occurrence of individual species, as well as the frequency of occurrence of flowers and fruit on individual species (Table 3), and the significance of differences was tested using Chi-Squared tests (Powell, 1996). The diversity and evenness of species distribution within each coupe were then calculated, using the Shannon- Weiner Diversity Index (Kent & Coker, 1992). The similarity of species composition between coupes was also tested using the Czekanowski Coeffcient (Kent & Coker, 1992). The Spearman Rank Correlation Coefficient (Powell, 1996) was used to establish whether the growth or density of stools and standards, time since coppicing or leaf litter cover were related to the ground flora cover, number of species per quadrat, species richness, diversity or evenness of species distribution. Results Vascular ground flora cover Significantly less ground flora cover was recorded in neglected coupes compared to restored coupes (Table 4), the 1998/99 coupe having the greatest cover and the 1960 coupe having the least. No statistically significant correlation between ground flora cover and growth or density of stools and standards, litter cover or time since coppicing was found. However, there was some evidence of negative correlation between ground flora cover and the height of coppice regrowth (rs = -0.759, p<0.1, n = 7). 22 Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora TABLE 3. Definitions and application of vascular ground flora measurement criteria. (Adapted from Kent & Coker, 1992; Powell, 1996). Measurement Definition Application Mean percentage cover Mean area covered by overall ground flora or individual species in the survey quadrats of a coupe. Comparison of overall field layer productivity or growth vigour of individual species between coupes. Species richness The total number of ground flora species recorded within the survey quadrats of a coupe. Comparison of the number of species able to tolerate the conditions throughout the various coupes. Mean number of species per quadrat The sum of species scores for the quadrats of a coupe, divided by the number of quadras. Comparison of the number of species able to tolerate the prevailing conditions in the various coupes giving less weight to species inhabiting only part of a coupe. Diversity (Shannon -Weiner index) S Diversity H' = X pi In pi i=l s = Number of species pi = Abundance of the ith species. In = log base n Quantitative comparison of the diversity of coupes, combining species richness with relative abundance of species. Evenness of species distribution (Shannon -Weiner index) H' Equitability J = Hmax IT = Diversity H max - In S s = Number of species Quantitative comparison of coupes in terms of relative abundance of individual species, which may be indicative of factors such as habitat suitability or inter-specific competition. Frequency of occurrence of species The number of quadrats within a coupe in which an individual species was recorded. Assessment of habitat preferences of species, allowing for those which tend not to have extensive ground cover. Similarity of species composition (Czekanowski coefficient) m 2Xmin (X;, Yj) i=l Sc = Quantification of similarities in species composition to give an indication of influences on vegetation. SXi + f Yi i=l i=l Xi? Yj = abundances of species i m X min (Xj, Yj) = sum of lesser i=i scores of species i m = number of Species Frequency of occurrence of flowers and fruit The number of quadrats in which flowers or fruit were present on an individual species as a proportion of the number of quadrats where that species was present. Comparison of the flowering vigour of a particular species in different coupes. Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora 23 Vascular ground flora species richness Significantly fewer vascular ground flora species were recorded in the neglected coupes compared to the restored coupes (Table 4). The greatest number of species was recorded in the 1996/97 coupe, and the fewest in the 1960 coupe. No statistically significant correlation was found between species richness and features relating to stools, standards, leaf litter or time since coppicing, although there was some evidence of negative correlation between species richness and leaf litter cover (rs = -0.714, p<0. 1 , n = 7). TABLE 4. Vascular ground flora characteristics of coupes, including statistical test results. Vegetation feature 98/99 97/98 96/97 95/96 93/94 c. 1960 Signifi- es. 1950 cance Mean percentage cover 133.83 40.67 63.33 72.00 68.33 19.83 25.50 ANOVA p = 0.0001 Species richness 24 18 26 19 21 7 8 x’= 18.76 df = 6 p<0.01 Mean number of species per quadrat 6.73 2.97 4.77 4.83 4.46 0.77 2.10 ANOVA p = 0.0001 Diversity (Shannon- Weiner index) 2.12 1.52 2.08 1.91 2.32 0.74 1.53 Equitability of species distribution (Shannon- Weiner index) 0.67 0.53 0.64 0.65 0.76 0.38 0.74 Number of vascular ground flora species per quadrat A significantly lower mean number of ground flora species per quadrat was found in neglected coupes (Table 4). The highest number of species per quadrat was recorded in the 1998/99 coupe and the lowest in the 1960 coupe. Although not statistically significant, there was some evidence of negative correlation between the number of species per quadrat and time since coppicing (rs = -0.723, p<0.1, n = 7), the height of coppice regrowth (rs = -0.723, p<0.1, n = 7) and the girth of coppice regrowth (rs = -0.723, p<0.1, n = 7). Diversity and evenness of species distribution The neglected coupes and the 1997/98 coupe were found to have relatively low ground flora diversity (Table 4). The coupe with the greatest diversity was the 1993/94 coupe. No correlation was found between diversity and other coupe features. The 1993/94 coupe and the 1950 coupe had the most even species distribution (Table 4), and the 1960 coupe had the most uneven species distribution. No correlation was found between evenness of species distribution and other coupe features. 24 Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora Vascular ground flora species composition Significant differences between the coupes were found in the percentage cover of eight species (Table 5). These were Epilobium montanum, Fragaria vesca Hedera helix and Mercurialis perennis, which had greatest cover in the 1998/99 coupe; Rubus fruticosus and Viola rivianax which had greatest cover in the 1995/96 coupe; and Helianthemum chamaecistus and Pteridium aquilinum. which had greatest cover in the 1993/94 coupe. In terms of percentage cover, Rubus fruticosus was dominant in all restored coppice coupes, although Fragaria vesca and Hedera helix were also abundant in the 1998/99 coupe. Hedera helix was dominant in the 1960 coupe and Mercurialis perennis and Hyacinthoides non-scriptus were co-dominant in the 1950 coupe. TABLE 5. Mean percentage cover of vascular ground flora species, including results of ANOVA statistical test (NS = Not significant). Species Mean Percentage Cover per Coupe ANOVA 98/99 97/98 96/97 95/96 93/94 1960 1950 Arum maculatum 0.8 1.2 0.0 0.0 0.5 1.5 1.3 NS Brachypodium sylvaticum 1.5 0.5 2.3 0.0 3.2 0.0 0.2 NS Carex flacca 0.0 0.0 0.0 0.7 1.3 0.0 0.0 NS C. sylvatica 0.0 0.0 0.2 0.0 0.3 0.0 0.0 NS Chamaenerion angustifolium 0.3 0.0 1.5 1.5 1.5 0.0 0.0 NS Cirsium arvense 0.2 0.2 1.2 0.7 0.0 0.0 0.0 NS C. vulgare 0.5 0.0 0.0 0.0 0.0 0.0 0.0 NS Cotoneaster sp. 0.0 0.0 0.7 0.2 0.0 0.0 0.0 NS Daphne laureola 0.0 0.0 0.7 0.3 0.0 0.0 0.0 NS Epilobium hirsutum 1.8 1.0 1.3 1.8 1.7 0.0 0.0 NS E. montanum 5.3 2.2 2.5 0.8 0.8 0.0 0.0 p=0.0001 Fragaria vesca 29.5 3.5 5.5 6.2 10.5 0.0 0.3 p=0.0001 Geranium robertianum 1.2 0.2 0.0 0.0 0.5 0.0 0.0 NS Geum urbanum 4.3 0.0 0.3 0.5 0.3 0.3 0.2 NS Hedera helix 26.8 0.2 0.2 0.0 0.0 16.2 0.0 p=0.0001 Helianthemum chamaecistus 0.0 0.2 4.2 3.3 5.3 0.0 0.0 p=0.0001 Hyacinthoides non-scriptus 0.0 0.0 0.0 0.0 0.0 0.0 8.2 NS Lonicera pericylmenum 0.0 2.7 0.2 0.0 0.0 0.0 0.0 NS Lotus corniculatus 0.2 0.0 0.5 0.3 0.0 0.0 0.0 NS Medicago lupulina 0.0 0.2 0.0 0.0 0.0 0.0 0.0 NS Mercurialis perennis 19.3 0.0 1.8 0.0 3.5 0.2 8.8 p=0.0001 Mycelis muralis 0.3 0.0 0.0 0.0 0.0 0.0 0.0 NS Potentilla erecta 0.0 0.0 0.2 0.0 0.0 0.0 0.0 NS P. sterilis 0.7 0.0 0.0 0.0 0.0 0.0 0.0 NS Primula veris 0.2 0.0 0.7 0.0 1.2 0.0 0.0 NS Pteridium aquilinum 0.0 1.3 0.2 2.8 4.2 0.2 0.0 p=0.007 Ranunculus repens 0.2 0.2 1.2 0.2 0.2 0.0 0.0 NS Rosa agg. 0.8 0.3 0.2 0.3 0.0 0.0 0.0 NS Rubus fruticosus 29.7 25.8 31.8 36.3 23.2 1.3 3.8 p=0.0001 Senecio jacobaea 4.3 0.0 0.0 0.0 0.0 0.2 0.0 NS Taraxacum officinale agg. 3.2 0.7 3.0 4.3 3.7 0.0 0.0 NS Teucrium scorodonia 0.0 0.0 1.0 4.8 2.0 0.0 0.0 NS Veronica chamaedrys 0.3 0.0 0.0 0.0 0.0 0.0 0.0 NS V. hederifolia 0.3 0.0 0.0 0.0 0.0 0.0 0.0 NS V. officinalis 0.0 0.2 0.3 0.0 0.3 0.0 0.0 NS Viola hirta 0.0 0.0 0.3 2.7 1.2 0.0 0.0 NS V. riviana 2.0 0.3 1.5 4.2 3.0 0.0 2.7 p=0.005 Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora 25 In terms of frequency of occurrence, Rubus fruticosus was again dominant in most restored coppice coupes, apart from the 1998/99 coupe, where Hedera helix had a slightly higher frequency (Table 6). H. helix was also dominant in the 1960 coupe, but Rubus fruticosus and Hyacinthoides non-scriptus were co-dominant in the 1950 coupe. Five species were found to have a significant tendency to occur only in areas coppiced between 1997 and 1999, where deer fencing was still in place. These were Epilobium montanum, Geum urbanum, Hedera helix. Lonicera periclymenum and Senecio jacobaea (Table 7). TABLE 6 Frequency of occurrence of vascular ground flora species (out of 30 quadrats surveyed per coupe). Species Coupe/Frequency of Occurence c. 1950 1998/99 1997/98 1996/97 1995/96 1993/94 c. 1960 Arum maculatum 4 5 0 0 1 2 5 Brachypodium sylvaticum 4 2 8 0 4 0 1 Carex flacca 0 0 0 3 1 0 0 C. sylvatica 0 0 1 0 1 0 0 Chamaenerion angustifolium 1 0 8 4 6 0 0 Cirsium arvense 1 1 5 4 0 0 0 C. vulgare 2 0 0 0 0 0 0 Cotoneaster sp. 0 0 4 1 0 0 0 Daphne laureola 0 0 2 1 0 0 0 Epilobium hirsutum 8 5 8 9 8 0 0 E. montanum 21 11 13 5 5 0 0 Frag aria vesca 20 9 15 12 19 0 2 Geranium robertianum 2 1 0 0 2 0 0 Geum urbanum 10 0 1 2 2 2 1 Hedera helix 29 1 1 0 0 9 0 Helianthemum chamaecistus 0 1 8 9 9 0 0 Hyacinthoides non-scriptus 0 0 0 0 0 0 18 Lonicera pericylmenum 0 8 1 0 0 0 0 Lotus corniculatus 1 0 2 1 0 0 0 Medicago lupulina 0 1 0 0 0 0 0 Mercurialis perennis 17 0 1 0 2 1 7 Mycelis muralis 2 0 0 0 0 0 0 Potentilla erecta 0 0 1 0 0 0 0 P. sterilis 2 0 0 0 0 0 0 Primula veris 1 0 2 0 6 0 0 Pteridium aquilinum 0 7 1 6 4 1 0 Ranunculus repens 1 1 2 1 1 0 0 Rosa agg. 3 1 1 2 0 0 0 Rubus fruticosus 27 28 30 30 27 7 18 Senecio jacobaea 16 0 0 0 0 1 0 Taraxacum officinale agg. 17 4 16 20 20 0 0 Teucrium scorodonia 0 0 3 7 3 0 0 Veronica chamaedrys 2 0 0 0 0 0 0 V. hederifolia 2 0 0 0 0 0 0 V. officinalis 0 1 1 0 1 0 0 Viola hirta 0 0 1 14 4 0 0 V. riviana 9 2 7 14 8 0 11 26 Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora TABLE 7 Significance of association between frequency of occurrence of individual species and the stage in the coppice rotation (Chi-squared tests - 2 degrees of freedom). Species t Significance Apparent Habitat Preferences Level (NS = Not Recent Older Neglected Significant) Coppice Restored Area (1997-99) Coppice (Coppiced Fenced (1993-97) 1950-60) Unfenced Arum maculatum 10.774 p<0.01 ✓ ✓ Brachy podium sylvaticum 6.047 p<0.05 ✓ ✓ Carex flacca 5.437 NS C. sylvatica 2.692 NS Chamaenerion angustifolium 23.060 p<0.001 ✓ Cirsium arvense 7.867 p<0.02 ✓ C. vulgare 5.048 NS Cotoneaster sp. 6.829 p<0.05 ✓ Daphne laureola 4.058 NS Epilobium hirsutum 19.465 p<0.001 ✓ ✓ E. montanum 44.176 p<0.001 ✓ Fragaria vesca 40.311 p<0.001 ✓ ✓ Geranium robertianum 3.244 NS Geum urbanum 7.038 p<0.05 ✓ Hedera helix 56.695 p<0.001 ✓ Helianthemum chamaecistus 36.204 p<0.001 ✓ Hyacinthoides non-scriptus 49.219 p<0.001 ✓ Lonicera pericylmenum 16.871 p<0.001 ✓ Lotus corniculatus 2.166 NS Medicago lupulina 2.512 NS Mercurialis perennis 19.471 p<0.001 ✓ ✓ Mycelis muralis 5.048 NS Potentilla erecta 1.340 NS P. sterilis 5.048 NS Primula veris 8.339 p<0.02 ✓ Pteridium aquilinum 5.575 NS Ranunculus repens 2.631 NS Rosa agg. 4.138 NS Rubus fruticosus 74.484 p<0.001 ✓ ✓ Senecio jacobaea 39.075 p<0.001 ✓ Taraxacum officinale agg. 60.120 p<0.001 ✓ Teucrium scorodonia 18.477 p<0.001 ✓ Veronica chamaedrys 5.048 NS V. hederifolia 5.048 NS V. officinalis 1.296 NS Viola hirta 27.853 p<0.001 ✓ V. riviana 5.395 NS Chamaenerion angustifolium, Cirsium arvense, Cotoneaster sp. Helianthemum chamaecistus, Primula veris, Taraxacum officinale, Teucrium scorodonia and Viola hirta were found to have a significant tendency to occur only in coupes coppiced between 1993 Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora 27 and 1997, where deer fencing had been removed. Brachypodium sylvaticum, Epilobium hirsutum, Fragaria vesca and Rubus fruticosus, were found to have a significant tendency to occur in a range of restored coppice coupes regardless of the year of coppicing or the presence of deer fencing. Arum maculatum and Mercurialis perennis were found to have a significant tendency to occur both in neglected coupes and recently restored coupes where deer fencing was present. Hyacinthoides non-scriptus had a statistically significant tendency to occur in neglected coupes, although it was present in only one of the neglected coupes. Similarity of vascular ground flora between coupes The similarity found between the vascular ground flora species composition of each pair of coupes ranged from 3.9% to 77.4% (Table 8). Between 3.9% and 24.7% similarity was found when comparing neglected coupes to restored coupes. Restored coupes were between 41.1% and 77.4% similar to one another. TABLE 8 Percentage similarity of ground flora species composition (Czekanowski coefficient). 1998/99 1997/98 1996/97 1995/96 1993/94 c.1960 c. 1950 20.2 17.5 17.5 14.4 23.9 13.2 c.1960 24.7 9.5 5.3 3.9 5.7 1993/94 47.9 59.7 73.2 72.1 1995/96 44.2 60.8 77.4 1996/97 49.3 68.2 1997/98 41.1 Frequency of occurrence of flowers and fruit The frequency of occurrence of flowers on Geum urbanum was significantly higher in restored compared to neglected coupes (x2 = 7.2, p<0.01). A significantly higher occurrence of flowers in recently restored coupes, where deer fencing was still present, compared to older restored coupes, was found on Fragaria vesca (x2 = 5.598, p<0.02), Taraxacum officinale (x2 = 15.721, pcO.OOl) and Viola riviana (x2 = 6.144, p<0.02). No significant differences were found for other ground flora species. Discussion Low ground flora cover in neglected coupes, and possible negative correlation between coppice regrowth height and vegetation cover, appears to concur with the findings of Grime (1979), who suggests that field layer production tends to decrease as the size of coppice regrowth increases. This may be due to increased shading and associated microclimate changes. The lower number of species in neglected coupes, compared to restored coupes, reflects the findings of various earlier studies (Mitchell & Kirby, 1989). The possible negative correlation between litter cover and species richness may suggest that a number of species cannot tolerate being smothered by leaf litter. Compared to the overall species richness of a coupe, the mean number of species per quadrat gives less weight to small atypical areas that may be inhabited by species not present throughout the majority of a coupe. The possible negative correlation between the number of species per quadrat and the height and girth of coppice regrowth, as well as time since coppicing, may suggest that increasing shade and associated microclimate factors 28 Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora results in fewer species being able to tolerate the conditions in the majority of the coupe. It is difficult to draw conclusions regarding differences between coupes in the diversity and evenness of species distribution, owing to the lack of correlation with other coupe features. This may be due to combined or conflicting influences on the vegetation, perhaps including factors that were not examined in depth during this study, such as variability of conditions within coupes. The ability of Mercurialis perennis and Hedera helix to take advantage of undisturbed habitats through lateral vegetative spread and shade tolerance (Grime et al., 1990), may explain their dominance in terms of percentage cover in neglected coupes. The greater cover of these species in recently coppiced coupes, but low cover in restored coupes coppiced prior to 1998/99, may be explained by an initial increase in growth following coppicing, followed by a slow decline in response to disturbance (Barkham, 1992b), or competition from species of open habitats. The significantly higher cover of many species in actively coppiced areas may be due to factors such as increased light levels and associated microclimate changes or reduced litter cover. Ability to tolerate microclimate stresses or disturbance during coppicing may also be important. The dominance of Rubus fruticosus , in terms of percentage cover, in every restored coupe is likely to be a response to increased light levels (Barkham, 1992b) and reflects the findings of Fuller and Warren (1993), who suggest that R. fruticosus is likely to develop between coppice stools within three years of coppicing and is likely to persist for up to eight years. This, and its dominance in terms of frequency of occurrence in most restored coupes, also reflects the findings of Van der Werf (1991), who states that it is a species which prefers coppicing to high forest management. The significant tendency of Rubus fruticosus as well as species such as Epilobium hirsutum and Fragaria vesca to occur in restored coupes of all ages may be an indication of their competitive natures and ability to tolerate open habitats as well as some shade (Grime et al., 1990). Responsiveness to increased light, or a tendency to colonise disturbed ground, are likely explanations for the tendency of a number of species to occupy only recently coppiced coupes. This may include species that have survived as buried seed in neglected coppice areas (Fuller & Warren, 1993). Subsequent reduction in frequency of occurrence may be due to a variety of factors. Some may be reduced by grazing following the removal of deer fencing, for example, Geum urbanum. which is rare in grazed habitats (Taylor, 1997). Van der Werf (1991) suggests that the species of recently coppiced areas are likely to include those that may not generally belong in forest ecosystems. This may describe Senecio jacobaea, which was found to occur almost exclusively in the most recently coppiced coupe and Epilobium montanum which does not have a long term ability to survive under woodland canopies (Fuller & Warren, 1993). Such species are often excluded in the later stages of the coppice rotation (Ash & Barkham, 1976), perhaps as a result of shade or increasing volumes of leaf litter. Evans and Barkham (1992) suggest that the response time of some perennial species to canopy gaps may be relatively long. Those species that tend to occur mostly in the older restored coupes, coppiced between 1993 and 1997, where the deer fencing has been removed, may therefore be more likely to consist of species of open habitats that are not rapid colonisers. The competitive abilities of certain species may also have been important in allowing them to persist among other light responsive species. Also, the early flowering of some species such as Primula veris, Taraxacum officinale and Viola hirta (Grime et al., 1990) may have enabled them to survive under a developing canopy and compete with species that flower later in the year. Some species may additionally have been favoured by grazing, following the removal of deer fencing, which may have eliminated some competitors. The tendency of Arum maculatum and Mercurialis perennis to occur in both neglected areas and recently coppiced coupes reflects the ability of these species to survive in shaded habitats (Grime et al., 1990), but may also suggest a tolerance of open habitats. However, it may be that they slowly decline in response to disturbance, following an initial reaction to Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora 29 increased light levels (Barkham, 1992b). Alternatively, their shade tolerance may enable them to survive in the shade of light responsive species for a time (Ash & Barkham, 1976), perhaps until they are eliminated through competitive exclusion. Because of the ability of these species to achieve wide ground cover in shaded areas (Barkham, 1992b), when other light responsive species have diminished, and the possibility that they cannot tolerate conditions throughout the coppice rotation, it may be argued that these species would benefit from non-intervention. For some species, such as Taraxacum officinale, the apparent increase in flowering vigour in areas of less shade may be explained by adaptation to open habitats. For species which flower in midsummer, such as Geum urbanum, flowering vigour may be increased by the removal of the woodland canopy. This concurs with the findings of Taylor (1997) that the seed production of G. urbanum was enhanced in the first two to three years following coppicing at Buff Wood in Cambridgeshire. It is recognised that there are various possible influences, which have not been discussed, that may have resulted in vegetation differences between coupes, even prior to coppice reinstatement, such as different soil conditions, propagule availability and influences from adjacent land. Nevertheless, the size of coppice regrowth and leaf litter cover appear to be important influences on the vegetation. Furthermore, the relatively low similarity of ground flora composition found when comparing neglected coupes to restored coupes may suggest that coppicing has had a considerable influence on flora composition. The greater ground flora cover, species richness and number of species per quadrat in certain coupes also appears to coincide with reinstated coppice management, as does the increased cover, frequency of occurrence and flowering vigour of various individual species. The fact that Arum maculatum and Mercurialis perennis were shown to be potentially disadvantaged by coppicing highlights the importance of retaining areas of non- intervention or high forest management in order to protect those species that depend on shaded, moist or undisturbed areas, especially as they are unlikely to re-colonise an area once they are lost from the vegetation (Brown & Oosterhuis, 1981). A number of species, however, have been shown to benefit from the conditions provided during discrete stages of a coppice rotation. This includes adaptable open ground species with good dispersal powers, which are not restricted to woodland habitats, such as Taraxacum officinale (Grime et al., 1990). However, it also includes a number of open woodland species, such as Geum urbanum (Taylor, 1997, Grime et al., 1990) and Teucrium scorodonia (Hutchinson, 1968), some of which were absent in the neglected coppice areas. Because many open woodland species have declined as a result of canopy closure (Van der Werf, 1991), the provision of suitable habitats for such species may justify the reinstatement of coppice management at Eaves Wood, as long as species of woodland shade continue to be protected through the maintenance of areas of non-intervention or high forest management. Vegetation survey prior to coppice reinstatement is, however, very important in order to ensure that the chosen management regime for any site is appropriate for the species present. This can also provide baseline information for monitoring the effects of coppicing. Acknowledgements This work was carried out in partial fulfilment of the regulations governing the award of MSc in Conservation and Recreation Management at the University of Hertfordshire, under the supervision of Dr Stuart Warrington, Department of Environmental Sciences. Dr John Foster of the Department of Environmental Sciences, University of Hertfordshire, is also thanked for his advice and feedback throughout the project. The National Trust, Bank House Farm, Silverdale, Lancashire, in particular Alan Ferguson (Warden-in-Charge), Steve Bradley (Warden) and Peter Simpson (Warden), are thanked for allowing the study to take place at Eaves Wood and for their co-operation throughout the project. 30 Effects of the Reinstatement of Coppice Management on the Vascular Ground Flora References Alexander, K., Lutley, W., Bullard, R and Hearn K. (1984) Biological Survey: Eaves & Waterslack Woods, Silverdale, Lancashire. National Trust Internal Report, Cirencester. Ash, J. E. and Barkham, J. P. (1976) Changes and variability in the field layer of a coppiced woodland in Norfolk, England. Journal of Ecology 64: 697-712. Barkham, J. P. (1992a) The effects of management on the ground flora of ancient woodland, Brigsteer Park Wood, Cumbria. Biological Conservation 60: 167-188. Barkham, J. P. (1992b) The effects of coppicing and neglect on the performance of the perennial ground flora. In: Ecology & Management of Coppice Woodlands (G. P. Buckley ed.): 115-143. Chapman & Hall, London. Brown, A. H. F. and Oosterhuis, L. (1981) The role of buried seed in coppice woods. Biological Conservation 21: 19-38. Evans, M. N. and Barkham, J. P. (1992) Coppicing and natural disturbance in temperate woodlands - a review. In: Ecology & Management of Coppice Woodlands (G. P. Buckley ed.): 79-98. Chapman & Hall, London. Fuller, R. J. and Warren, M. S. (1993) Coppiced Woodlands: Their Management for Wildlife. 2nd. ed. JNCC, Peterborough. Goldsmith, F. B. (1992) Coppicing - a conservation panacea? In: Ecology & Management of Coppice Woodlands (G. P. Buckley ed.): 306-311. Chapman & Hall, London. Grime, J. P. (1979) Plant Strategies and Vegetation Processes. Wiley, Chichester. Grime, J. P. Hodgson, J. G. and Hunt, R. (1990) The Abridged Comparative Plant Ecology. Chapman & Hall, London. Hambler, C. and Speight, M. R. (1995) Biodiversity conservation in Britain; science replacing tradition. British Wildlife 6: 137-147. Hutchinson, T. C. (1968) Biological Flora of the British Isles: Teucrium scorodonia L. Journal of Ecology 56: 901-911. Kent, M. and Coker, P. (1992) Vegetation Description & Analysis - A Practical Approach. Belhaven Press, London. Kirby, K. J. (1988) A Woodland Survey Handbook - Research & Survey in Nature Conservation No. 11. JNCC, Peterborough. Kirby, K. J. (1993) Coppice restoration for nature conservation: how much and where. In: Coppice Restoration Seminar Proceedings (R. Lightbown & A. Searle, ed.): 15-24. Institute of Chartered Foresters. Kirby, K. J. (1996) Conservation of habitats. In: Conservation Biology (I. F. Spellerberg (ed.): 141-153. Longman, Essex. Mitchell, P. L. and Kirby, K. J. (1989) Ecological Effects of Forestry Practices in Long-established Woodland and Their Implications for Nature Conservation. Occasional Paper No. 39, Oxford Forestry Institute, Oxford. National Trust, (c. 1993) Eaves Wood, Silverdale: Management Plan. National Trust Internal Report, Silverdale. Peterken, G. F. (1992) Coppices in the lowland landscape. In: Ecology & Management of Coppice Woodlands (G. P Buckley ed.): 3-17. Chapman & Hall, London. Powell, S. (1996) Statistics for Science Projects. Hodder & Stoughton, London. Taylor, K. (1997) Biological Flora of the British Isles: Geum urbanum. Journal of Ecology 85: 705-720. Van der Werf, S. (1991) The influence of coppicing on vegetation. Vegetatio 92: 97-110. 31 THE WASPS AND BEES (. HYMENOPTERA : ACULEATA ) OF CORNELIAN AND CAYTON BAYS AND OSGODBY POINT IN WATSONIAN YORKSHIRE MICHAEL E. ARCHER 17 Elmfield Terrace, Malton Road, York Y031 1EH Aculeate wasps and bees require open terrestrial habitats. The open landscape that we now know is largely a result of human activity. The habitats originally used by the aculeate wasps and bees would mainly be disturbed areas at the coast (e.g. sand dunes, erosion of hard rock cliffs and soft rock cliffs and slips) or inland (e.g. river banks and inland sand dunes). Cornelian and Cayton Bays are areas subject to frequent landslips and Osgodby Point hard rock cliffs are being eroded by the sea. Thus these three sites could be modern- day representatives of the original disturbed habitats. The three sites are situated on the coast just to the south of Scarborough (TA0685) and are made up of boulder clay cliffs and a sandstone headland. The sites are in the ownership of the National Trust and the local authority. They are situated in the North York Moors and Hills Natural Area. The study area was about 23 hectares, but probably covered about 39 hectares when others’ records are considered. The main habitats within these sites are woodland, dense shrub, flowery basic enriched grassland and marsh. There is abundant dead wood, which in sunny situations is used by aerial nesters. The dry friable bare banks, particularly where the soft sandstone comes to the surface in sunny situations, are used by subterranean nesters. Pollen and nectar sources are readily available from the flowers of trees (sycamore), shrubs (sallow, blackthorn, hawthorn) and herbs (dandelion, kidney vetch). The wide variety of habitats provide the prey, needed by wasps. The aims of this paper are to describe the aculeate wasp and bee fauna of the three sites, to estimate the potential number of species that could be present, carry out comparisons with other Yorkshire sites and evaluate the conservation value of the sites. The three sites, for the purposes of analysis, will be treated as one site. Methods Between 1984 and 2000, 16 visits were made to the three sites, distributed throughout the year as follows: April (2 visits), May (3), June (3), July (3), August (4) and September (1). During these visits, which usually lasted about three hours, all species of aculeate wasps and bees were recorded and usually collected with a hand net for identification. Some records have been made available by the following: J. D. Coldwell (Cayton Bay, July 1995), W. A. Ely (Cornelian Bay, May 1976). J. H. Flint (Cayton Bay, June 1986, five visits August 1965-1971), A. Grayson (Cornelian Bay, June 1995), P. Kendall (Cayton Bay, July 1983), T. H. Riley (Cornelian Bay, June 1973) and D. H. Smith (Cornelian Bay, July 1987). The additional records added two species not found by the author. In the following account, the nomenclature can be related to that of Fitton et al. (1978). Species present and seasonal progression of species A full list of recorded species is given in the Appendix. At the family level, Table 1 shows the taxonomic distribution of species and records. The Archer sample consists of 13 social wasp and bee species and 195 records of 57 solitary species. A record, in the Archer sample, represents a specimen differing in one of the following three variables: name, sex and day of visit. The solitary wasp family, Sphecidae, and the solitary bee families, Andrenidae and Halictidae, are the dominant families in terms of both the number of species and records. Table 2 shows the number of solitary species and when species were first recorded for each month of the Archer sample. The most productive months for the species of solitary wasps were June and July with most species being first recorded during June. No solitary Naturalist 127 (2002) 32 Wasps and Bees (Hymenoptera: Aculeata) in Watsonian Yorkshire wasps were found during the spring months (April to May). The species most evident are the aerial nesters ( Trypoxylon figulus, T. clavicerum, Ectemnius cavifrons ) usually flying around and landing on upright dead wood in sunny situations, and the subterranean nesters (. Priocnemis schioedtei, Odynerus spinipes, Diodontus tristis, Mellinus arvensis). A O. spinipes nesting aggregation was found on a boulder clay bare bank facing south, usually with its parasite, Chrysis viridula, in attendance. Males and females of P. schioedtei were found flying and running over bare exposed boulder clay while males and females M. arvensis were associated with sandy outcrops. TABLE 1 The number of records and species of the Archer sample and total number of aculeate wasps and bees recorded from Cornelian and Cayton Bays with Osgodby Point. Archer Records Archer Species Total Species Solitary Wasps Chrysididae 13 5 5 Pompilidae 17 7 8 Eumenidae 8 3 3 Sphecidae 89 19 20 Total Solitary Wasps 127 34 36 Solitary Bees Andrenidae 23 7 7 Halictidae 37 11 11 Megachilidae 3 3 3 Anthophoridae 5 2 2 Total Solitary Bees 68 23 23 Total Solitary Species 195 57 59 Social Wasps & Bees Vespidae 4 4 Apidae 9 9 Total Social Species 13 13 Total Wasps and Bees 70 72 TABLE 2 The number of species and when species were first recorded per month of solitary wasps and bees from the Archer sample for Cornelian and Cayton Bays with Osgodby Point. April May June July August September No. species Wasps 0 0 23 22 16 5 Bees 6 9 9 9 9 5 No. species first recorded Wasps 0 0 23 7 4 0 Bees 6 4 6 3 3 1 The most productive months for the solitary bees were May, June, July and August with most species being first recorded during April and June. The species most evident were the spring species ( Andrena scotica with its cleptoparasite Nomada marshamella, A. haemorrhoa ) and the species that appear throughout the spring and summer ( Halictus Wasps and Bees (Hymenoptera: Aculeata) in Watsonian Yorkshire 33 rubicundus, Lasioglossum f rate llum, L. rufitarse, L. villosulum). All these bees are subterranean nesters. Estimating the potential number of solitary and bee species Twenty-one of the Archer sample of 57 solitary species were only recorded on one occasion, giving rise to concern as to how many species had not been found. Recent advances in non-parametric statistical procedures offer some hope of addressing this problem. The presence/absence quantitative estimate of Chao (in Colwell & Coddington, 1994) is based on the number of species that are recorded on one (unique species) or two (two occasion species) samples or visits. The jackknife procedure (Heltshe & Forrester, 1983) only depends on the unique species. The software to carry out the statistical procedures is provided by Pisces Conservation Ltd. The statistical procedure takes 1, 2, etc. samples at random 16 times (i.e. the number of samples), each time calculating a mean estimate of the potential number of species with a measure of the error of the estimate. With a small number of samples the estimates are erratic, but as more samples are selected the estimates may stabilise giving confidence in them. Since the estimates for the Chao (Fig. 1) and Jackknife (Fig. 2) procedures more-or-less stabilise and the final estimates based on all 16 samples for the two procedures (Table 3) are very similar, there is confidence that there are on an average nearly 80 potential species of solitary wasps and bees for the three sites. Just over 70% of these potential number of species have been found by the author. Another 20-22 species remain to be found. TABLE 3 Non-parametric estimates of species richness of solitary wasps and bees based on the Archer sample from Cornelian and Cayton Bays with Osgodby Point. Chao estimate Jackknife estimate No. species - recorded 57 57 - estimated 79 77 95% confidence limits 57-100 67-87 % species recorded of estimate 72.2 74.0 Comparisons with other Yorkshire sites Another problem in the study of any site is the difficulty of knowing when the species list is sufficiently complete so that comparisons with other sites may reasonably be carried out. One way to resolve this problem is the use of the species-area relationship, where the number of species and the area of sites, both expressed as natural logarithms (In), can show a positive relationship (Usher, 1986). If the number of species in relation to the area of a site falls within the range of other sites which show a statistically significant species-area relationship, then the site may reasonably be compared with other sites. If the number of species in relation to the area of the site falls below the values of the other sites than this could indicate either more species could be found at that site, or that the site consists of habitats which are particularly unfavourable for aculeate wasps and bees. The dot for the current site (either the smaller area of the Archer sample or the larger area of all recorders) falls within the range of 19 sites from the north and north midlands of England (Archer, 1999) and so the species list from the current study site can be considered sufficiently complete to make valid comparisons with other sites. Cleptoparasitic load The cleptoparasitic load (CL) is the percentage of aculeate species that are cleptoparasites (or parasitoids) on other host aculeates. Wcislo (1987) showed that parasite behaviour among aculeate Hymenoptera correlated with geographical latitude. Thus the parasitic rates are higher in temperate regions as host populations are more Species Richness Estimates Species Richness Estimates 34 Wasps and Bees (Hymenoptera: Aculeataj in Watsonian Yorkshire 100 80 60 40 20 0 0 2 4 6 8 10 12 14 16 No. Samples FIGURE 1 rhe Chao presence/absence estimate of species for Cornelian and Cayton Bays with Osgodby Point. FIGURE 2 The Jacknife estimate of species for Cornelian and Cayton Bays with Osgodby Point. 35 Wasps and Bees (Hymenoptera: Aculeata) in Watsonian Yorkshire synchronised in their life-history characteristics. This finding probably does not hold for desert regions where the occurrence of rainfall would tend to synchronise life history characteristics. From a review of the literature Wcislo found that the CLs for bees in Europe varied between 16% and 33%, with a range of 17%. Archer (1999) found that the CLs for the solitary bees from north and north midlands sites of England vary between 22% and 37%, with a range of 15% which is similar to the range found by Wcislo. The CL for the solitary bees from the current study site (Table 4) falls within this range. Wcislo gives no CLs for solitary wasps but Archer (1999) found that the north and north midlands of England CLs vary between 10% and 22%, with a range of 12%. The narrow range of this variation indicates that the argument Wcislo developed for the bees also applies to the solitary wasps. The CL for the solitary wasps from the current site (Table 4) falls within this range. Archer and Burn (1995) discussed why the CLs for the solitary bees are higher than the CLs for the solitary wasps. They argue that it is probably a consequence of food-chain relationships. All the social species are host species, except for the Psithyrus species, which are social parasites on the Bombus species. TABLE 4 The relative frequency of the cleptoparasitic (or parasitoid) species among the solitary species from Cornelian and Cayton Bays with Osgodby Point. No. hosts No. cleptoparasites Cleptoparasitic Load (H) (C) CL = 100 x C/(H+C) Solitary Wasps 31 5 13.9 Solitary Bees 17 6 26.1 Aerial nester frequency The aerial nester frequency (AF) is the percentage of host aculeate species that have aerial nest sites. The aerial nesters used old beetle burrows in dead wood, hollow stems, e.g. bramble, crevices in mortar walls or old snail shells. Subterranean nesters nest in the soil, usually in burrows dug by themselves, but sometimes holes and crevices are used after being altered. Archer (1999) found that the AFs for the solitary wasps from the north and north midlands vary widely between 0% and 84% while the AF for British species is 46%. The AF of the solitary wasps from the current study site (Table 5) falls within this range but is higher than the British percentage. Archer (1999) found that the AFs for the solitary bees from the north and north midlands of England vary less widely - between 7% and 23% - while the British percentage is 18%. The AF of the solitary bees for the current study site (Table 5) falls within this range but is lower than the British percentage. In summary the current study site is relatively rich in aerial nesters for solitary wasps but relatively poor in aerial nesters for the solitary bees. Since the study site is rich in aerial nesting sites the small number of aerial nesting solitary bees was not expected. Perhaps more aerial nesting solitary bee species will be found with more sampling, or maybe their habitat requirements are not all present for their continued existence. Of the species of social wasps Vespula rufa and Paravespula vulgaris are mainly subterranean nesters and Dolichovespula sylvestris and D. norwegica are mainly aerial nesters. A nest of the social wasp, Dolichovespula sylvestris, was found in a hollow of the boulder clay cliffs. Males of the social wasp, Vespula rufa, were once found in a mating swarm on the tops of the shrubs. The host species of the bumble bees are mainly or entirely subterranean nesters, although Bombus pratorum has been found in aerial situations such as old bird’s nests. 36 Wasps and Bees (Hymenoptera: Aculeata) in Watsonian Yorkshire TABLE 5 The nesting habits of the host solitary species recorded from Cornelian and Cayton Bays with Osgodby Point No. aerial No. subterranean Aerial nester frequency nesters (A) nesters (S) AF = 100 x A/(A+S) Solitary Wasps 17 14 54.8 Solitary Bees 2 15 11.8 The conservation value of the site Two Yorkshire rarities have been found: Priocnemis cordivalvata which is at the northern edge of its range in Yorkshire and Coelioxys rufescens which is found in England, Wales and southern Scotland. Four species are nationally rare or scarce according to Shirt (1987) and Falk (1991): Priocnemis cordivalvata, Ectemnius ruficornis and E. sexcinctus which are at the northern edge of their ranges in Yorkshire, and Priocnemis schioedtei which is found in England, Wales and Scotland. Recent work by the Bees, Wasps and Ants Recording Society indicates that the statuses of P. schioedtei, E. ruficornis and E. sexcinctus should be downgraded. Within a national context these three species have an Archer Widespread status (Archer, 1999), and within a Yorkshire context a Frequent Status (Archer, 1993). Giving each solitary species a regional status, a regional quality score and species quality score (RSQS) (Archer, 1993) can be calculated (Table 6). Four species are given a national Archer Scarce status: Chrysis viridula, Priocnemis cordivalvata , Anoplius concinnus and Diodontus tristis. The RSQS for the study site at 2.6 is similar to those calculated for Allerthorpe Common (2.7) and Keswick Fitts (2.7) so qualifying the study site as important in a Yorkshire context. TABLE 6 The regional and Archer national quality scores of the solitary wasps and bees from Cornelian and Cayton Bays with Osgodby Point. Scores Status value (A) No. species (B) Quality (A x B) Regional Status Common 1 36 36 Frequent 2 12 24 Occasional 4 7 28 Rare 8 0 0 Nationally scarce 16 4 64 Total (Quality Score) 59 Species Quality Score 152/59 = 2.6 152 National Status Universal 1 39 39 Widespread 2 16 32 Restricted 4 0 0 Scarce 8 4 32 Total (Quality Score) 59 Species Quality Score 103/59 = 1.7 103 37 Wasps and Bees (Hymenoptera: Aculeata) in Watsonian Yorkshire Giving each species a national status, a national quality score and species quality score (NSQS) (Archer, 1999) can be calculated (Table 6). The NSQS for the study site at 1.7 is similar to those calculated for Skipwith Common (1.6) and Burton Leonard Lime Quarries (1.6). From a national perspective these sites could be considered to form a second tier of Yorkshire sites. Examples of Yorkshire first tier sites would be Strensall Common (2.5), Duncombe Park (2.2) and Crow Wood (2.5). Conclusions • The number of aculeate species recorded from Cornelian and Cay ton Bays with Osgodby Point indicate that this is a good site. • The estimates of the potential number of species that could be present are stable indicating that, on average, a further 20-22 species could be found. • The site has the expected number of solitary species for its area, and so it can properly be compared with other sites. • The site has similar cleptopasitic load to that of other sites as predicted by Wcislo (1987). • The site has a higher than average aerial nester frequency (AF) for the solitary wasps but a lower than average AF for the solitary bees. • The conservation value of the study site is important both in a national and regional context. References Archer, M. E. (1993) Recorder’s fourth report on the aculeate Hymenoptera in Watsonian Yorkshire and the development of a quality scoring system. Naturalist 118: 13-15. Archer, M. E. (1999) The aculeate wasps and bees (Hymenoptera: Aculeata) of the Ainsdale Formby sand dunes on the Lancashire coast compared with other northern sites. Br. J. Ent. Nat. Hist. 12: 1-10. Archer, M. E. and Burn, J. T. (1995) The aculeate wasps and bees of Crow Wood, Finningley in Watsonian Yorkshire, with the introduction of a new national quality system. Br. J. Ent. Nat. Hist. 8: 49-59. Colwell, R. K. and Coddington, J. A. (1994) Estimating terrestrial biodiversity through extrapolation. Phil. Trans. R. Soc. Lond. B. 345: 101-118. Falk, S. (1991) A review of the scarce and threatened bees, wasps and ants of Great Britain. Research and Survey in Nature Conservation No. 35. Nature Conservancy Council, Peterborough. Fitton, M. G. et al. (1978) In Kloet G. S.and Hincks, W. D. (eds.), A check list of British Insects. Part 4: Hymenoptera. Handbk Ident. Br. Insects 11: (4). Heltshe, J. F. and Forrester, N. E. (1983) Estimating species richness using the Jackknife Procedure. Biometrics 39: 1-11. Shirt, D. B. (ed.) (1987) British Red Data Books: 2. Insects. Nature Conservancy Council, Peterborough. Usher, M. B. (1986) Wildlife Conservation Evaluation. Chapman and Hall, London. Wcislo, W. T. (1987) The role of seasonality, host synchrony, and behaviour in the evolutions and distributions of nest parasites in Hymenoptera (Insecta), with special reference to bees (Apoidea). Biol. Rev. 62: 515-543. Appendix Chrysididae. Chrysis angustula Schenck, C. impressa Schenck, C. ruddii Shuckard, C. viridula Linn., Trichrysis cyanea (Linn.). Pompilidae. Dipogon subintermedius (Magretti) (= nitidas ), D. variegatus (Linn.), Priocnemis cordivalvata Haupt, P. fennica Haupt, P. parvula Dahlbom, P. schioedtei Haupt, Arachnospila anceps (Wesmael), Anoplius concinnus (Dahlbom). Eumenidae. Odynerus spinipes (Linn.), Ancistrocerus parietum (Linn.), A. scoticus (Curtis). Book Reviews 38 Vespidae. Dolichovespula norwegica (Fab.), D. sylvestris (Scopoli), Vespula rufa (Linn.), Paravespula vulgaris (Linn.). Sphecidae. Trypoxylon clavicerum Lepeletier, T. figulus (Linn.), Crossocerus annulipes (Lepeletier & Bridle), C. dimidiatus (Fab.), C. elongatulus (Vander Linden), C. megcephalus (Rossius), C. podagricus (Vander Linden), C. pusillus Lepeletier & Bridle (= varus), C. quadrimaculatus (Fab.), C. tarsatus (Shuckard), Ectemnius cavifrons (Thomson), E. ruficornis (Zetterstedt), E. sexcinctus (Fab.), Psenulus pallipes (Panzer), Pemphredon inornatus Say, P. lugubris (Fab.), Diodontus tristis (Vander Linden), Passaloecus corniger Shuckard, Mellinus arvensis (Linn.), Argogorytes mystaceus (Linn.). Andrenidae. Andrena chrysosceles (Kirby), A. clarkella (Kirby), A. fucata Smith, A. fulva (Muller in Allioni), A. haemorrhoa (Fab.), A. nigroaenea (Kirby), A. scotica Perkins. Halictidae. Halictus rubicundus (Christ), Lasioglossum cupromicans (Perez), L. fratellum (Perez), L. nitiusculum (Kirby), L. punctatissimum (Schenck), L. rufitarse (Zetterstedt), L. smeathmanellum (Kirby), L. villosulum (Kirby), Sphecodes geojfrellus (Kirby) (= fasciatus ), S. gibbus (Linn.), S. monilicornis (Kirby). Megachilidae, Osmia caerulescens (Linn.), O. rufa (Linn.), Coelioxys rufescens Lepeletier & Serville. Anthophoridae. Nomada goodeniana (Kirby), N. marshamella (Kirby). Apidae. Bombus hortorum (Linn.), B. lapidarius (Linn.), B. lucorum (Linn.), B. pascuorum (Scopoli), B. pratorum (Linn.), B. terrestris (Linn.), Psithyrus bohemicus (Seidl), P. vestalis (Geoffrey in Fourcroy), Apis mellifera Linn. BOOK REVIEWS Maggots, Murder and Men. Memories and Reflections of a Forensic Entomologist by Zakaria Erzinclioglu. Pp. 256. Harley Books, Great Horkesley, Colchester. 2000. £13.95 paperback. The author, known to many naturalists as Zak, is widely respected not only for his forensic work but also for his contribution to entomology in general. As he states, forensic science is indeed “often very exciting, even exhilarating” - and this work proves just that. The author demonstrates through first-hand knowledge how the facts are gathered together during a criminal investigation and are transformed into the solution of the crime, stressing how “there is no substitute for deep thought, hard work and imagination”. Although the author exemplifies his own work by many case studies, these are somewhat repetitive in terms of their underlying principles, namely where and when was the crime perpetrated; sometimes, however, such work can provide details of the circumstances of the death and indeed can occasionally point the finger at a particular individual. Nevertheless, the book contains a rich variety of fascinating cases, not only those personally investigated by the author, but also those explaining the deaths of important historical figures. As well as providing a most readable and entertaining book, to be dipped into rather than read at one sitting, the author shows his undoubted enthusiasm for the subject and his respect for the moral, legal and social implications of such sensitive work. MRDS Photographic Guide to the Butterflies of Britain & Europe by Tom Tolman. Pp. 320, 500+ colour photos, & colour distribution maps for each species. Oxford University Press, 2001. £35.00 hardback, £16.50 paperback. This is a most attractive book with full colour throughout, and up to five colour photos to the page, plus very clear colour distribution maps covering the whole of Europe. Text and Book Reviews 39 illustrations are usefully placed together, with brief notes on distribution, description, flight period, habitat and sometimes conservation. Unfortunately, as far as Great Britain is concerned, the maps are not very accurate. Whilst it is appreciated that the relatively small size limits the detail which can be included, there seems little excuse for placing the northern limit of the Brimstone across the middle of Scotland, when it lies across northern Yorkshire! The odd error can be forgiven, but these maps contain many more, including the placing of Brown Hairstreak well into Yorkshire when it has never ever been recorded there. In addition the British element of the maps fails to take into account the many recent changes in butterfly distribution. The information appears to have been based on data collected 30 to 40 years ago. However, the book is worth buying just for the huge collection of over 500 photographs. Many of them are quite stunning and some show extremely rare species. A few are so dark as to be almost useless in aiding identification. It is good that the country in which each photo was taken is clearly stated, as most species vary geographically in some degree. However, this does reveal that many of the pictures were taken in Greece (including the Small Tortoiseshell), which does not necessarily give the most typical colouration for those species which are widespread. In spite of some shortcomings this is a valuable reference book. There are so many species to contend with in Europe and so few books that attempt to cover them all. This new guide would provide an excellent companion to the Collins Field Guide to the Butterflies of Britain and Europe by Tolman & Lewington (HarperCollins 1997), which uses paintings to illustrate the species. You could get the best of both worlds by comparing the photographs of the one with the paintings of the other. HMF The Butterflies of Pakistan by T. J. Roberts. Pp. 200 (large format c.20 x 28 cm), 47 colour plates, 9 figures, Oxford University Press, 2001, £9.99 hardback. This is something of a landmark publication, being the first ever attempt to list and describe all 320 species of butterfly found in Pakistan. As such it will be of special interest to any UK naturalist visiting that part of the world as well as anyone wishing to be better informed about the world of butterflies in general. This book includes introductory chapters covering life cycles, habitats and other general topics, including a brief, but very interesting, section on migration. The bulk of the book is taken up with species coverage and colour plates painted by the author. Unfortunately, the paintings are not to scale, but they are extremely detailed and possess a certain charm in their unsophisticated style. The information for each species includes a description, and notes on status. An additional and extremely interesting section on habitats is included where the information is available. There are many seemingly unfamiliar species listed here, like the Sorrel Sapphire Heliophorus sena, the Common Crow Euploea core, and the Blue Pansy Junonia orithya, yet when you look at the pictures you realise that most of the family groups are totally familiar even though many of the species carry strange sounding names. So you have the Clouded Yellows, the Brimstones, the Fritillaries, the Vanessids, the Browns and the Blues. The Skippers include the Awls and the Flats, the Hoppers and the Darts. You may not have heard of these before, but the plates reveal that they simply extend the rather confusing array of similar-looking Skipper butterflies we have in Europe. Pakistan is also host to many of our familiar butterflies like the three common Whites, the Brimstone Gonepteryx rhamni, the Red Admiral Vanessa atalanta and so on. It is interesting to compare their habits in Pakistan with those at home. The Farge White Pieris brassicae for instance breeds on the northern plains in the coolest months and the offspring migrate into the Himalayas to produce successive generations in spring and summer. 40 Book Reviews At £9.99 this substantial hardback book is an exceptional bargain and deserves a place in every lepidopterist‘s library! HMF Biological Collections and Biodiversity edited by B. S. Rushton, P. Hackney and C. R. Tyre. Pp. x + 326, inch numerous line drawing & b/w plates. Westbury Publishing, Otley, for the Linnean Society of London, 2001. £35.00 hardback. This long-awaited volume, based on an Annual Regional Conference of the Linnean Society held in Belfast in August 1996, is a curious mix of papers, some rather dusty now in view of the lengthy gestation process and some on the fringe of or indeed outside the topic under review, a classic example of the latter being (to give it its full title) a “Checklist of marine phytoplankton in a subtropical lagoon system in Baja California Sur, Mexico, from 1980 to 1989”. Although one recognises the role that biological collections play in cataloguing and preserving biodiversity, in the reviewer’s opinion ‘Biological Collections’ and ‘Biodiversity’ do not fit comfortably within this disparate collection of 33 (often short) papers and 32 abstracts of other papers and posters. As a participant in the conference, the reviewer can testify to its success in terms of enlightenment and enjoyment. The editors cannot be criticised for the resulting publication - they have made a valiant effort to portray it, but the wide-ranging topics covered fail to provide the in-depth treatment this subject requires, and the contents reflect only the viewpoints of those authors who chose to contribute. There is an underlying feeling of disheartenment conveyed through past and present devaluation of biological collections. Despite these criticisms, the book contains a considerable amount of useful and interesting information, the contribution on “The monetary value of herbarium collections” being particularly thought-provoking, and the publishers are to be congratulated on the excellence of the finished project. MRDS Small Freshwater Creatures by Lars-Henrick Olsen, Jacob Sunesen and Bente Vida Pederson. Pp. 229 Oxford University Press. 2001. £12.50 hardback. This book, a translation of a work in Danish, aims to introduce freshwater animals - in spite of the title not always small - to beginners, and is not designed specifically for the needs of British users. The front and back end papers portray silhouettes of adult and larval stages respectively of a wide range of animals and indicate the pages on which they are described, which is a good idea. More than 500 species, some identified only to genus, are illustrated and briefly, and not always accurately, described in what is taxonomically a somewhat curious sequence. Illustrations of large forms, such as dragonflies, are often attractive and helpful, but those of microscopic animals, with which describer and illustrator are obviously unfamiliar, are often crude and sometimes misleading. Many of the animals included are not found in Britain, and in some groups most, or even none, of the species shown occurs here. There is a miscellany of general topics (18 pages) at the end and a short glossary. The last entry begins “Zooplankton are very small animals ...” thus perpetuating a grammatical horror apparently of trans-Atlantic origin. Zooplankton is a collective term for small open- water animals and is singular. While its ease of use and generally attractive illustrations may whet the appetite of beginners, the shortcomings of this work are such that one can give it only a muted welcome. GF BIRDS ON THE SPURN PENINSULA by Ralph Chislett Parts I and II (1996), edited by Michael Densley. Hardback 218 pages with coloured dust jacket, coloured frontispiece and twelve photographic illustrations in black and white. This enlarged edition is the first time that Part II has appeared in print. Not surprisingly this book virtually sold-out remarkably quickly, and only recently the printers have discovered a small quantity of the remaining books. Before releasing them on the open market, the publisher has offered this limited stock to YNU members for the same price as when it was published in 1996 - £14.95 per copy post free. Owing to a slight mix-up at the time, the newly released book has not previously been advertised in The Naturalist - thus members now have a chance to purchase a copy. The James Reckitt Charity of Hull and other individual benefactors have made a substantial contribution towards the publishing cost in order to keep the selling price relatively low. Apply to: Clive Varty, 26 Craggwood Road, Horsforth, Leeds LSI 8 4PB. Tel (0113) 258 7627 THE HARROGATE AND DISTRICT NATURALISTS’ SOCIETY A NEW BOOK on the BIRDS OF THE HARROGATE DISTRICT by JOHN R. MATHER MBOU A complete historical review including records from the 18th century to the present time. 3 1 2 pages, many line drawings, fully referenced, with a foreword by D. I. M. Wallace. Published October 2001. Price £10.00 plus £2 postage and packing. Please order your copy from: Mrs J. McClean, 6 Rossett Park Road, Harrogate, North Yorkshire HG2 9NP YNU PUBLICATIONS FOR SALE A Fungus Flora of Yorkshire. 1985. 296 pp. Hardback. £10.00 incl. p&p. Butterflies and Moths of Yorkshire. 1 989. 380 pp. Softback or unbound. £1 2.50 incl. p&p. Mammals of Yorkshire. 1985. 256 pp. £7.00 incl. p&p. Provisional Keys to British Plant Calls. 1 986. 85pp. £5.50 incl. p&p. Moths and Butterflies of Spurn, 1 995. 1 24 pp. £6 incl. p&p. Cheques should be made payable to Y.N.U. From: MrsJ. Payne, 15 Broad Lane, Cawood, Selby, North Yorkshire, Y08 3SQ Telephone: 01 757 268242 Printed in Great Britain by Titus Wilson & Son, Kendal ISSN 0028-0771 Latest publication of the Yorkshire Naturalists’ Union ALJEN PLANTS OF YORKSHIRE GEOFFREY WILMORE The first comprehensive modem publication dealing with the alien plants of any major British county, detailing the flora (almost 1300 species) of Yorkshire’s wool waste dumps, railway sidings, sewage works, shoddy fields, industrial wasteland, dockland and oil mill sidings. Pp. 316, including 15 line drawings; published January 2000 Normal price: £15.00 (plus £2.00 per copy p.&p.) Reduced price for members of the YNU: £12.00 (plus £2.00 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Mr John A. Newbold, Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS THE FRESHWATER CRUSTACEA OF YORKSHIRE a faunistic & ecological survey GEOFFREY FRYER The crustacean fauna of Yorkshire reflects the great physiographic diversity of the region. Adopting an ecological approach, this book considers the Yorkshire fauna in relation to climate, topography, geology, soils and water chemistry, always keeping in mind that it is dealing with living organisms whose habits, requirements and physiological limitations determine exactly where they live. Matters covered include the ecological background; faunal assemblages and their regional attributes; an analysis of the factors that determine distribution patterns, many of which are mapped; wide geographical aspects; and conservation. Large areas, such as the Pennines, Howgill Fells, North Eastern uplands and the lowland plains are surveyed. So too are localised regions including Whernside, the Malham area, lowland heaths, and the largest lakes, as well as habitats such as upland tarns, seepages, cold springs, small lowland ponds, inland saline waters. Notes are given on every species recorded, including parasitic forms. Price £8.00 (plus £2.00 p.&p.) Special offer to members of the Yorkshire Naturalists’ Union £6.00 (plus £2.00 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Professor M. R. D. Seaward, Department of Environ- mental Science, University of Bradford, Bradford BD7 1DP. A QUARTERLY JOURNAL OL NATURAL HISTORY LOR THE NORTH OL ENGLAND Red in Tooth and Claw: 1. Studies on the History of the Wild Cat Felis silvestris Schr. in Yorkshire - Colin A. Howes Aquatic Plants at North Cave Wetlands, a New Yorkshire Wildlife Trust Reserve Comprising Former Gravel Workings - R. Goulder Distribution of the Banded Demoiselle Calopteryx splendens along the River Derwent, North Yorkshire - Richard Sunter Coleoptera Report for 1995-2001 - R. J. Marsh Published by the Yorkshire Naturalists’ Union Editor M. R. D. Seaward MSc, PhD, DSc, FLS, The University, Bradford BD7 1DP Notice to Contributors to ‘The Naturalist’ Manuscripts (two copies if possible), typed double-spaced on one side of the paper only with margins at top and left-hand at least 2.5cm wide, should be submitted. Latin names of genera and species, but nothing else, should be underlined. S.I. Units should be used wherever possible. Authors must ensure that their references are accurately cited, and that the titles of the journals are correctly abbreviated. Volumes of The Naturalist for the years 1886 to 1975 have been retrospectively numbered 11 to 100 to accord with numbering before and after this period (see YNU Bulletin no. 3, pp. 21-22 1985); please cite these volume numbers in all references. 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The editor, on behalf of the YNU, wishes to record this deep appreciation of this imaginative gesture. © Yorkshire Naturalists’ Union — 2002 Single copies may be made of single articles in this journal provided that due acknow- ledgement is made and the copies are for non-profit making educational or private use. Copying of more than one article or multiple copying of a single article is forbidden unless special permission has been obtained from the Yorkshire Naturalists’ Union. Permission is granted for the use of brief quotations in published work provided that acknowledgement of the source is clearly stated, but the use of substantial sections of text and any illustrative matter requires the express permission of the Yorkshire Naturalists’ Union. Registered Office (for all enquiries, correspondence and payments): Mr John A. Newbould, Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS (tel: 01305-837384; email: janewbould@aol.com) The Naturalist is issued free to individual members of the Yorkshire Naturalists’ Union and to Affiliated Societies. Institutions and Subscribers £24.00 Registered Charity No. 224018 41 RED IN TOOTH AND CLAW: 1. STUDIES ON THE HISTORY OF THE WILD CAT FELIS SILVESTRIS SCHR. IN YORKSHIRE COLIN A. HOWES Doncaster Museum & Art Gallery, Doncaster DN1 2AE Part 1 of the Presidential Address presented to the Yorkshire Naturalists’ Union at Doncaster, 2nd December 2000 Preface Inspirational teachers can have indelible effects on their charges leading to unexpected though rewarding quests in life. One such, Hilda Smith of Scarborough, would regale our year with fascinating stories which included the gory legend of a wild cat that savagely attacked a Knight Templar. Through the writings of Ted Hughes (1960) in his poem Esther’s Tomcat the saga became particularly memorable: A Tomcat sprang at a mounted knight Locked round his neck like a trap of hooks While the knight rode fighting its claws and bite . . . A later mentor, Mike Clegg, then assistant curator at Scarborough’s Wood End Natural History Museum, also revealed himself to be familiar with T’ Cat ‘n ’ Man legend. Decades later, on coming to live and work in South Yorkshire, I encountered the story on its native soil, for here the legend, Hilda Smith, Ted Hughes and Mike Clegg had their origins. The numerous tellings and transmogrifications of the story (over 40 published examples collected to date) seem to spring from two root versions, a perfunctory form collected from the vernacular by the naturalist the Rev. William Bingley, published in his Animal Biography of 1802, and a lengthy ‘gothic horror’ version fabricated by the historian and journalist C. W. Hatfield in his Village Sketches published by the Doncaster Gazette in 1849. The gist of the tale was that in 1455, Sir Percival Cressacre of Barnbrough Hall, returning on horseback from an encounter with the Franciscan Grey Friars of Doncaster, was attacked by a wild cat, which issued forth from the local woodland. A running battle ensued, culminating in Sir Percival taking refuge in the porch of St. Peter’s Church, collapsing through loss of blood onto the cat; thus t’man killed t’cat and t’cat killed t’man (Twigg 1879). The physical evidence on which the legend has its basis are precised in the following rhyming couplets (Bingley 1802): And still the tessellated floor Shows traces of the purple gore, Of both the baron and his foe - At least tradition says ’tis so; And on his marble tomb displayed, Full length in effigy is laid; While at his feet, lies large as life, The cat which caused the mortal strife. The ‘ purple gore’ is the iron oxide in the stonework of the porch in St. Peter’s Church, Barnburgh. The ‘ cat which caused the mortal strife’ is actually a dog at the foot of the remarkable medieval life-size wooden ‘ effigy ’ of a knight in armour, situated in the Cressacre tomb within the church. The Cressacre arms (on the exterior of the church tower) also feature a ‘cat-a-mount’ (cat of the mountains) as an heraldic device. Puzzlingly, the truly fascinating folkloric interpretations of the saga are not included in any compendium of English folk tales and knowledge of the tale is largely confined to Barnbrough (‘t cat ‘n’ man town) and the Dearne valley pit village communities. The faunal elements of the legend, which include wolves, owls and of course the wild cat, were rationalised by Victorian historians and naturalists such as James (1865), Hatfield (1849) and Harting (1890), who refer to the ‘hercinian’ forests which in unspecified times past Naturalist 127 (2002) 42 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat were deemed to have clothed Pennine South Yorkshire and which were evidently populated by ravening beasts exemplified by wild cats. Imbued with this early interest in cats ancient and modern, beasts which had largely eluded historians, had confused archaeologists and had been ignored by naturalists, a quest was undertaken to search for further evidence of the wild cats (Red in Tooth and Claw: part 1) and ultimately to investigate the ‘wildlife’ of their domestic conterparts in Yorkshire (Red in Tooth and Claw: part 2). Historical evidence of the Wild Cat in Yorkshire Despite an extensive and disparate body of literature from Quaternary, archaeological, topographical, local history and folklore studies, little is actually known of the history of the wild cat Felis silvestris Schr. or the introduced con-specific domestic cat Felis catus Linn, in Yorkshire (Howes 1984, 1985). This study seeks to assemble and review these sources and to add a substantial body of new research from place name and ecclesiastical archives, to refine our sketchy knowledge of its former distribution and decline. Material from excavations at the Mesolithic site at Thatcham, Berkshire, carbon dated 10,050-9600b.p., revealed that even at this early date a modern warm temperate fauna including wild cat and seventeen other mammal species had already established itself in Britain (King 1962, Yalden 1982). Excavations at the Mesolithic seasonal hunting camp at Star Carr in the Vale of Pickering, North Yorkshire produced evidence that most of the Thatcham fauna was already established in Yorkshire by 9,500 b.p. (Fraser & King 1954). Although no cat remains were identified, the faunal composition and habitat evidence at Starr Carr strongly suggested that wild cat could have been part of the local fauna. In undated post-glacial deposits in Teesdale cave on the Durham side of Teesdale (Backhouse 1881) the remains of wild cat were found in association with wolf, pine marten, polecat, otter, badger, red and roe deer, all of which, apart from polecat and otter, were present in the Thatcham and Star Carr faunas. So by implication, despite the absence of direct proof, the wild cat may well have been established in the Yorkshire area by the Mesolithic period. In Yorkshire at Pifflehead Wood, Newtondale (SE/8395), further undated circumstantial evidence was a shelter identified by Simms (1972) as ‘possibly being a wild cat den', containing amongst other prey remains the bones of a small beaver {Castor fiber). Beaver is known to have been present in Yorkshire from 9,600 b.p. at Star Carr (Fraser & King 1954) and place-name evidence suggests its presence into historic times, possibly up to the 10th century, so the den could have been occupied at any time between those dates. The earliest positively dated wild cat evidence is a single bone from an early Iron Age village site dated 2,500-2,400 b.p. on Castle Hill, Scarborough (TA/0489) (Rowntree 1931, Rutter 1956). An atlas, a phalange and part of an immature lower jaw were excavated from the Iron Age village on the northern escarpment of the Wolds at Staple How (SE/8974) dated 2,560-2,450 b.p. (King 1963) and a femur and humerus were found at an Iron Age lakeside site on the northern shore of Lake Pickering at Thornton-le-Dale (SE/8382) a site thought possibly to have been under human occupation up to Roman times (Bate 1931). From the Roman occupation the identification of wild cat from skeletal evidence becomes increasingly difficult, due to the introduction into Britain during the 1st or 2nd century of the domesticated cat, a form considered to have been derived by selective breeding from the north African race of the wild cat Felis s. lybica (Kolb 1977). To the difficulties implicit in identifying incomplete skeletal remains from archaeological sites are added the potential effects of hybridisation and evolutionary trends towards a smaller size (Kolb 1977). Archaeologists have been understandably reluctant to differentiate cat material on the basis of structure and comparative measurements. Thus Bate (1931) judged Iron Age material from Thornton-le-Dale to be of wild cat solely on the grounds that the domestic form was not known in Britain at such an early date, and remains from Roman and later sites are generally recorded as domestic cat purely because of association with sites of human occupation. 43 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat Cats are recorded in relatively few archaeological excavation reports; this may reflect the primary interest of the archaeologist in mammals of economic importance, such as cattle, sheep etc., or may suggest infrequent occurrence. Indeed, where cat evidence is recorded, numbers are lower than species of economic value. Table 1 gives examples from the 2nd to the 17th century of cat skeletal remains in Yorkshire. As all the sites are of human occupation, these remains are likely by inference to be of Felis catus, although there is a possibility that hunted or trapped wild cats may be represented since remains of other wild species e.g. Roe Deer ( Capreolus capreolus ) have also been found in association with human occupation sites. Since Kitchener (1995) has identified reliable structural features in cat skull and jaws to enable differentiation between F. sylvestris and F. catus, it would be opportune to re-examine skeletal material from excavations in order to confirm late dates for sylvestris and early dates for catus. TABLE 1 Cat remains from Roman and later archaeological sites in Yorkshire Period Locality Cats (min. no. or status) Material Association speices Source 2nd Century 5th Century Doncaster (Roman town) Langton (well) 1 1 Tibia Jaw, scapula, limb bones 2 skulls (cranial fragments only) 1,5,6, 8, 11, 13 Doncaster Mus. colln. Dell (1932) 11th- 14th Century Petergate, York 2 2 tibiae 1,4, 6, 7, 8, 11, 12, 13 Ryder (1971) 1300-1450 Fangfoss 1 Mandible 1,2, 6, 8, 13 Youngson (1978) 1350-ca. 1400 Chapel Lane, Staith, Hull Scale Lane, Lowgate, Hull 1 Humerus, ulna 15 1,2, 8, 11, 13, 15,18 Phillips (1980) 1360-1380 (cesspit) 1 2 bones (unspecified) 21, 22 1,6, 8,11,13, 15, 17 Phillips (1980) Late 13th-early 14th Century Chapel Lane, Staith, Hull 1 1 bone (unspecified) 20,22 1, 2, 3, 4, 5, 7, 8, 9, 10 Phillips (1980) 13th-early 16th Century Warram Percy Scale Lane, Lowgate, Hull Common (unspecified bones) 11,13 1,8,13, 14, 15, 16, 17 Ryder (1974) 2nd quarter of 14th Century (garde-robe pit) Scale Lane, Lowgate, Hull 2 5 bones (unspecified) 19 Phillips (1980) 16th Century (well) 1 1 bone (unspecified) 1, 8, 13, 16, 17 Phillips (1980) Late 17 th Century Bedern, York ? ? ? Buckland (1974) Key to species: 1 Cow; 2 Dog; 3 Fallow deer; 4 Goat; 5 Hare; 7 Mouse; 8 Pig; 9 Rabbit; 10 Rat; 11 Red deer; 12 Roe deer; 13 Sheep; 14 Blackbird; 15 Chicken; 16 Duck; 17 Goose; 18 Grouse; 19 House sparrow; 20 Mallard 44 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat Place Name Evidence The term ‘cat’ often forms the first element in the names of topographical features and associated place names. Research into the origins of such names shows that some originate from the Old English ‘catt’ in the sense of ‘wild cat’ and therefore probably allude to the former association between the animal and the locality, while some may refer to a topographical structure which in some respect resembles a cat. Other etymological roots may bear a coincidental phonetic similarity to ‘cat’; examples include Catterick, known to the Romans as ‘Cataracta’, meaning waterfall, although the present form refers to the Welsh ‘cader’, meaning ‘hill fort’. The term ‘Caty’, as in Caty Well, is derived from an association with St. Catherine. Many place names however are derived from personal names such as the Old Danish ‘Kate’ or Old Norse ‘Cada’ and ‘Rati’ as in Cat Flatts and the Old English ‘Catta’ and ‘Cada’ as in Cat Foss. By examining the fifteen (7th series) 1 inch to 1 mile Ordnance Survey maps which cover the Yorkshire area and the ten volumes of the English Place Name Society dealing with the three Yorkshire Ridings (Smith 1937, 1961-1963, 1969) some 80 examples of place names containing the element ‘cat’ have been found and are presented in Appendix 1 . An analysis of these using Smith {loc. cit .) showed that 24 relate to personal names, five showed phonetic similarity to ‘cat’ but 29 are judged to allude to wild cat; the locations of all these are shown in Figure 1. Place names not located on Ordnance Survey maps but which appear in other documentary sources have been awarded map references centred on the main village or parish in which the place name is situated. Figure 1 shows that the names associated with wild cats are exclusively distributed along the Pennines or Pennine foothills, mainly at altitudes above the 400 ft contour. Tantalisingly, the distribution of 19 of the 22 sites for which the etymological derivation is uncertain also shows a close correlation with the upland wild cat sites. This may provide the possibility of further circumstantial evidence of early wild cat occurrence. Places associated with personal name roots are chiefly connected with phases of Scandinavian settlement and are thus distributed along the east coast and in the rich lowland agricultural areas of Holderness and the Vale of York. Although unconfirmed, Wiley Cat Wood in the Cleveland uplands of north-east Yorkshire could imply the former existence of wild cats in the area, a possibility enhanced by allegation that the last wild cat in Yorkshire was also a resident of these north-eastern uplands. The date at which a place or geographical feature is first named after the local occurrence of a wild cat, if indeed this was the reason for it being so named, potentially gives a clue as to when the species occurred in the district concerned. The earliest dates when these names appeared on manuscripts, maps etc. have been obtained where available from the relevant English Place Name Society volumes and are included in Appendix 1. Earliest documentary dates range from 1086 (Domesday Book) to the latest in 1866. That 63% of these fall in the mid- 19th century probably reflects the fact that detailed maps giving minor place names (tithe, estate and Ordnance Survey maps) became more readily available during this period. Licenses to Hunt Wild Cat from the 12th- 17th Centuries The granting of Royal licenses to hunt certain beasts of the chase, or to exterminate certain species regarded as vermin, frequently included the wild cat as potential quarry, thus providing indirect evidence of its past occurrence in certain counties, forests and manors. Miller (1804), Hunter (1828), Harting (1890) and Large (1954) collectively quote eleven examples of such documents dating from the reign of King John (1199-1216) up to 1630. Despite inferences to the contrary none of these actually concern Yorkshire, identifiable localities referring to Buckinghamshire, Essex, Huntingdonshire, Northamptonshire, Oxfordshire, Rutland, Sutherland and Perthshire. Evidence of wild cats being hunted in Yorkshire does however exist. On the return of Edward 1st to England in 1274 from crusade in the east, it was decided, in consultation with his council, to investigate improper alienations or encroachments on royal lands and Red in Tooth and Claw: 1 Studies on the History of the Wild Cat 45 • = Place name element attributed to Wild Cat. O = Place name element of uncertain origin. P & + = Place name attributed to personal name. A = Place name element with phonetic resemblance only. ^ = Parish where bounties were paid for Wild Cats. FIGURE 1 Possible former distribution of Wild Cat in Yorkshire based on place name evidence and bounty payment records in Churchwardens’ accounts. privileges which had accrued during the previous reign. From 1274 to 1275, from 1279 to 1281 and again from 1293 to 1294 appointed Commissioners held ‘sessions’ in each administrative division (‘hundred’) (or ‘wapentake’ in Yorkshire) and jurors were called to give written verdicts on evidence of rights and privileges claimed by local landowners and clerics. Examples of claimed rights to hunt various specifically identified quarry animals provide circumstantial evidence of these species occurring in delineated parts of Yorkshire during the 13th century (English 1996). The Abbot of St. Mary’s, York claimed to have the chase through the whole of the forest of the king [in the region of] York for fox, cat, hare and badger (English 1996). In 1252-53 Roger Hardy was allowed to hunt with hounds for the wild cat, hare, fox and badger in the Forest of Pickering (Rimington 1956), and in 1338 Henry Percy claimed the right to hunt the wild cat, fox, badger and roe deer in the manor of Seamer within the Forest of Galtres (Gill 1852, Anon. 1906). 46 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat Fur Trade Miller (1804), followed by Hunter (1828) and Gill (1852), showed that in addition to being hunted for sport and as vermin, cats were killed for their fur. The pelts of cats, along with those of rabbit, fox and lamb, were all locally obtainable but held a low prestige value, as evidenced in both ecclesiastical and civil laws of the medieval period. Archbishop William Corboyle’s Canon of 1127 ordained that no abbess or nun could use more costly apparel than was made of lamb or cat skin. The ‘Sumptuary Laws’ of 1363 and 1532 stipulate that unlike the pelts of imported fur-bearing species such as sable ( Mustela zibellina), mink (. Mustela lutreola), lynx ( Felis linx) and genet ( Genetta genetta), the use of which was restricted to the nobility and richer merchant classes, only cat, fox, rabbit and lamb could be worn by the lower orders of society, e.g. trades people owning more than 40 shillings worth of goods (Veals 1966). In the absence of statistical or anecdotal evidence, this low prestige value may be indicative of the abundance and accessibility of cats in the wild during this period. Locally acquired wild cat skins could possibly have been used by members of the Skinners Guilds which existed in medieval York and Beverley, pelts being collected from trappers and villagers by travelling peddlers, as was customary at that time. Direct reference to cats being used for fur in Yorkshire comes from the Ordinances of the Skinners of the City of York of 1500, which stipulated a price for the preparation of ‘catte skins’ (Raine 1943). It is not possible to determine whether these were domestic or wild animals or both, and as beaver (Castor fiber) and mink were also mentioned in the Ordinances, neither is it possible to establish firmly whether they were collected locally, traded from other parts of Britain or indeed imported from elsewhere in Europe. Bounty Payments for Wild Cats in Churchwardens’ Accounts of the 16th- 18th Centuries In 1566 an Act for the ‘Preservation of Grayne’ came into force, its aim being to encourage the destruction of a wide range of what were regarded as ‘ravening byrdes and vermyn’. Although forming a highly productive source of biological records from the late 16th to the early 19th centuries, the practice of paying head money declined rapidly from the first quarter of the 19th century and the Act was finally repealed in 1863. With at least eleven literature references to wild cats in Yorkshire parish records (viz. Eastwood 1851 and 1862, Hatton & Fox 1880, Clarke & Roebuck 1881, Roberts 1882, Harting 1890, Wroot 1895, Grabham 1907, Denny 1910, Taylor 1956, Howes 1973), evidence would seem to abound. In fact these oft repeated references relate to only two parishes, Ecclesfield, where bounties were paid for wild cats in 1589 and 1626 and Shipley, where payments were made for two cats in 1676, one in 1678 and two in 1679. For this study the churchwardens’ accounts from some eighty-eight Yorkshire parishes have been examined and in seven of these records bounty payments for a total of seventy wild cat heads have been located. The 64 records additional to those already published relate to Masham in North Yorkshire (27 from 1653 to 1776) and four parishes in South Yorkshire, Thorp Salvin (34 cats from 1699 to 1769), Whiston (1 in 1710), Harthill (1 in 1773) and Wadworth, where payment was made for one as recently as 1788 (see Appendix 2). Figure 1, which also shows the distribution of the parishes where bounty payments were made for wild cats, serves to give a more realistic impression of wild cat distribution from the 16th to 18th century. In counties to the south of Yorkshire records are scarce and relatively early, possibly signifying an early date of decline: for instance, in the parish of Worksop on the Nottinghamshire border with South Yorkshire, payments were made for two wild cats in 1585 and four in 1586 (White 1904). To the north of Yorkshire in Cumberland, MacPherson (1892) mentions the presence of wild cat in the ecclesiastical records from the parish of Martindale in 1739, and Watson (1886) notes bounty payments for twelve wild cats during the 1760s in the parish of Crossthwaite. Perhaps the most frequent records in parish accounts from the north of England, possibly reflecting relative abundance, were 47 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat from the district of Corbridge near the Durham/Northumberland border, where bounty payments for at least 414 wild cats were recorded in the incomplete documents surviving for the period 1677-1724 (Nelson 1881). With regard to abundance relative to other targeted carnivores, bounty payments for wild cat have only been located in the accounts of seven (8%) out of 88 parishes examined. This indicates their scarcity relative to fox, present in 34 (38%) parishes; otter, present in 13 (15%) parishes and badger, present in 12 (13%) parishes. Only Pine Marten was scarcer, being represented in only four parishes. To achieve a more refined comparison an analysis of all the 10,787 carnivore (and hedgehog) bounty payments from Yorkshire parish records showed that from 1550-1599 wild cats constituted 3% of bounties. This level fell to 0.3% of bounties from 1600-1649, accounted for 1% in 1650-1699 and 1% in 1700-1749, falling to 0.1% in 1750-1799 and was unrepresented during the 19th century (see Figure 2). I l I FIGURE 2 Numbers of Wild Cat bounties in 88 sets of churchwardens’ accounts from Yorkshire parishes compared with cats as % of total carnivore and hedgehog bounty payments. Although usually referred to as wild cats, there is no entirely certain way of distinguishing between wild and domestic animals from this archival source, though presumably churchwardens only knowingly paid for what they considered to be the former. Matheson (1940) sounds a note of caution by quoting a vestry minute from a Montgomeryshire parish, which showed that churchwardens had been deceived into paying for domestic cats. With significant financial inducement to defraud the system by substituting domestic cats, it could be said that the authenticity of the Yorkshire examples may rest on their early dates and infrequent occurrence. Whereas the larger carnivores like badger ( Meles meles), fox (Vulpes vulpes) and otter ( Lutra lutra ) generally commanded a bounty of 1 shilling, particularly through the 18th and early 19th centuries, this was only occasionally the case with wild cats during the 18th century (see Appendix 2 for examples from the parishes of Whiston in 1710, Harthill in 1773 and Wadworth in 1788). The smaller bounties (2d, 3d, 4d and 6d) often paid for cats 48 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat during the 16th and 17th centuries are probably a reflection of contemporary monetary values and also perhaps the smaller size of cats and kittens. Since these sums were typically paid for the smaller mustelids, particularly in the 18th and 19th centuries, this raises the posibility that the term ‘wild cat’ as used in the accounts was merely an alternative vernacular name for weasel, stoat or polecat. However, when wild cat bounties appear, they are generally higher than for the smaller mustelids. The relative scales of the bounty tariff therefore gives useful indication that the ‘wild cat’ is genuine. Gamekeepers’ Records From the 18th century, and probably up to the Second World War, cats which strayed into game preserves were regarded by game-keeping staff as vermin and killed, often in large numbers. These were probably all domestic and feral animals, though occasionally particularly large specimens were claimed to be wild cats, e.g. one at Northowram in 1830 (Johnson 1965). Some estate archives contain reports of vermin killed on specific game preserves, though information on species and numbers is more frequently obtained by examining gamekeepers’ gibbets or lines. Examples dating from 1866 to 1930 of vermin records which include cats have been located for six areas, details of which are given in Table 2. Although a small and variable sample, cats, as a percentage of the total vermin, appear to increase during this period. The only clear finding is that around the turn of the 19th century cats were relatively abundant on gamekeepers’ gibbets, while today they are generally absent. TABLE 2 Records of cats killed by gamekeepers 1866-1930 Locality Grid ref. Date No. of Cats Total head of ‘vermin’ Cats as %of ‘vermin’ Source Hooton Pagnell SE/40 1866-68 31 627 4.9 Ruston & Witney (1934) Irton TA/08 1885 Present ? ? Slater (1885) Irton TA/08 1890 100 755 13.2 Gyngell (1905) Suffield SE/99 1904 1 53 1.9 Sheppard (1904) Irton TA/08 4.3 1905 31 240 12.9 Gyngell (1905) Osmotherley SE/49 1908 28 100 28 Booth & Fortune (1908) Over Silton SE/49 1908 3 4 75 Booth & Fortune (1908) Market Weighton SE/84 5.7.1930 A few ? ? Booth (1930) Allusions in 19th Century Topographical Accounts During the 19th century there were several published allusions, both historical and contemporary, to wild cats occurring in Yorkshire. Some were merely general allusions used by authors attempting to illustrate the untamed nature of certain remote parts of the Yorkshire landscape in times past. However, it has been possible through this study to identify the basis of most of these statements. Gill (1852), referring to medieval times, describes the Forest of Galtres as ‘abounding with wild cats’, a claim based on the licence to hunt wild cats given to Henry Percy in 1338. Barker (1854) noted that after the Norman period, Wensleydale, to the west of Bainbridge and Askrigg, was a ‘wild forest inhabited by the . . . wild cat’. Graves (1808) included the wild cat in his catalogue of Cleveland animals, thus supporting the likelihood of Wiley Cat Wood (NZ/6414) near Guisborough 49 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat referring to this species. In Twist Wood, Northowram (SE/1126) a cat, allegedly a wild cat, was killed in late May 1830 ( Halifax Chronicle 1830, Johnson 1965). The 23 lb animal, crushed by a ‘Samson stone’ set up by the local gamekeeper, was alleged to have caused great destruction amongst rabbits, poultry etc. In support of the Barnburgh ‘Cat and Man Legend’ which sparked off this entire investigation. Hunter (1828) commented that wild cats still haunted the woods of South Yorkshire and were spoken of as objects of terror. James (1865) in his ‘ Early History of Sheffield’ used the tenor of Hunter’s 1828 account in describing the ‘thick hercynion forests which covered the slopes and valleys of Hallamshire’, which he says were roamed by the wild cat, ‘then a formidable beast’. Clarke and Roebuck (1881), later quoted by Denny (1910) and Fortune (1916), give what is regarded as the last record of wild cat in Yorkshire, one trapped by Mr John Harrison on his farm at Murton (SE/5388) near Hawnby, one winter around 1840. In addition, they claim that other testimony (not stated) confirms the opinion that the Hambledon Hills were the wild cat’s ‘latest’ haunt. It is strange that no contemporary report of such a noteworthy record was published, even in the Yorkshire literature and despite Clarke and Roebuck both serving as editors of the Naturalist and being prolific contributors of notes and papers, the record was never published by them in its pages. Although Barker (1854) alluded to wild cats as being ‘still occasionally found in the woods of Wensleydale’, thereby post-dating the 1840 Murton record, Clarke and Roebuck (1881), though unable to verify Baker’s allusion, conceded that in all probability wild cats once existed in the fells of the north west. Distribution, status and decline The preferred habitat of upland woodland, the borders of forest and open hilly ground where they hold territories of 60-70 ha (Kolb 1977) would, even by medieval times, have been restricted and fragmentary in much of Yorkshire, due to agricultural developments. This suggests that conditions may never have been suitable for them in the Vales and lowland of central and eastern Yorkshire, despite the various permissions of the 13th and 14th centuries to hunt wild cats in these areas. The Pennine distribution pattern which persuasively emerges from place name studies (see Figure 1) seems not to have survived into the 16th century according to negative evidence from ecclesiastical records, though allusions of former occurrence are still weakly echoed in 19th century topographical sources. However churchwardens accounts of the 16th to the 18th centuries provide robust evidence of an undocumented distribution along the wooded parishes on or adjacent to the Permian ridge from Masham in the north to Thorpe Salvin in the south. On the information available, it is difficult to plot precisely the wild cat’s decline in Yorkshire. Figure 2 shows the period during which bounty payments were made, with numbers peaking in the 1660s and 1670s and again in the 1720s and 1730s. Even during these periods, cats only represented 1% of total carnivore bounties, tailing off to 0.1% by the last half of the 18th century and vanishing altogether by the 19th century, a demise somewhat earlier than the claimed date of 1840 for the death of the ‘last’ wild cat in Yorkshire. Were even a residual population to have persisted after this time, wild cats would surely have featured, as did polecat, pine marten and otter, in the north regional natural history journals which proliferated during the mid- to late 19th century. Although plausibly argued by Langley and Yalden (1977) that the demise of the wild cat and the other rare carnivores in England came about by intensification of persecution through the 19th century boom in gamekeeping, it would seem likely from evidence presented here that in Yorkshire at least, its decline and ultimately the eradication of viable populations had realistically taken place a century earlier. The wild cat’s fate in Yorkshire probably followed the fate of its preferred habitat, though its final extermination could possibly have been brought about by persecution, isolation of populations and hybridisation with domestic (house) cats. That cats were killed by gamekeepers during the 19th and early 20th centuries is not in dispute, though with the relative frequency of culled cats increasing through this period (see Table 2) it is likely that 50 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat keepers were trapping wandering farm and feral domestic cats rather than wild cats. In Britain today, wild cats are confined to remote areas of Scotland north of a line between Edinburgh and Glasgow (Easterbee et al., 1991), normally occurring at relatively low altitudes. The estimated total pre-breeding population was calculated in 1995 to be about 3,500, with population density gradients evidently increasing from west to east and from north to south (Harris et al., 1995). 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Appendix 1: ‘Cat’ as an element in Yorkshire place names Key to etymological roots: C = cat, P = personal name, PH = phonetic resemblance, ? = uncertain origin English Place Name Society volume reference: W = West Riding, E = East Riding, N = North Riding Place name Grid ref. English Place Name Etymological Soc. vol. & page root Earliest date Cat Babbleton SE/6228 W4 7 9 Cat Babbleton TA/0074 ? Cat Beeston SE/2830 W3 217 P Catcliffe SK/4288 W1 179 c 1086 Cat Clough SE/1101 W1 232 c 1841 Cat Clough (Thurlstone) SE/2303 W1 341 c 1843 Cat Crag (Aldfield) S E/2669 W5 194 c 1841 Cat Crag (Dacre) SE/1960 W5 140 I Cat Craggs (Blubberhouses) SE/1650 W5 121 c Cat Crags (Hazelwood) SE/0958 W5 74 9 53 Red in Tooth and Claw : 1 Studies on the History of the Wild Cat Place name Grid ref. English Place Name Etymological Soc. vol. & page root Earliest date Cat Moss (Stutton) SE/4711 W6 26 ? Cat Flats (Marske) NZ/6322 N 154 P Catfoss Grange TA/1448 E 67 P Catfoss Hall Farm TA/1446 E 67 P Catfoss Manor TA/1447 E 67 P Catgate (Thornton-in-Craven) SD/9048 W6 33 ? Catgill SE/0654 W6 63 C 1539 Catgill (Carleton) SD/9749 W6 30 C 1843 Cat Hill SE/4203 W1 96 c 1841 Cat Hill SE/2405 W1 308 c 1605 Cathill Plantation (Barwick) SE/4037 W4 110 7 Cat Hole SE/0937 7 Cat Hole (Rishworth) SE/0318 W3 73 c Cat Hole (Todmorden) SD/9324 W3 180 ? Cat Holes (Bowland) SD/65 W6 213 ? Cat Holes (Bradfield) SK/2692 W1 232 ? Cat Holes (Marsden) SE/0411 W2 278 C 1850 Cat Holes SD/6590 W6 266 c 1603 Cat Holes Ings (Marsden) SE/0411 W2 278 c 1850 Cat Holes (Nesfield) S E/0949 W5 69 c 1847 Cat Not (Bowland Forest) SD/65 W6 210 7 Cat Knot SD/7159 W6 210 7 Cat Lane (Balne) SE/5819 W2 15 ? Cat Loup Dub (Dacre) SE/16 W5 143 C Catlow Fell S D/7 060 W6 210 C 1665 Catlow Low Laithe SD/7158 W6 210 c Catlow Gill (Carleton) SD/9749 W6 30 c Cat Moss (Rishworth) SE/0318 W3 73 c 1775 Cat Nab TA/2173 p Catrigg SD/8367 W6 155 c Catrigg Barn SD/8368 W6 155 c Catrigg Beck SD/8367 W6 155 c 1858 Catrigg Force SD/8367 W6 155 c Catriggs SD/9089 ? Catshaw SE/2003 W1 341 c 1647 Catshaw Cross (Thurlstone) SE/2003 W1 341 c 1843 Catstones (Baildon) SE/1539 W4 159 c Catstones SE/0638 W4 165 c 1849 Catstones SD/9640 W6 13 ? Catstones (Mytholmroyd) SE/0126 W3 164 C 1866 Catstones (Rishworth) SE/0318 W3 75 c Catstones (Shipley) SE/1437 W3 267 7 Catstones (Todmorden) SD/9424 W3 180 7 Catstone Wood (Hartwith) SE/2165 W5 146 7 Cattal SE/4454 W5 17 P Cattal Grange SE/4355 P Catall Lodge SE/4553 W5 17 P Cater Croft (Almondbury) SE/1515 W2 259 Ph Catterick SE/2299 N 242 Ph Catterick Bridge SE/2397 N 242 Ph 54 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat Place name Grid ref. English Place Name Etymological Soc. vol. & page root Earliest date Catterson (Almondbury) SE/1515 W2 259 ? Catterton S E/5 145 W4 236 Ph Catterton Beck SE/5046 W4 236 Ph Catton SE/3778 N 183 P Catton Moor SE/3878 P Catton Park SE/7352 P Catton Wood SE/1545 W4 197 P Catwick NZ/9005 N 118 P Catwick TA/1245 E 73 P Catwick Grange TA/1246 P Catwick House TA/1245 P Caty Well (Warley) S E/05 25 W3 126 P Caty Well Wood S E/05 25 W3 126 P High Catton SE/7153 E 186 P High Catton Grange SE/7352 E 272 P Little Catwick TA/1244 P Little Cattail (Goldsborough) SE/4454 W5 17 P Low Catton S E/7 05 3 E 186 P Low Catton Grange SE/7150 P Near Cat Clough SK/1799 ? Wiley Cat Wood NZ/6414 ? Appendix 2: Bounty payments for ‘Wild Cats’ in Churchwards’ Accounts from Yorkshire Parishes Bradford (Hatton & Fox 1880, Roberts 1882, Wroot 1895, Taylor 1956) Date Record 1676/77 Allowed to the churchwardens of Shipley for 6 urchanes [hedgehogs] a grey [Badger] and 2 Wild Cats. 1678/79 Allowed to the churchwardens of Shipley for 6 urchanes hedgehogs] & for a Fox head & for a Wild Cat. 1679 To the churchwardens of Shipley for 2 hedgehogs & a Wild Catt. £. s. d. -. 2. 4. -. 2. 2. -. -. 6. Ecclesfield (Eastwood 1851, 1862, Clarke & Roebuck 1881, Harting 1890, Grabham 1907 Denny 1910). Date Record £. s. d 1589 Item for Catte head. -. -. 4 1626 A Wylde Catte head. -. -. 2 Harthill (Sheffield City Archives PR 47/61) Date Record £. s. d 4.6.1773 To 1 Wild Cat to Thos. Pitchfork. -. 1. - Masham (NTCRO PR//MAS 3/1/1 Microfilm no. 995) Date Record £. s. d 1652 To John Thwaite for 7 Fox heads & 1 Wild Cat head. -. 8. - 1656 To Richd. ... for 1 Fox head, 2 badger heads & 2 Wild Cat heads. -. 4. - 1659 [Account book too tightly bound for microfilm to show amount paid]. 1659 1662 1663 1664 1665 1667 1671 1673 1776 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat 55 Marmaduke ... for 1 Wild Cat head To Robt. Pickersgill for 1 Fox head, 1 Fowmard [pole cat] head & 1 Wild Cat head -. 1. 6. To Sami. Beckwith for 1 Wild Cat head. -. -. 3. To Geo. Beckwith for 1 Wild Cat head. -. -.3. To Robt. Pickersgill man for 1 Wild Cat head. -. -. 4. To Sami. Beckwith for 1 Badger head & 2 Wild Cat heads. -. 2. -. To Robt. Pickersgill man for 1 Wild Cat head. -. -. 6. For 1 Wild Cat head to . . . Beckwith. -. -. 6. To . . . for 2 Badger heads & 1 Wild Cat head. -. 2. 3. To ... Beckwith man for 1 Wild Cat head. -. -. 6. To John Parker for 1 Wild Cat head. -. -. 4. Item Robt. Johnson for 1 Wild Cat head. -. -. 6. Item to Marmaduke Cross’s man for 2 Fowmard heads & 1 Wild Cat head. -. 1. -. Item to Thomas Jackson for 2 Fowmard heads & 1 Wild Cathead. -. 1. -. Item to Squire Beckwith man for 14 Fox heads, 1 Wild Cat head & 1 Fowmard head. -.14. 9. Item to Thos. ... for 3 Wild Cats heads & 1 Badger head. -. 2. 6. Item to Robt. Thompson of Swinton for 2 Fowmard heads & 1 Wild Cat head. -. 1. -. Item to Wm. ... for 6 Fox heads & 1 Wild Cat head. -. 6. 6. Item to Robt. Aiscough for 1 Wild Cat head. -. -. 6. Item to ... for 1 Wild Cat head. -. -. 6. Thorp Salvin Date 1699 4.7.1700 20.5.1707 9.4.1711 5.11.1711 12.12.1713 21.3.1718 26.10.1718 12.1720 10.9.1722 2.11.1724 13.1.1725 23.1.1725 3.3.1725 6.8.1725 3.3.1727 14.1.1729 29.6.1727 1732 1733 1737 (Sheffield City Archives MD/1236/1-64, 65-107; 1238/24, 30) Record Pd. to Henry Foster for 1 Wild Cat head. Pd. to Jo. Turner for 1 Wild Cat head. Pd. to Jo. Turner for 1 Wild Cat head. Pd. to Robt. Barker for 1 Wild Cat head. Pd. Jonathon Horwoods for 5 Badger heads & 1 Wild Cat head. Myself for 1 Wild cat head. Jonathon Horwoods for 1 Fox heads & 1 Wild Cat head. Pd. John Horke for 1 Wild Cat head. Pd. to Robt. Dunston for 1 Wild Cat head. Pd. to Ed. Alin for 1 Wild Cat head. Pd. to Arthur Barlow 1 Wild Cat head. Pd. Harry Foster for 1 Wild Cat. Pd. to John Alin for 1 Wild Cat. Pd. to Thos. Wildsmith for 1 Wild Cat head. Pd. to Ed. Alin for 1 Wild Cat head. Pd. to Jas. Crawshaw for 1 Wild Cat head. Pd. to Jas. Crawshaw for 1 Wild Cat head. Pd. to Jas. Crawshaw for 1 Wild Cat head. Pd. to Jas. Crawshaw for 2 Badger heads, 1 Fox head & 2 Wild Cat heads. Pd. to Sami. Booth for 1 Wild Cat head. Pd. for 2 Wild Cat heads & 1 Pole Catt head. Pd. to Jas. Ryall for 1 Fox head & 1 Cat head Pd. to Anthony Barlow for 6 Wild Catts. Pd. to Anthony Barlow for 1 Wild Cat head. Pd. to John Berry for 1 Wild Cat head. s. d. -. 4. -. 4. -. 4. -. 4. 5. 4. -. 4. 1. 4. -. 4. -. 4. -. 4. -. 4. -. 4. -. 4. -. 4. -. 4. -. 4. -. 4. -. 4. 3. 8. -. 4. 1. 6. 2. -. 2. -. -. 4. -. 4. 56 Red in Tooth and Claw: 1 Studies on the History of the Wild Cat Date Record £. s. d. 6.5.1769 Pd. for 1 Fox & 1 Wild Cat. -. 1. 4. Wadworth (Doncaster MBC Archives P21/3/B2-3) Date Record £. s. d. 23.5.1788 To a Wild Cat head. 1. Whiston (Sheffield City Archives PR 37/43) Date Record £. s. d. 1710-1711 Given to Robert Natoris man by ye Townes Consent for killing ye Wild Catt. 1 . BOOK REVIEW Guardians of the Salmon by Gordon H. Bielby. Pp. 144, with b/w historical photographs. Halsgrove, Tiverton. 2001. £16.99 hardback. The literature of angling is vast, but apart from one or two classics such as Walton’s Compleat Angler, few fishing books are appreciated by naturalists. This book is different for it is not a book about angling and is written by a non-angler. As th author says “It contains nothing about how, when and where to go angling for salmon” (or indeed any other species). Gordon Bielby is a Yorkshireman born in Scarborough; except for a short spell in Canada with Alberta Fish and Wildlife, he has spent his whole career on the environmental side of river management in the southwest of England. For all who are seriously interested in river conservation this book is a must. The research has been painstaking and extensive, complemented by annual reports, newspaper archives and museum records. There are hundreds of personal reminiscences and anecdotes from long departed netsmen, water baliffs and poachers. The anecdotes are presented in the form that make reference easy, with a dry wit which seems to indicate the author’s disapproval of some of the methods of ‘conservation’. The slaughter of ‘predators’ on a vast scale, including cormorants, shags, herons, seals, otters, even large brown trout and eels, is recorded in a chapter entitled “Predators must Die”. It was a foolish bird, seal or porpoise which strayed into the southwest river catchments. Three shillings (15p) was the reward for the head and stomach of a seal! All these efforts were in the interests of conserving the Atlantic Salmon. The illustrations show the scale of slaughter of the salmon by the netsmen. Of course this was a period in which the mystery of the salmon’s journeys in the seas had not been solved. Even today pressure is being put on the EC to agree to the culling of cormorants which frequent gravel pits and open water in Britain. Human predators were severely dealt with; even transportation was proposed. The many stories of encounters with the poachers and their nefarious methods reflect the hard times in which the rural population lived. Our new and thriving biodiversity industry has suddenly become concerned with preserving the genetic strains of wild salmonids. It will be hard to ignore the fact that millions of salmon ova of Scottish origin were planted in southwest rivers between 1864 and 1950. This is a book that should give food for thought for the modern day conservationists and their misguided allies who are recommending similar methods of eradication in the interests of individual species. LM AQUATIC PLANTS AT NORTH CAVE WETLANDS, A NEW YORKSHIRE WILDLIFE TRUST RESERVE COMPRISING FORMER GRAVEL WORKINGS 57 R. GOULDER Department of Biological Sciences, University of Hull, Hull HU 6 7RX Email: rgoulder@hull.ac.uk Introduction North Cave Wetlands, also known as the John Dealtry Reserve, comprise approximately 40 ha of former sand and gravel workings (GR: SE 883 331) situated about 1 km north-west of the village of North Cave, East Yorkshire. The site, comprising six large pits (Fig. 1) which were excavated in the 1980s and 1990s (Table 1), was acquired by the Yorkshire Wildlife Trust in late 2000 (Anon., 2001). The following account is based on visits to the reserve made during June and July 2001. Pits 1, 2, 3, 5 and 6 have become flooded, forming lakes that are, in places, many metres deep. The lake margins are generally sand or gravel. There are trees and shrubs along the western sides of Lake 1 , parts of the eastern and western sides of Lake 2, and the southern side of Lake 3; otherwise there is little shading. There are some wet, flushed, areas of shore where there is up- welling of ground water, most notably at the south west corner of Lake 5 and at the north east corner of the irregularly-shaped eastern section of Lake 5. Lake 5 is linked to its eastern section by a water-filled channel, 3-5 m wide and at least 2 m deep (Fig. 1). This eastern section of Lake 5 is joined to Lake 6 by a shallow neck of water. There is a pipe through which water flows from Lake 2 into Lake 1 . Lake 1 has an outflow pipe on its north side that discharges into the deeply-excavated drainage channel, Black Dyke, that runs along the northern boundary of the site. The water was very transparent in all five lakes and all had similar pH and conductivity; on 6 July, measurements made with appropriate meters showed a pH range of 6.6 to 6.8 while that for conductivity was 929 to 1002 pS cm1. Pit 4 was not water-filled. It was a large excavation the bottom of which was several metres below the water level in the adjacent Lake 1. On the bottom of Pit 4, in its north- west corner, there was a small (longest side about 30 m), shallow (depth <1 m), triangular pond. There were also muddy areas, other shallow pools and flooded scrapes, and ditches that were filled by seepage water from Lake 1, from which water was pumped back into Lake 1. During summer 2001 this pit was being back-filled with subsoil and rubble. The reserve is an important site for waterfowl. Ducks and geese were abundant on the lakes and had heavily grazed some of the shore vegetation. Between the lakes, the site is mostly open grassland, with areas of bare, recently-worked ground; there are few trees. In this study aquatic macrophytes at North Cave Wetlands were recorded to provide a base line for reference as the site matures. Aquatic plants within 1 km beyond the site boundary were also recorded as these neighbouring plants are potentially responsible for past and and future colonisation. Aquatic Plants at North Cave Wetlands Aquatic macrophytes were recorded during June-July 2001 in and around the margins of the five lakes. The eastern section of Lake 5 (henceforth referred to as Lake 5e) was treated as being part of Lake 6, with which it is continuous. Recording was done by walking around the circumference of the lakes and by grapnel hauls. Aquatic macrophytes in the ponds and wet places at the bottom of Pit 4 were also recorded. The checklist used was that of aquatic plants that occur in England and Wales (Palmer & Newbold, 1983) except that all Juncus species and charophytes were included. Nomenclature follows Stace (1997) for vascular plants and Moore (1986) for charophytes. Twenty-six species of aquatic macrophyte were recorded. Their distribution is shown in Naturalist 127 (2002) 58 Aquatic Plants at North Cave Wetlands, a New Yorkshire Wildlife Trust Reserve metres FIGURE 1 Map of North Cave Wetlands. Stippling indicates the area of water in August 2000; the water levels are higher in winter. Table 1 with an indication of their abundance. The number of species per lake ranged from ten in Lake 2 to 17 in Lake 5e/6. The older lakes (1-3) supported no more species than the newer ones. The most frequently found submerged species was Potamogeton pusillus which was present in all the lakes and abundant in all except Lake 3. Zannichellia palustris was also found in all the lakes. Otherwise, the submerged species showed notable between-lake variation. Chara vulgaris was abundant in Lakes 1, 3, 5 and 5e/6 but was not recorded in Lake 2. Furthermore, there may be varietal differences between the lakes: Chara vugaris var. vulgaris and var. longibracteata were found in Lakes 1 and 5e/6 while var. papillata was found in Lakes 3 and 5. Myriophyllum spicatum was abundant in Lakes 1-3 but was unimportant in Lake 5e/6 and was not recorded in Lake 5. Potamogeton crispus was abundant in Lake 3 but was not found in the other lakes, while Potamogeton pectinatus was recorded only in Lakes 2, 3 and 5e/6. The aquatic plants at the lake margins also showed between-lake variation. Only Agrostis stolonifera and Ranunculus sceleratus were found around all five lakes. Apium nodiflorum was found only by Lake 5e, Phragmites australis only by Lakes 3 and 5e, Typha latifolia only by Lakes 2, 5 and 5e, Veronica anagallis-aquatica only by Lakes 5 and 5e/6, and Veronica beccabunga only by Lakes 1, 2, 5 and 5e/6. The importance to the marginal plants of the limited areas of wet, flushed, shore with upwelling groundwater was very apparent. The flushed area at the south-west corner of Lake 5 supported luxuriant 59 Aquatic Plants at North Cave Wetlands, a New Yorkshire Wildlife Trust Reserve TABLE 1 Aquatic plants at North Cave Wetlands; June-July 2001 Lake 1 1983 Lake 2 1987 Lake 3 1990 Lake 5 Lake 5e/6f 1994 1995/1996 Pit 4 1999 Submerged plants Chara vularis t +++ +++ +++ +++ Myriophyllum spicatum +++ +++ +++ - + - Potamogeton crispus* - - +++ - - - P. pectinatus* - ++ ++ - + - P. pusillus +++ +++ ++ +++ +++ +++ Zannichellia palustris + ++ + +++ +++ + Plants growing around margins and in muddy places Agrostis stolonifera ++ ++ + + + ++ Apium nodiflorum - - - - + - Callitriche sp. - - - - - + Eleocharis palustris - - + - - ++ Equisetum fluviatile* - - - - - ++ Juncus articulatus + - - ++ + +++ J. bufonius + + - ++ ++ +++ J. conglomeratus* + - + + - + J. effusus + - - + + + J. inflexus + - + ++ + + Mentha aquatica - - + - - - Phragmites australis - - + - ++ + Ranunculus aquatilis* - + + - - + R. peltatus* - - - - - + R. sceleratus ++ + + ++ ++ +++ Rorippa nasturtium-aquaticum - - - ++ + ++ Samolus valerandi* - - - - - + Typha latifolia - ++ - + ++ + Veronica anagallis-aquatica - - - ++ ++ ++ V. beccabunga ++ + - ++ + ++ The lakes are arranged in age order; approximate dates when excavation was completed are shown. ^Indicates species that were not also found within 1 km outside the Wetlands site boundary. fCharophytes were determined by P. J. Cook and included Chara vulgaris var. vulgaris (Lakes 1 and 5e/6), C. vulgaris var. papillata (Lakes 3 and 5) and C. vulgaris var. longibracteata (Lakes 1 and 5e/6). + = Plants sparsely or locally distributed in small patches; ++ = plants frequent around the margins, or forming local patches of a few square metres in area; +++ = plants abundant around margins or forming substantial submerged beds; - = not recorded. $The eastern section of Lake 5 is included with Lake 6. vegetation which included Juncus articulatus, J. bufonius, J. ejfusus, J. inflexus, Ranunculus sceleratus, Rorippa nasturtium-aquaticum, Typha latifolia and Veronica anagallis-aquatica. Similarly, the flushed areas around the north-east corner of Lake 5e supported Apium nodiflorum, Juncus articulatus, J. bufonius, J. inflexus, Phragmites australis, Ranunculus sceleratus, Rorippa nasturtium-aquaticum, Typha latifolia, Veronica anagallis-aquatica and V. beccabunga. More aquatic species (20) were recorded in the non-flooded Pit 4 than in any of the lakes. Species found only in Pit 4 were Callitriche sp. on mud, Equisetum fluviatile and 60 Aquatic Plants at North Cave Wetlands, a New Yorkshire Wildlife Trust Reserve Samolus valerandi (three plants only) in muddy areas with up-welling seepage water, and flowering stands of Ranunculus peltatus in the small triangular pond. Aquatic plants within one kilometre of North Cave Wetlands Aquatic macrophytes at sites outside the reserve, but within 1 km of its boundary, were recorded during July- August 2001. Most of the ponds and watercourses shown on the 1:25000 scale OS maps within 1 km of the site were visited. Standing-water sites were: (1) the ornamental fishing lake (SE 897 328), immediately north of the church and just within 1 km east of the Wetlands boundary, which was mentioned in 1769 and probably dates from the mid-18th century (Kent, 1979); (2) two ponds at SE 890 325, about 600 m south-east of the Wetlands boundary, which possibly originated as medieval fish ponds (May & May, 1999); (3) a gravel-pit pond at Crosslands (SE 879 322), 500 m south of the Wetlands site, excavated in the 1970s, (4) a gravel-pit pond in North Cave village (SE 888 321), 700 m south of the Wetlands boundary, excavated in the 1980s. Stream sites were along much of North Cave Beck from the head of the fishing lake downstream almost to the dismantled railway line (SE 889 320), about 1.75 km of watercourse. The beck is a clear-water, spring-fed, calcareous stream with silt and gravel bed; much of its course is artificially aligned and is tree-shaded. At its closest, the beck is within about 500 m of the Wetlands boundary. Dykes that drain the low land, altitude <10 m, to the west of North Cave include Black Dyke and Ings Drain. Black Dyke forms the northern boundary of the Wetlands site; aquatic plants in the dyke along this boundary were recorded in two 100 m lengths, one upstream and one downstream of the outflow from Lake 1. Two further 100 m lengths of Black Dyke were surveyed adjacent to the Dryham to Hotham Common farm road (SE 874 331) about 500 m west (downstream) of the Wetlands site. There was an appreciable flow along the dyke but most of this water originated as the discharge from Lake 1. Water plants in Ings Dyke were recorded along about 300 m of the watercourse, eastwards from Dryham (SE 876 326) to within about 125 m of the Wetlands boundary. There was little flow in this dyke. Aquatic plants in a wet horse pasture (SE 891 325), adjacent to North Cave Beck and about 600 m south east of the Wetlands site were also recorded. Thirty-four species of aquatic macrophyte were recorded within 1 km of the Wetlands boundary, i.e. plants on the Palmer and Newbold (1983) checklist plus all J uncus spp. and charophytes. Many of these neighbourhood plants, 15 species, were not found on the Wetlands site (Table 2). Also found at neighbourhood sites, but not on the checklist, were the aquatic moss, Fontinalis antipyretica, in North Cave Beck, and the Galingale, Cyperus longus, an obvious introduction in the Crosslands gravel pit. Some of the neighbourhood plants, shown in Table 2, are also likely to be recent introductions: e.g. Typha angustifolia and perhaps T. latifolia and Iris pseudacorus, again at the Crosslands gravel pit; I. pseudacorus and varigated Glyceria maxima at the gravel pit in North Cave; I. pseudacorus and T. latifolia in the beck where it flows at the bottom of gardens. Water lilies, still in pots, at the gravel pit in North Cave are not shown in Table 2. There were also seven species of aquatic macrophyte that were recorded at North Cave Wetlands but not at neighbourhood sites (Table 1). Discussion The aquatic-plant species richness in the gravel pits at North Cave Wetlands was around the middle of the range found for other ponds formed by mineral extraction in East Yorkshire. The mean number of taxa per pit, including the non-flooded Pit 4, equalled 14.5 (range 10-20). This may be compared with other East Yorkshire ponds: i.e. means of 13.8 (4-23) for 11 gravel-pits, 15.8 (8-26) for nine clay-pits, and 14.4 (2-25) for ten borrow-pits alongside railway lines (Linton & Goulder, 2000). The between-lake variation in the distribution of the submerged species (Table 1) may 61 Aquatic Plants at North Cave Wetlands, a New Yorkshire Wildlife Trust Reserve TABLE 2 Aquatic plants recorded within 1 km of North Cave Wetlands; July- August 2001 Ponds Streams Dykes Wet pasture Agrostis stolonifera + + + - Alisma plantago-aquatica* - + - - Apium nodiflorum - + + + Callitriche sp. + - - - Caltha palustris * - + - + Carex acutiformis* - - + - C. riparia* + + + - Chara vulgaris + - - - Eleocharis palustris + - - + Glyceria maxima* + + - + G. notata/fluitans* + - + + Iris pseudacorus* + + - - Juncus articulatus + + - + J. bufonius + - + + J. effusus + + — - J. inflexus + + + + Lemna minor* - - + - Mentha aquatica + + + - Myosotis scorpioides* + + - + Myriophyllum spicatum + - + - Phalaris arundinacea* + + - - Phragmites australis + - + - Potamogeton pusillus + - + - Ranunculus circinatus * + - - - - R. sceleratus + - + - Rorippa nasturtium-aquaticum + + + - Solanum dulcamara* + + - - Sparganium erectum* + + + - Typha angustifolia* + - - - T. latifolia + + - - Veronica anagallis-aquatica - - + - V. beccabunga + + + + V. catenata* + - - - Zannichellia palustris + - - - indicates species that were not also found on the North Cave Wetlands site. + = Present; - = not recorded. reflect the chance element in plant dispersal. The variation was also potentially related both to the age of the lakes and to competition between species. Thus Myriophyllum spicatum was abundant in the older Lakes 1, 2 and 3 but was relatively sparse in Lakes 5e/6 and was not recorded in Lake 5. Furthermore, the relative scarcity of Zannichellia palustris in Lakes 1, 2 and 3, which contrasted with its abundance in Lakes 5 and 5e/6, may be related to its being out-competed by M. spicatum. It will be interesting to see whether M. spicatum extends its range and replaces Z. palustris in Lakes 5 and 5e/6. The greater importance of marginal plants in Lakes 5 and 5e/6 (Table 1) was probably due to the presence of flushed areas at their margins. Also, extensive erosion faces around the margins of the older Lakes 1, 2 and 3 tended to make them less hospitable habitats. The presence of more aquatic plant species, largely plants of muddy places, in Pit 4 than in any of the lakes (Table 1) probably reflected the greater habitat diversity of Pit 4, 62 Aquatic Plants at North Cave Wetlands, a New Yorkshire Wildlife Trust Reserve although plant propagules may have arrived at Pit 4 along with back-fill material. None of the aquatic vascular plants found at North Cave Wetlands (Table 1) have outstanding conservation value. None are rare or threatened in Britain: i.e. known from 15 or fewer 10-km squares. Nor are any of them among the scarce species: i.e. known from 16-100 10-km squares (Preston & Croft, 1997). From a regional perspective. Palmer and Newbold (1983) reported them all as being frequent in the then Yorkshire Water Authority area. At the local level of East Yorkshire, however, some of the plants do have conservation interest. Crackles (1990) regarded Ranunculus aquatilis as uncommon (i.e. believed then to occur at between 12 and four localities) although probably under-recorded. She also regarded Potamogeton pusillus and Samolus valerandi as uncommon. These too, however, are probably under-recorded; both were found in several out of thirty-five 100 m lengths of dyke in the Hull Valley by Goulder (2000). Also, some of the other water plants in the North Cave pits were considered to be infrequent in East Yorkshire by Crackles (1990): i.e. Potamogeton pectinatus (in Lower Derwentland, which includes North Cave, although frequent in the Hull Valley) and the probably more scarce Ranunculus peltatus. There are also few records from East Yorkshire of the charophytes that were found at North Cave Wetlands. Moore (1986) has no post- 1960 records for Chara vulgaris var. vulgaris, and has only single post- 1960 records, in the extreme north east of the county, for C. vulgaris var. longibracteata and C. vulgaris var. papillata. Charophytes are, however, probably extensively under-recorded or in the case of C. vulgaris agg. not determined to varietal level; recent critical determinations have shown var. papillata and var. longibracteata to be the most frequently encountered taxa in south east Yorkshire (P. J. Cook, pers. comm.). The large size of the lakes at North Cave Wetlands, relative to the size of most standing waters in East Yorkshire, gives botanical conservation value to the reserve. There are extensive stands of submerged hydrophytes, especially of Potamogeton pusillus and Chara vulgaris, which are unusual in the county. Furthermore, there is the potential for further colonisation and the long-term development of a more diverse freshwater flora; larger ponds may develop greater species richness (Helliwell, 1983). Also, the water plants are probably an important food resource for the waterfowl that are the principal wildlife interest at the site. Species of Potamogeton, and Chara vulgaris can be important components of waterfowl diet (Preston, 1995; Knapton & Petrie, 1999). Such grazing may, to some extent, suppress vegetation but is sustainable in larger water bodies (Jupp & Spence, 1977; Mitchell & Wass, 1996; Strand & Weisner, 2001). Aquatic plants in general are surprisingly mobile and utilise diverse and effective dispersal strategies (Cook, 1987; Barrat-Segretain, 1996; Preston & Croft, 1998). Thus up to 26 taxa (Table 1) have managed to arrive at North Cave Wetlands over roughly twenty years since extraction of gravel began. Most of them were probably not originally present on the site as it was formerly arable land and because there are no water bodies shown on the pre-excavation 1:50000 scale OS map apart from the northern boundary ditch (Black Dyke) which is species poor: i.e. in July 2001 this ditch where it runs alongside the reserve contained only six aquatic species - Lemna minor, Myriophyllum spicatum, Potamogeton pusillus, Ranunculus sceleratus, Rorippa nasturtium- aquaticum and Veronica beccabunga. Of these, the plants of M. spicatum and P. pusillus were loose shoots that had clearly come through the discharge pipe from Lake 1, while L. minor was not found in the pits. Mechanisms used by water plants to reach the site probably included wind dispersal and carriage by waterfowl. Phragmites australis and Typha latifolia produce wind-dispersed seeds, and both occur within the neighbourhood (Table 2). Seeds and vegetative propagules of vascular plants are carried on feathers and feet or in the guts of waterfowl (Proctor, 1968; Vivian-Smith & Stiles, 1994). Preston (1995) notes that species of Potamogeton are rapid colonisers of new habitats such as gravel pits, and considers that the carriage of their seeds in the guts of birds is a significant dispersal mechanism. Similarly, charophytes, which are also often early colonisers of new waters, may be transported as oospores and vegetative fragments in the guts of waterfowl (Moore, 1986). Many of the water plants recorded at North Cave Wetlands probably originated from 63 Aquatic Plants at North Cave Wetlands, a New Yorkshire Wildlife Trust Reserve freshwater sites in the immediate neighbourhood. Nineteen of the taxa found on the site were also present within 1 km of the site boundary (Table 2). Some of these species were especially abundant at neighbourhood sites; e.g. Apium nodiflorum in Ings Drain and Myriophyllum spicatum in the fishing lake near to North Cave church. Thus they provided a rich source of propagules potentially available for colonisation of the newly-dug pits. Proof of place of origin is not available, although verification of the source of clonal species should be possible using molecular genetic fingerprinting techniques as used, for example, with the Flowering-rush Butomus umbellatus by Fernando and Cass (1996). Since there were also 15 species that were recorded in neighbourhood waters but not at North Cave Wetlands (Table 2), there remains scope for further colonisation from local sites. The seven species recorded at the Wetlands but not at neighbourhood sites (Table 1 ) may have come from further afield. Proctor (1968) showed that viable seeds are retained by ducks and geese for over 24 hours and so may be carried for long distances. It is also possible that some plants have arrived from further afield and, having colonised the Wetlands, have later spread to neighbouring waters. For example, Potamogeton pusillus was found rooted in Black Dyke about 500 m downstream of the site boundary. Thus, although P pusillus does occur in some East Yorkshire dykes (Goulder, 2000), its origin here may have been the loose shoots which reach the dyke in the discharge from Lake 1. Management of the Wetlands by the Yorkshire Wildlife Trust will influence the development of the aquatic plant flora. The Trust’s proposals are described in its planning application to the East Riding of Yorkshire County Council (01/04197) which was approved by the authority in November 2001. After back filling. Pit 4 will be flooded to create a reed- bed habitat with water depth of about 20-60 cm. By January 2002 the back filling had buried some aquatic plants which, in summer 2001, were found only in Pit 4, especially Equisetum fluvicttile, Ranunculus peltatus and Samolus valerandi. However, these species might reappear since the plans for the whole site include the creation of several small shallow ponds, and some reprofiling of Lakes 5 and 6 to form shallow foreshore and marshy areas. Ideally, the relatively floristically-rich flushed areas on the margins of Lake 5 and its eastern extension will be incorporated into the re-profiling scheme. The plants of Phragmites australis required to establish the reed bed in Pit 4 will best be grown from locally gathered seed. The final layer of fill into which the reeds are planted may need to be of soil. Thus new aquatic species might be introduced if they are part of the seed bank in this soil. Otherwise, the existing dispersal mechanisms, which have so far brought a diversity of plants to the site, can be expected to continue. Acknowledgements I am grateful to the Yorkshire Wildlife Trust for access to the site, to other land holders for access around North Cave, to Peter Edmonds and ARCO Ltd for the map of the site, to Mike Juggins for information on dates of excavation, to C. D. Preston for confirming Potamogeton pusillus, to Peter. Cook for identifying charophytes, and to Derrick Boatman for his ecological input and helpful criticism of the manuscript. References Anon. (2001) New reserve for Trust. Yorkshire Wildlife Winter/Spring: 5. Barrat-Segretain, M. H. (1996) Strategies of reproduction, dispersion, and competition in river plants: a review. Vegetatio 123: 13-37. Cook, C. D. K. (1987) Dispersion in aquatic and amphibious vascular plants. In: Plant Life in Aquatic and Amphibious Habitats (ed. Crawford, R. M. M.):~ 179-190, Blackwell Scientific Publications, Oxford. Crackles, F. E. (1990) Flora of the East Riding of Yorkshire. Hull University Press and Humberside County Council, Hull. Fernando, D. D. and Cass, D. D. (1996) Genotypic differentiation in Butomus umbellatus (Butomaceae) using isozymes and random amplified polymorphic DNAs. Canadian Journal of Botany 74: 647-652. 64 Aquatic Plants at North Cave Wetlands, a New Yorkshire Wildlife Trust Reserve Goulder, R. (2000) Aquatic macrophytes and conservation in the drains of the Hull Valley. Naturalist 125: 17-28. Helliwell, D. R. (1983) The conservation value of areas of different size: Worcestershire ponds. Journal of Environmental Management 17: 179-184. Jupp, B. P. and Spence, D. H. N. (1977) Limitations of macrophytes in a eutrophic lake, Loch Leven. II. Wave action, sediments and waterfowl grazing. Journal of Ecology 65: 431-446. Kent, G. H. R. (1979) North Cave. In: A History of the County of York: East Riding. Volume 4 (ed. Allison, K. J.): 19-37, Oxford University Press, Oxford. Knapton, R. W. and Petrie, S. A. (1999) Changes in distribution and abundance of submerged macrophytes in the Inner Bay at Long Point, Lake Erie: implications for foraging waterfowl. Journal of Great Lakes Research 25: 783-798. Linton, S. and Goulder, R. (2000) Botanical conservation value related to origin and management of ponds. Aquatic Conservation: Marine and Freshwater Ecosystems 10: 77-91. May, S. and May, G. (1999) Walking Round North Cave. Privately printed. Mitchell, S. F. and Wass, R. T. (1996) Grazing by black swans ( Cygnus atratus Latham), physical factors, and the growth and loss of aquatic vegetation in a shallow lake. Aquatic Botany 55: 205-215. Moore, J. A. (1986) Charophytes of Great Britain and Ireland. Botanical Society of the British Isles, London. Palmer, M. and Newbold, C. (1983) Focus on Nature Conservation No. 1, Wetland and Riparian Plants in Great Britain. Nature Conservancy Council, Shrewsbury. Preston, C. D. (1995) Pondweeds of Great Britain and Ireland. Botanical Society of the British Isles, London. Preston, C. D. and Croft, J. M. (1997) Aquatic Plants in Britain and Ireland. Harley Books, Colchester. Preston, C. D. and Croft, J. M. (1998) Britain’s changing aquatic flora. British Wildlife 10: 18-28. Proctor, V. W. (1968) Long-distance dispersal of seeds by retention in digestive tract of birds. Science 160: 321-322. Stace, C. (1997) New Flora of the British Isles. 2nd ed. Cambridge University Press, Cambridge. Strand, J. A. and Weisner, S. E. B. (2001) Dynamics of submerged macrophyte populations in response to biomanipulation. Freshwater Biology 46: 1397-1408. Vivian-Smith, G. and Stiles, E. W. (1994) Dispersal of salt marsh seeds on the feet and feathers of waterfowl. Wetlands 14: 316-319. BOOK REVIEW The Fever Trail. The Hunt for the Cure for Malaria by Mark Honigsbaum. Pp. xviii + 333 (inch 5 maps), plus 16pp. b/w & colour plates. Macmillan, London. 2001. £18.99 hardback. Until it became known in the mid- 17th century that the bark of the Cinchona tree could stave off fevers, no one was safe from the scourge of malaria. Two-thirds of this book documents the search for this miraculous bark and the quinine it contains; it honours the men who risked their lives for it, including Richard Spruce (1817-1893), the famous Yorkshire botanist and explorer who spent 15 years in the Amazon and Andes. Whilst in Ecuador, Spruce collected 100,000 seeds and 600 seedling plants of Cinchona which were successfully shipped to England; it was from these shipments that the Cinchona plantations in and industry of S.E. Asia were developed. Despite all the endeavours of such men, so vividly described by the author, malaria still kills up to three million people each year. Thoroughly recommended, not only to those concerned with ethnobotany and medical history, but also to those with wider natural history interests. MRDS 65 DISTRIBUTION OF THE BANDED DEMOISELLE CALOPTERYX SPLENDENS ALONG THE RIVER DERWENT, NORTH YORKSHIRE RICHARD SUNTER 3 Kingsley Garth, Outwood, Wakefield WF1 2LD The Banded Demoiselle Calopteryx splendens (Harris, 1782) is a lowland riparian species with a preference for relatively unpolluted, slow flowing rivers with silt beds. In Yorkshire the main population appears to occur along the River Derwent and other tributaries of the Ouse in the Vale of York, for example the River Wharfe. In west and south Yorkshire it is relatively scarce, probably as a result of pollution of the main rivers from the traditional heavy industries that dominated this part of the county. However, it has been recorded from the River Dearne in Barnsley, the Don at Doncaster (SE/5603) and the Torne catchment at Potteric Carr (SE/6000) and Torne Bridge (SE/6703 to 6903). This paper reports on the results of a survey undertaken in 1999 and 2000 to determine the distribution of the Banded Demoiselle along the Derwent. The Derwent rises on Fylingdales Moor from whence it passes 5 km to the west of Scarborough. At Mowthorpe Farm the Sea Cut has been constructed to channel excess water directly to the North Sea at Scalby Mills. Between its source on the North York Moors and East Ayton the river is swift flowing with a stony bottom and is therefore far from optimal for the Banded Demoiselle. At East Ayton the river enters the Vale of Pickering, a wide expanse of intensively farmed arable countryside, where along much of its course it has been extensively straightened and has the appearance of a canal, with opportunities for surveying restricted to a few road bridges. Approximately 5 km north of Malton the Derwent meets the River Rye, after which it flows between Malton and Norton, down towards Stamford Bridge and into the Vale of York. The Lower Derwent Valley has been designated as a SSSI and through much of this the river passes meadowland which extensively floods in the winter. Eventually at Barmby the Derwent meets the Ouse which is tidal at this point. However, the construction of the Barmby Barrage almost eliminates any effect of the tide in the lowest sections of the Derwent. Methodology The Derwent and a small part of the Rye at Ryton (SEM96754) were surveyed by walking along the river bank and counting the number of Banded Demoiselle seen. The areas around Elvington Bridge, Cottingwith and Wheldrake Ings were surveyed on 9 June 1999. The area around Stamford Bridge was surveyed on 10 June 2000, the section between Kirkham Priory and Howsham on the 11 June, around Malton, including the Rye at Ryton, on the 16 June and the section between Bubwith and the confluence with the Ouse on 27 June. Unfortunately, due to time constraints and lack of public access along parts of the riverbank, the whole course of the river was not surveyed. In addition, a small section of the Derwent at Brompton Ings (SE/954794) was surveyed from the bridge and the Sea Cut at Mowthorpe Farm, near Scarborough (SE/983884) was also surveyed. Results Calopteryx splendens was recorded from almost all 1 km squares surveyed between the confluence of the Derwent with the Ouse to just below that with the Rye above Malton. The few 1 km squares where the species was not recorded were generally those in which the river only passed through the square for a short distance and/or where the river was obscured by tree cover. It was not found above grid reference SE8072 or seen at the recording stations at Brompton Ings in the Vale of Pickering, the Rye at Ryton, at the Sea Cut at Mowthorpe Farm or in the Forge Valley. All records of the Banded Demoiselle are given in Appendix 1 below and the distribution of records is shown in Figure 1 . Naturalist 127 (2002) 66 Distribution of the Banded Demoiselle Calopteryx splendens along the River Derwent FIGURE 1 The Distribution of Records of the Banded Demoiselle Calopteryx splendens on the River Derwent, North Yorkshire 1999-2000. Hatched areas indicate 1 km squares through which the Derwent passes; areas shaded black indicate 1 km squares in which the Banded Demoiselle was recorded. Northings are the lines below the number in the extreme left-hand column. The first record of the Banded Demoiselle in the south-west corner is therefore SE/68-28. Distribution of the Banded Demoiselle Calopteryx splendens along the River Derwent 67 The Banded Demoiselle was found to be most abundant where the bank-side and adjacent land was dominated by rank grassland and/or scrub, and reasonably so in arable areas when it was seen settling in fields of wheat to the north of Stamford Bridge. The species appeared to be least abundant where the bankside vegetation had been grazed by cattle. Discussion The Banded Demoiselle appears to be common and widespread between the confluence of the Derwent with the Ouse and the Rye respectively. Given the range of situations and bank-side habitats in where it was found, including wooded banks, marsh/tall herb communities and arable fields, it is likely to be encountered in most of the 1 km squares of this stretch of the river. Above the confluence of the Derwent with the Rye the situation is less clear. Its apparent absence may merely reflect the poor public access and consequent difficulty in recording. However the river does change its character in the Vale of Pickering, where it has been straightened, appears to have a swifter flow and the surrounding countryside is dominated by intensively farmed arable landscapes, probably subjected to insecticide sprays. Above East Ay ton the river becomes very fast flowing and with a rocky bottom appears unsuitable for this species. The upstream limit of the Banded Demoiselle would therefore appear to be somewhere between the confluence of the Derwent with the Rye and East Ay ton, which poses the question: what are the determining physical and biotic factors that limit the upstream range of this species? Given the serious floods along the Derwent during November 2000, including the flooding of Malton, Norton, Stamford Bridge, and Barlby, it is probable that pressure will be exerted on the Environment Agency to implement extensive flood defence schemes along parts of the river system. As this may have a negative impact on the population of the Banded Demoiselle, identification of the factors affecting the Banded Demoiselle distribution is therefore of crucial importance. At the other extreme at Barmby the Banded Demoiselle was not found during a quick survey of a short part of the Ouse, although it was found within 100 m of the Barmby Barrage on the Derwent with a total of 75 recorded in grid SE 69-29. The apparent absence of the Banded Demoiselle along the Ouse may reflect either the relative lack of recording or the tidal nature of the Ouse, which may prevent the successful colonisation of this species. Conclusion The River Derwent supports a healthy population of the Banded Demoiselle and indeed this species has been recorded from its confluence with the Rye down to the Ouse. The status of this species in the Vale of Pickering is less clear, but it is certain that somewhere between the Rye and East Ayton the river is unable to support populations of this species. The factors determining this are currently unknown. Appendix 1 Records of the Banded Demoiselle along the River Derwent 1999-2000 Date Site Name Grid Reference Number of Adults District 09/06/99 Derwent (Elvington Bridge Section) SE706477 29 Selby 09/06/99 Pocklington Canal (East Cottingwith Section) SE702427 1 Market Weighton 09/06/99 Derwent (Wheldrake Ings Car Park) SE694445 3 Selby 10/06/00 Derwent (Stamford Bridge) SE71-55- 4 Stamford Bridge 10/06/00 Derwent (Stamford Bridge) SE70-55- 49 Stamford Bridge 10/06/00 Derwent (Stamford Bridge) SE70-54- 25 Stamford Bridge 10/06/00 Derwent (Stamford Bridge) SE71-55- 8 Stamford Bridge 68 Distribution of the Banded Demoiselle Calopteryx splendens along the River Derwent Date Site Name Grid Reference Number of Adults District 10/06/00 Derwent (Stamford Bridge) SE71-56- 36 Stamford Bridge 10/06/00 Derwent (Stamford Bridge) SE72-56- 2 Stamford Bridge 11/06/00 Derwent (Howsham Wood) SE74-63- 5 Malton 11/06/00 Derwent (Kirkham Priory) SE73-65- 5 Malton 11/06/00 Derwent (Kirkham Priory) SE74-65- 8 Malton 11/06/00 Derwent (Howsham Bridge) SE73-62- 4 Malton 11/06/00 Derwent (Kirkham Priory /Howsham Wood) SE74-64- 7 Malton 16/06/00 Derwent (Old Malton) SE80-72- 28 Malton 16/06/00 Derwent (Malton) SE78-71- 4 Malton 16/06/00 Derwent (Malton) SE77-70- 13 Malton 16/06/00 Derwent (Malton) SE77-69- 18 Malton 16/06/00 Derwent (Malton) SE76-69- 28 Malton 27/06/00 Derwent (Upstream from Barmby) SE70-29- 23 Selby 27/06/00 Derwent (Derwent Bridge, Bubwith) SE70-36- 16 Selby 27/06/00 Derwent (Breighton) SE70-34- 34 Selby 27/06/00 Derwent (Breighton) SE70-35- 5 Selby 27/06/00 Derwent (Barmby Barrage) SE68-28- 3 Selby 27/06/00 Derwent (Barmby) SE68-29- 1 Selby 27/06/00 Derwent (Upstream from Barmby) SE69-29- 75 Selby BOOK REVIEW The Flora of County Cavan by P. A. Reilly. Pp. iv + 177 (incl. 2 maps), plus 8 pp. colour plates. National Botanic Gardens, Glasnevin, Dublin. 2001. 17.00 (including postage and packing), available from National Botanic Gardens, Glasnevin, Dublin 9, Ireland, paperback. Although less elaborate than many other recently published county Floras of Britain and Ireland, this is a valuable contribution to our knowledge of a large inland county (746 square miles) of Ireland, all too often ignored in the past by botanists in their desire to study oceanic environments. The Flora covers the charophytes and vascular plants, their distribution based on only five districts, the topography of which are elaborated in the introductory matter; the latter also reviews the physical landscape, including geology (the accompanying map unhelpfully inserted), landscape history and botanical exploration (which contains some particularly useful biographical information). Statistical treatment of the number of taxa found up to 1898 is provided, but the current number of taxa found in the county, recent extinctions, etc. are less apparent. The work also contains almost 50 pages of most useful appendices and indexes on such topics as plant lists previously published or occurring as annotations in books, other documentary and bibliographic sources, areas of scientific interest and a topographical index. All in all, a most useful guide to accompany the field botanist and an important source of background information to past and present botanical activity. MRDS 69 COLEOPTERA REPORT FOR 1995-2001 R. J. MARSH 11 Crusader Drive, Doncaster DN5 7RX This report deals with the Coleoptera excluding the family Carabidae and the sub-family Aleocharinae, and is the first report to be presented since that which appeared in The Naturalist 116:101-106 (1991). Prior to that paper, the late John Flint produced periodic Coleoptera reports in this journal. The records that follow are the substance of annual Coleoptera reports presented to the Entomological Section of the YNU at their indoor meetings during the years 1995-2000, plus additional records for 2001. These reports included new county and vice-county records and other records worthy of note. There are several coleopterists working actively on the beetles of the county, including M. L. Denton, W. R. Dolling, W. A. Ely, M. Hammond, P. Kendall, F. E. Kenington, R. Merritt and E. J. Smith, and my thanks are due to these gentlemen for their dedicated efforts, and to all others who have submitted records to me. The initials used in the following pages are those of K. N. A. Alexander, E. W. Aubrook, L. Auckland, H. Britten, J. D. Coldwell, J. Cooter, H. H. Corbett, M. L. Cox, R. Crossley, J. Denton, M. L. Denton, T. Dixon, W. R. Dolling, W. A. Ely, G. N. Foster, E. F. Gilmour, A. Godfrey, M. Hammond, P. M. Hammond, D. G. Hemingway, J. B. Job, C. Johnson, P. Kendall, F. E. Kenington, R. Lawson, A. Lazenby, B. Levey, C. H. C. Lyal, R. J. Marsh, D. Maude, R. Merritt, M. G. Morris, A. Parker, K. G. Payne, P. A. Seccombe, P. Skidmore, E. J. Smith, T. Stainforth, R. R. Uthoff-Kaufmann, G. B. Walsh and P. F. Whitehead. I express my gratitude to Mike Denton who allowed me to extract data from his card index for Calderdale, which contained many records not on the YNU database, and for offering critical comments on the draft of this manuscript, and to Colin Howes who permitted me to extract data from the Coleoptera records held on record cards at the Doncaster Museum. The thanks of the Yorkshire Naturalists’ Union are due to the many individuals, landowners and organisations that have permitted access to their properties for the purposes of coleopterology during the period. Until 1990 all Coleoptera records were held and maintained by John Flint using a system originally begun by W. J. Fordham and continued by E. G. Bayford, G. B. Walsh, W. O. Steel and others. These records were kept on loose sheets on a species-by-species basis, but owing to the increasing volume of, and detail within, them, this methodology clearly could not be maintained especially when site details were required. After 1990, all these records were transferred to a computerised database system, completed in 1994, and employing RECORDER 3.3, developed by Dr Stuart Ball of JNCC. This site-based system allows an extensive range of rapid information retrieval, of which the old ‘paper’ system was totally incapable. In the list that follows, species nomenclature generally follows that of Kloet and Hincks (1977). I am aware of the nomenclatural changes, particularly within the genus Apion, where the family name Brentidae is gaining acceptance (Gonget, 1997), and the subgenera of Apion listed in Kloet and Hincks (1977) which have now been elevated to generic rank, and I have adopted these changes in this report. Morris (1993) gives corrections to his Handbook (Morris, 1990) and a revised checklist. Morris (1991) also puts forward a ceutorhynchine checklist as a supplement to and modification of that in Kloet and Hincks (1977). Johnson (1990) summarised the known distribution of the Ptiliidae of Yorkshire. Responsibility for any errors or omissions with respect to the list that follows rests entirely with the author. National Status is included where appropriate and the definitions of status categories (JNCC, 1992) are as follows: RDB1 - endangered - taxa in danger of extinction and whose survival is unlikely if causal factors continue operating. Naturalist 127 (2002) 70 Coleoptera Report for 1995-2001 RDB2 - vulnerable - taxa believed likely to move into the Endangered category in the near future if the causal factors continue operating. RDB3 - rare - taxa with small populations that are not at present Endangered or Vulnerable, but are at risk. Notable A - species which do not fall within the RDB categories but are thought to occur in 30 or fewer 10 km squares of the National Grid, or for less well-recorded groups, within seven or fewer vice-counties. Notable B - species which do not fall within the RDB categories but thought to occur in between 31 and 100 10 km squares of the National Grid. The following recording symbols are used: # New county record; * New vice-county record; “recent” = post- 1980. Haliplidae Haliplus heydeni Wehncke - Notable B - (*61) Burton Constable Park (TA1836) 8/5/1999, MH. (*62) Old Malton Moor (SE8174) 9/8/1987, RM det GNF. (*65) Garsdale Head (SD7894) 18/5/1992, RM det GNF. Recent extensive surveying by RM and MH has yielded many new records for the county. Dytiscidae Hydroglyphus pusillus (F.) - Notable B - (*61) Dunnington (SE6753) 9/10/1997, MH. (*62) Rawcliffe Meadows (SE5755) 1 8/5/1998, MH. Recent surveying by MH and RM has yielded many new county records. Coelambus nigrolineatus (von Steven) - Notable A - (*61) North Duffield Carrs (SE6938) 3/11/1999, MH). (*63) Rother Valley (SK4581) 31/7/1995, RM. (*65) Nosterfield (SE2779) 11/9/1998, MH. Copelatus haemorrhoidalis (F.) - (*62) Strensall Common (SE6459) 8/4/1998, MH det MLD. (64) Cawood (SE5737) 23/4/1998, MH, being the first vice-county record since Askham Bog in the 1930s (GBW). Rhantus suturalis (Macleay) - Notable B - (*62) Rawcliffe Meadows (SE5755) 7/10/1997, MLD. Before 1997, this species was very rarely recorded in the county. Extensive survey work by MH and RM has greatly extended the knowledge of the species in Yorkshire. Gyrinidae Gyrinus distinctus Aube - RDB3 - (*61) Leven Canal (TA1045) 24/4/2000, WRD. Welton Waters (SE9525) 24/4/2000, MH. (#63) Beighton Ponds (SK4483) 9/6/1978, WAE. Catcliffe Flash (SK4288) 1995, RM. (*64) Barlow Common (SE6328) 28/4/2000, MH. Hydrophilidae Helophorus strigifrons (Thomson) - Notable B - (*61) Bubwith Bridge (SE7036) 16/4/1990, MLD. (*62) Strensall Common (SE6459) 29/3/2000, MH. (*63) Scout Dike (SE2305) 20/6/1995, JDC. Until 1995 very rarely recorded, but now there are numerous records due to the efforts of MH and RM. Enochrus ajfinis (Thunberg) - (*61) Skipwith (SE6537) 28/8/1995, RM. (*62) Strensall (SE65) 1997, MH. (#63) Upperhead Dike (SE1301) 24/4/1985, DM det MLD. (*64) Askham Bog (SE5748) 26/7/1998, MH. As a result of a species split (Foster, 1982) and confusion with E. isotae Hebauer, the records of E. ajfinis in Yorkshire prior to 1995 (i.e. those dating back to 1908) have to be regarded, in the absence of voucher specimens, as E. ajfinis sensu lato. Authenticated records of E. ajfinis sensu stricto therefore are those given above. Cymbiodyta marginella (F.) - (*62) Strensall (SE6362) 5/9/1997, MH det MLD. (*63) Potteric Carr (SE5900) 1981, GNF. This hydrophilid remained largely unrecorded in Yorkshire since the early captures by Hey and Fordham, with very few other records, until much recent survey work by MH, RM and others has produced many records from all VCs except 65, possibly demonstrating an increase in range and frequency. Coleoptera Report for 1995-2001 71 Histeridae Sphaerites glabratus (F.) - RDB3 - (62) Garbutt Wood (SE5083) 25/4/1999, LA. Duncombe Park (SE68) 6/8/2000, MLD. A rare northern species of decaying fungi and dung. Five records only on the YNU database and unrecorded in the county since 1938. Abraeus granulum Erichson - Notable A - (#63) Worsborough (SE30) 1/1997, DG det JDC (Coldwell, 1997). Rivelin (SK38) 1/9/1998, EJS. The only county records. An indicator of ancient woodland, particularly in red-rotted heartwood of old oaks. Gnathoncus nannetensis (Marseul) - (61) Skipwith Common (SE63) 7/9/2000, DGH det MLD. (62) Helmsley (SE58) 6/8/2000, MLD. (64) Brayton Barff (SE53) 8/9/2000, AG det MLD. A local species from bird nests, carrion, and bracket fungi. Very few records, and unrecorded since 1966. Gnathoncus nanus (Scriba) - (*61) Welwick (TA32) 10/9/1999, FEK det RJM, in mealhouse debris at a farm. (*63) Margery Wood (SE20) 2/7/1988, MLD, in carrion. Another very local beetle of carrion and bird nests. The only other county record is from Pannal Ash in 1937 (RRUK) when the species was taken from carrion-baited pitfalls. Saprinus aeneus (F.) - (63) Hatfield Moor (SE70) 1992, PS. There are only six other, very old (pre-1928), records of this supposedly common species for the county. Hydraenidae Hydraena testacea Curtis - Notable B - (*61) Hornsea Mere (TA1947) 26/10/1996, RJM teste CJ. There has been a marked increase in recent records especially from VC61, being due to the collecting efforts of RM and MH. Ptiliidae Nossidium pilosellum (Marsham) - Notable B - (#61) Danthorpe (TA2432) 31/7/2001 WRD teste CJ. In bracket fungi on a tree stump. Ptenidium intermedium Wankowicz - (*61) Bubwith (SE7036) 14/5/1988 (RJM). Wheldrake (SE7044) 7/9/1989, RJM. (61) Hornsea Mere (TA1947) 26/10/1996, RJM det CJ. This very local and scarce species may be found in well-established good quality wetlands. It has since been found at several sites in VC61 . Euryptilium gillmeisteri Flach - (#64) Fountains Abbey (SE2768) 3/9/1995, RJM det CJ. This is the first British record (Johnson, 2001). Acrotrichis sanctaehelenae Johnson - (#62) Rawcliffe Meadows (SE55) 10/1997, RJM det CJ. Collected from plant refuse at the margin of a pond, and described as new to Britain on specimens from Essex collected in 1985 (Johnson, 1987). Leptinidae Leptinus testaceus (Muller, P. W. J.) - (*63) Treeton Wood (SE4487) 22/9/1997, EJS. T. P. Wood (SE1317) 13/8/2001, MLD. This species is parasitic in the fur of voles and other small rodents and probably under-recorded. Leiodidae Triarthron maerkeli Markel - Notable B - (#62) May Moss (SE89) 7/1996 and 9/1996, LA det MLD. On both occasions in flight interception trap. A leiodid of probably subterranean fungi, and very rarely recorded nationally (Cooter, 1996). Leiodes obesa (Schmidt) - (*63) Eggborough Power Station (SE5824) 3/6/1983, RJM det JC, one specimen on a windowpane in a power station outbuilding. Thorne Moor (SE7415) 10/8/1990, RJM teste JC, a single specimen in a pitfall trap. Associated with subterranean fungi, this species is almost certainly greatly under-recorded owing to its obscure lifestyle. Leiodes triepkii (Schmidt) - (#63) Frickley Park (SE4608) 8/6/1985, RJM det JC, by sweeping vegetation in woodland ride in the early evening; again, associated with subterranean fungi. Agathidium rotundatum Gyllenhal - (*63) Wintersett (SE3714) 29/4/1999, MLD. Bole Edge, Strines (SK29) 4/6/2000, EJS. Very rarely recorded, and associated with rotting wood. 72 Coleoptera Report for 1995-2001 Nemadus colonoides (Kraatz) - (*61) Skipwith Common (SE6437) 7/9/2000, DGH det MLD, being unrecorded since Waterhouse’s 1871 record from Studley (VC64). Typically an insect of bird nest debris in rotting trees. Catopidius depressus (Murray) - Notable B - (#63) Hatfield Moor (SE60) 1992, PS. This is the only county record. Silphidae Dendroxena quadrimaculata (Scopoli) - Notable B - (63) King’s Wood, Elsecar (SK3999) 1997, Coldwell, 1999. Seckar Wood (SE3314) 16/5/1998, MLD. The first county records for 50 years. Staphylinidae Micropeplus tesserula Curtis - (*65) Catterick (SE1697) 19/5/2001, WRD, found by beating fungoid sticks over a sheet. This species is known in the county from only two other records, Sandall Beat (VC63) and Bishopwood, Selby (VC64). Phyllodrepa puberula Bernhauer - Notable A - (*63) Crosland Moor, Huddersfield (SE1214) 16/11/2000, MLD. This, a single specimen found on a window pane in a house, constitutes the only record other than an early 1900’s record from Saltburn. Omalium exiguum Gyllenhal - Notable B - (*63) Melton Wood (SE5103) 1/5/1987, RJM. Blackmoorfoot (SE0912) 21/8/1999, MLD. These are only the second and third county records. Micralymma marina (Strom) - (*61) North Ferriby (SE9826) 12/6/1999, MLD, WRD. A local staphylinid of intertidal shorelines and estuaries where it may be found in cracks in rocks. Ochthephilus andalusiacus (Fagel) - (#62) Helmsley (SE6183) 13/6/2000, RJM det PMH. This is the first Yorkshire record for this inhabitant of riverbanks, particularly in shingle. Ochthephilus omalinus (Erichson) - (*62) Hutton Common (SE7088) 14/8/1993, MLD. Helmsley (SE6183) 13/6/2000, RJM teste PMH, and in company with O. andalusiacus. (*63) Elland Gravel Pits (SE1362) 19/4/2000, MLD. (#64) High Batts (SE3076) 8/4/1993, JBJ det MLD. Platystethus nodifrons (Mannerheim) - (*62) Duncombe Park (SE6082) 29/5/1997, RJM. (*64) Cawood (SE5737) 4/1990, KGP det MLD. Breary Marsh (SE2541) 7/1999, RJM. All the other 20 or so records on the YNU database emanate from several localities in the Lower Derwent NNR in VC61. Stenus solutus Erichson - (*61) Bubwith Bridge (SE7036) 30/12/1989, MLD. (*62) Rawcliffe Meadows (SE5755) 18/4/1998, WRD. All known Yorkshire records are post- 1982, with only a very few records to date. Stenus umbratilis Casey - (*65) Mickle Fell (NY8319) 19/8/2000, WRD. Taken beside a peaty bog pool amongst sedges and previously recorded in Yorkshire only from Malham Tarn in VC64. Leptacinus formicetorum Markel - (63) Hardcastle Crags (SD9630) 1990, RC det MLD. Green Wood (SE2108) 28/5/1998, MLD. The first county records since 1948. There are no other recent records for this associate of the ants Formica rufa and F. lugubris. Neobisnius villosulus (Stephens) - (*62) Helmsley (SE68) 28/6/2000, MLD. Taken by sieving riverbank shingle deposits and only the second county record. Philonthus rubripennis Stephens - (64) Fountains Abbey (SE2768) 8/2000, RJM. The first record for this vice-county since an early Studley record dated 1871 (EAW), and is a species, at least in Yorkshire, apparently restricted to riverside shingle deposits. Philonthus spinipes Sharp - (#63) Wharncliffe Wood (SK3094) 7/2000, EJS teste MLD. The third known British locality for this species, which was noted as new to Britain in 1997 (Allen & Owen, 1997) and subsequently reported from Warwickshire (Lane & Mann, 1999). Platydracus fulvipes (Scopoli) - Notable B - (#62) Staxton (TA0279) 20/5/1998, LA det MLD. The only known county record for this large predatory species. 73 Coleoptera Report for 1995-2001 Ocypus fuscatus Gravenhorst - Notable B - (*63) T. P. Wood (SE1317) 7/2/1981 (MLD). Elland Gravel Pits (SE12) 7/5/1983, MLD. These are the first county records of this rare species since Fordham’s Bubwith record of 1911. Heterothops praevius Erichson - (*61) Elstronwick (TA2332) 14/9/1999, WRD. The first for the vice-county and the first county record since 1967. The other two county records are pre-1900. Mycetoporus despectus Strand - Notable B - (#63) Boothroyd Wood (SE1812) 1/6/1980 (MLD). Blackmoorfoot (SE1010) 2/4/1999, MLD. (*64) Breary Marsh (SE2541) 9/6/1999, MLD. This recently described species has hitherto been confused with the common M. lepidus. The specimen standing as M. lepidus from Boothroyd Wood near Huddersfield, collected in 1980 (MLD), was in 1997 re-examined by MLD and found to be the new species, and the first authenticated record for the county. Mycetoporus punctus (Gravenhorst) - Notable B - (*62) Jugger Howe Wood (SE9399) 8/8/1993, LA det MLD. Fylingdales Moor (SE8897) 8/1996, PMH. Marsh (1991:105) noted that a record of this species from (VC61) Thornton Ellers in 1987 was apparently a new vice-county record. Since then it is noted (Denton, 1995:41) that the species was recorded from Spurn on 15/6/1947 and this latter therefore constitutes the first VC61 record. Sepedophilus immaculatus (Stephens) - (61) Halsham (TA2827) 20/3/2000, WRD, the first VC61 record for 50 years. This follows a spate of recent records in VC63 for this hitherto rarely recorded species. Lamprinodes saginatus (Gravenhorst) - Notable A - (*63) Blacker Dam (SE2804) 9/3/1996, JDC. Gypsy Marsh (SE4102) 30/3/1996, JDC. A species very rarely recorded in the county. Tachinus lignorum (L.) - Notable B - (63) Cawthorne (SE2808) 1986, EJS. The first county record since 1931. SCARABIIDAE Aphodius zenkeri Germar - Notable B - (#61) Allerthorpe Common (SE7657) 5/8/1989, RJM. (*63) Hatfield Moor (SE6906) 1991, PS. These remain the only known county records for this species, said to be associated particularly with deer dung (Jessop, 1986). Hoplia philanthus (Fuessly) - (*63) Brodsworth Hall (SE5007) 23/6/2001, PAS det RJM. Several specimens of this scarabiid were seen around flowers and shrubs. There are only eight other county records, from VC62 and VC64. SCIRTIDAE Elodes tricuspis Nyholm - (#64) Breary Marsh (SE2541) 9/6/1999, RJM. This species, only recently described (Nyholm, 1984), was swept from streamside vegetation. The species has only been recognised from a half dozen or so specimens nationally (Allen, 1993). Prionocyphon serricornis (Muller, P. W. J.) - Notable B - (62) Duncombe Park (SE6082) 2/9/1994, AG, bred from tree rot-hole debris. (#63) Cusworth Park (SE5403) 8/6/1994, AG teste MLD. Bretton Lakes (SE2812) 12/1995, AG, both records again bred from rot-hole debris. Brodsworth Hall (SE5007) 26/6/2000, RJM, swept from ivy at base of old ash. Recent studies of the invertebrate fauna of tree rot-holes by AG have yielded a number of records of this hitherto virtually unknown species in the county. Previous to these studies, there was only a single, unconfirmed, 19th century record from Scarborough. The larvae develop in water-filled rot-holes in various deciduous tree species. Dryopidae Dryops luridus (Erichson) - (*61) Howden (SE7528) 7/1985, PK. Pocklington (SE8049) 9/8/1997, RJM det CJ. (*62) World’s End Plantation (SE6659) 10/1998, MH. (#63) Thrybergh (SK4796) 12/6/1980, WAE. 74 Coleoptera Report for 1995-2001 Buprestidae Agrilus laticornis (Illiger) - Notable B - (62) Rawcliffe Meadows (SE5755) 7/6/1997, MLD. The first county record since 1942, and associated with oak, willow and hazel. Elateridae Fleutiauxellus maritimus (Curtis) - Notable A - (*61) Spurn (TA4112) 17/6/1986, MLD. (64) Grass Wood (SD9865) 1 1/6/1995, JDC. Usually found in shingle beds by rivers, this is a species with a northern and western distribution. Oedostethus quadripustulatus (F.) - Notable A - (62) Rawcliffe Meadows (SE5755) 1997, MLD. The first VC62 record since Fordham’s 1919 capture. Most records are recent ones from VC61, and this is a declining species mainly owing to the loss of its riverside wet meadow habitat. Eucnemidae Epiphanis cornutus Eschscholtz - (#62) Deepdale (SE9291) 7/2000, LA det WRD. Widely distributed nationally but extremely scarce, and associated with spruce logs, this constitutes the first known county record and was taken in a flight interception trap. Anobiidae Ochina ptinoides (Marsham) - (*63) Brodsworth (SE5007) 26/6/2000, RJM. This rare species occurs in ivy, in which it bores the stems, on old trees. There are only four other county records. Hemicoelus fulvicornis (Sturm) - (61) Escrick (SE6343) 17/7/2001, RC det RJM. (*62) Duncombe Park (SE6082) 26/6/1993, MLD. (63) Bretton Lakes (SE2812) 5/6/1999, RJM. (64) Coburnhill Wood (SE4535) 20/7/2001, RJM. These are the first Yorkshire records since 1955 for this mainly southern insect. Dorcatoma flavicornis (F.) - Notable B - (*61) Leven Canal (TA0945) 30/7/1999, WRD, one specimen reported from Salix sp. (*62) Beech Wood (SE5982) 6/8/1995, KNAA. Beningbrough Park (SE5357) 20/7/1999, KNAA. (*63) Hatfield Moor (SE6906) 1992, PS. High Hoyland (SE2509) 30/7/2000, EJS. Of the ten Yorkshire records, eight are post-1977. Anitys rubens (Hoffmann) - Notable B - (*62) Beech Wood (SE58) 6/8/1995, KNAA. (*63) Hatfield Moor (SE60) 1992, PS. There are only three other county records. Melyridae Dolichosoma lineare (Rossi) - Notable B - (#61) Spurn (TA4112) 29/6/1991, MLD, (Denton. 1995). Welwick Saltmarsh (TA3419) 6/8/1999, WRD. A saltmarsh and tidal creek species with a hitherto Kent to Lincolnshire coastal distribution. Nitidulidae Meligethes planiusculus (Heer) - (*63) Orgreave Common (SK4286) 19/7/1995, AL. Treeton Wood (SK48) 4/10/1997, AL. Associated with Echium vulgare Viper’s Bugloss. The only other known county record is that from Mulgrave Woods (VC62) in the 1930s (HB). Cryptarcha strigata (F.) - Notable B - (63) Bretton Lakes (SE2821) 9/6/1986, MLD. The first VC63 record since 1910. (*64) Brayton Barff (SE5830) 19/7/2000, AG det MLD. A subcortical species and associated with the Goat Moth Cossus cossus (L.). Cryptarcha undata (Olivier) - Notable B - (#64) Brayton Barff (SE5830) 23/7/1996, AG det PFW. The same habitat comments apply to this species as with C. strigata ( see above). Specimens have been found on several occasions since 1996 at the same site (AG) and this constitutes the only known Yorkshire locality. Sphindidae Sphindus dubius (Gyllenhal) - Notable B - (*63) Haw Park (SE3615) 12/5/1999, DGH teste MLD. New to the vice-county and the second Yorkshire record, otherwise being known only from Scarborough around 1900 (recorder unknown). Coleoptera Report for 1995-2001 75 CUCUJIDAE Pediacus dermestoides (F.) - (62) Duncombe Park (SE6482) 26/6/1993, PK. The first record for VC62 since 1951. (*63) Bretton Lakes (SE2812) 14/4/1996, JDC. (*64) Fountains Abbey (SE2768) 2/9/1995, MLD. There are now only five county records. SlLVANIDAE Oryzaephilus surinamensis (L.) - (61) Welwick (TA3421) 10/9/1999, FEK. This introduced species of stored products was found in mealhouse debris at a farm site, and was previously only recorded from Hull in 1922 (GBW). Psammoecus bipunctatus (F.) - (61) Hornsea Mere (TA1947) 13/6/1996, RC det RJM and also in 1998 (RJM). Last recorded from this site in 1977, then a new county record. Hornsea remains the only Yorkshire site now known to harbour the species, which may be found in marsh litter and between the stems of Typha and Phragmites. There is an old Doncaster record dated 1911 (HHC), and the species was found in Bronze Age peat deposits on Thorne Moor (PS). Cryptophagidae Telmatophilus schoenherri (Gyllenhal) - (*63) Rushy Moor (SE5712) 14/7/1982, PS. Worsborough (SE3403) 1985, JDC. Rockley Dike (SE3042) 14/7/1985, WAE. These constitute the only known Yorkshire records. Cryptophagus saginatus Sturm - (*61) Welwick (TA3421) 13/9/1999, WRD det RJM. Found in mealhouse debris at a farm. (*63) Thorpe in Balne (SE5911) 10/4/1987, RJM det CJ, in piles of roadside mowings. Blackmoorfoot (SE0912) 16/8/1999, MLD det CJ. Micrambe lindbergorum (Bruce) - (61) Haverfield Quarry (TA32) 16/8/1996, WRD teste CJ. Keyingham (TA2325) 21/8/1999, WRD teste CJ. On both occasions the insects were found on dead thistles, and are the first county records since 1963. Atomaria lohsei (Johnson & Strand) - RDB3 - (*62) May Moss (SE89) 27/7/1996, LA det CJ. (*63) Haw Park (SE3615) 15/8/1992, RJM det CJ. Langsett (SE1900) 9/3/1998, EJS det CJ. A recent arrival in Britain and associated with conifers, either under bark or in wood debris, or in flight interception traps as at May Moss. Atomaria pusilla (Paykull) - (63) Owston (SE5511) 27/7/2000, RJM. Very rarely recorded in the county, and apparently declining with only three post- 1970 records. Erotylidae Tritoma bipustulata (F.) - Notable A - (63) Hatfield Moor (SE60) 1991, PS. Only three other county records, all pre-1966. Phalacridae Phalacrus substriatus Gyllenhal - (*63) Langold Holt (SK5685) 14/7/1984, EJS det CJ. Blackmoorfoot (SE0912) 22/2/1987, MLD. Associated with rusts and smuts on various Carex species, there is an old unsubstantiated record from Scarborough around 1900 (RL). CORYLOPHIDAE Sericoderus lateralis (Gyllenhal) - (*61) Elstronwick (TA2332) 3/10/1997, WRD. (*63) Whitley Bridge (SE5620) 2/11/1990, RJM. (*64) High Batts (SE3076) 24/6/2000, RJM. Until the Whitley Bridge capture there was only one county record of this minute inhabitant of compost heaps, that from Scarborough in 1960 (EWA). The insect can be very numerous when found. Corylophus cassidoides (Marsham) - (61) Hornsea Mere (TA1947) 10/1996, RJM. A Yorkshire rarity of marshland ground litter, this species was last recorded from the same site in 1915 (TS), and has been found in Bronze Age peat deposits on Thorne Moor (PS). There are no other county records. 76 Coleoptera Report for 1995-2001 COCCINELLIDAE Stethorus punctillum Weise - (*62) Forge Valley (SE9886) 8/6/2000, RJM. (#63) Warren Vale (SK4397) 12/7/1996, AL. This minute ladybird is said to feed on Psilacotes mites, and is reputedly common in the south of England, these being the only known county records. Scymnus schmidti Fursch - Notable B - (61) Haverfield Quarry (TA3220) 15/5/1999, WRD det RJM. Last recorded at Spurn in 1950 (WDH) and only the second county record. Latridiidae Aridius australicus (Belon) - (#63) Thorne Waterside (SE6714) 14/8/1982, PS. An introduction (as Lathridius norvegicus Strand) to the British Isles (Allen, 1952), this appears to be the only known Yorkshire record. Enicmus brevicornis (Mannerheim) - Notable B - (#61) West Newton (TA1937) 24/8/1999, WRD. Elstronwick (TA2332) 11/9/1999, WRD. (*63) Blackmoorfoot (SE1010) 3/9/1999, MLD. Found variously by general sweeping in woodland and by sieving piles of twiggy garden refuse. These are the only county records. Cisidae Cis setiger Mellie - (*63) Hatfield Moor (SE6906) 1992, PS. Only two other county records are known. COLYDIIDAE Aglenus brunneus (Gyllenhal) - (#63) Armthorpe (SE6305) 6/8/1966, PS. This, the only known county record, appears to be hitherto unpublished as such, although the insect is included in Skidmore (1983). Tenebrionidae Alphitophagus bifasciatus (Say) - (61) Howden (SE7724) 6/7/1992, PK. The first VC61 record since 1949. (63) Kettlethorpe (SE3216) 9/7/1999, MLD. The first VC63 record since the 1930s. Melandryidae Abdera flexuosa (Paykull) - Notable B - (62) Helmsley (SE68) 5/6/2000, RJM. Forge Valley (SE9886) 8/6/2000, RJM. Rarely recorded in the county, the first VC62 records since 1935 and usually associated with fungi on old willows and alders. Hypulus quercinus (Quensel) - RDB2 - (#62) Helmsley (SE68) 6/2000, RJM and MLD. This saproxylic rarity is restricted to four widely separated localities in Britain, and there is a sub-fossil record from the peat deposits on Thorne Moor. Mordellidae Mordellistena neuwaldeggiana (Panzer) - (*63) Blaxton Common (SE6801) 8/7/2000, PK det BL. (#64) Bishopwood (SE5633) 1/8/1992, PK det BL. Oedemeridae Oedemera lurida (Marsham) - (61) Haverfield Quarry (TA3219) 27/6/1998, WRD. The first VC61 record for over 100 years. (*63) Tingle Bridge (SE3901) 9/6/1993, JDC. Since 1993 there has been a flood of records of this insect, which is clearly extending its range and increasing in frequency. Cerambycidae Arhopalus rusticus (L.) - (#61) Wheldrake (SE7044) 1/8/1996, TD det MLD. A specimen, probably emanating from a nearby conifer plantation, was found in a bird-watching hide on a reserve area. The species is associated with pine plantations in the south of England and in Scotland. Agapanthia villosoviridescens (Degeer) - (#63) Little Moor (SK5287) 10/6/2000, WAE. 77 Coleoptera Report for 1995-2001 Potteric Carr (SE6000) 7/2001, AP teste RJM. These would appear to be the most northerly records for this longhorn. Twinn and Harding (1999) in their BRC Atlas show a record for grid square SE08 in VC65 - this is an error. Saperda scalaris (L.) - Notable A - (*63) Little Don Valley (SE1900) 27/6/1995, JDC. Previously known in the county only from old woodland sites in VC62, the species is mainly northern and western in the British Isles. Bruchidae Bruchus rufimanus Boheman - (63) Thrybergh (SK4796) 5/12/1992, AL. This record was the first for the county since 1951 but since 1992 the species has been frequently reported. Chrysomelidae Oulema melanopus (L.) - (#63) Pollington (SE6119) 5/1998, RJM. (*64) Bishop Monkton Ings (SE3166) 9/1998, WRD. Owing to confusion with O. rufocyanea (see comments under the following entry) all previous records of this species, in the absence of voucher specimens, must be viewed with caution. Oulema rufocyanea (Suffrian) - In a recent paper (Cox, 1995), this species was brought forward as British and distinct from O. melanopus, both species being confused in collections. All Yorkshire captures of ‘ melanopus ’ are now examined critically as a matter of course, and in consequence, the first authenticated Yorkshire specimens of O. rufocyanea may be recorded from the following vice-counties: (*61) Kelsey Hill (TA2326) 19/8/1994, WRD. (*62) Forge Valley (SE9886) 7/9/1996, RJM. (#63) T. P. Wood (SE11) 3/12/1980, MLD. (*64) Bishop Monkton (SE3166) 5/9/1998, MLD. Virtually all Yorkshire ‘melanopus ’ specimens taken in the field prove to be O. rufocyanea. Crioceris asparagi (L.) - (#63) Rossington (SK6298) 7/6/2000, RJM. A pest of cultivated asparagus in some parts of the south of England, this is apparently the first Yorkshire record. The specimen was swept in grassland near allotment gardens where asparagus was being cultivated. Chrysolina oricalcia (Muller, O. F.,) - Notable B - (*63) Falthwaite (SE3003) 19/6/1996, JDC. Handsworth (SK4186) 10/6/2000, AL. There are very few recent records for the county. Chrysomela aenea L. - (*64) High Batts (SE3076) 21/6/1992, MLD, PK. (*63) Canklow Wood (SK4391) 28/5/1990, WAE. Most records for this conspicuous alder Alnus glutinosa feeder are from VC62 and the great majority are post- 1979. The species appears to have become much more frequent throughout the county during the last decade. Calomicrus circumfusus (Marsham) - Notable A - (#63) Denby Dale (SE2108) 8/8/1998, MLD. A species associated with Ulex and Cytisus. Longitarsus anchusae (Paykull) - Notable B - (63) Hatfield Moor (SE6906) 11/8/2000, PS. Associated with Echium species, this is the first recorded occurrence in the county since 1914. There is also an undated Doncaster record (EGB). Longitarsus exoletus (L.) - (61) Hedon (TA2129) 15/6/1999, WRD det MLC. The first VC61 record since the 1930s. (63) Lindrick Dale (SK5482) 9/8/1986, WAE. The first VC63 record since the 1920s. (*64) High Batts (SE3076) 24/6/2000, WRD det MLC. A little recorded species on Echium vulgare. Longitarsus ganglbaueri Heikertinger - Notable A - (*63) Hatfield Moor (SE60) 1992, PS. The first county record for over 60 years. Longitarsus kutscheri (Rye) - (*61) Wintringham (SE87) 23/8/1976, WRD det MLC. Beverley (TA0139) 13/10/1999, WRD det MLC. The first VC61 records for this species which is associated with Plantago. Longitarsus reichei (Allard) - (*61) Brandesburton (TA0947) 24/7/1999, WRD det MLC. (#63) Fan Field (SK5381) 13/9/1980, WAE. A little recorded species in the county. Lythraria salicariae (Paykull) - Notable B - (*63) Hatfield Moor (SE60) 1995, PS. There is only one other (unsubstantiated) county record, dated 1879, from the Whitby area. Psylliodes chalcomera (Illiger) - Notable B - (#63) Elland Gravel Pits (SE1121) 7/6/1990, 78 Coleoptera Report for 1995-2001 MLD. Associated with thistles Cirsium species, this remains the sole county record. Psylliodes laticollis Kutschera (= weberi Lohse) - (*61) Roos (TA3030) 18/4/1999, WRD. (*62) Rawcliffe Meadows (SE5755) 22/2/1997, MLD. (*63) Hipperholme (SE12) 9/5/1980, MLD. All county records are post- 1980 with most being post- 1997. Anthribidae Anthribus fasciatus (Forster) - Notable A - (#63) Medge Hall (SE7412) 8/6/1976, PS. This is the only county record of this species and appears not to have been published previously. Rhynchites cavifrons Gyllenhal - Notable B - (#62) Appleton-le-Moors (SE7387) 25/5/1985, KGP. (*63) Seckar Wood (SE3314) 16/5/1998, MLD. Apparently only five county records for this, one of the “leafroller”, species; it is mainly southern English in its distribution. Rhynchites interpunctatus Stephens - Notable B - (#63) Blaxton (SE6803) 1 1/6/1968, PS. Thrybergh (SK49) 7/7/1985, WAE. Thrybergh (SK4695) 1986, EJS. These remain the only known county records, the Blaxton record not, apparently, having been published previously. The species is another “leafroller”, recorded from oak, and again southern English in distribution. Rhynchites tomentosus Gyllenhal - Notable B - (63) Hatfield Moor (SE70) 1992, PS. The first county record since 1966. Brentidae Pseudapion rufirostre (F.) - (*61) Patrington Haven (TA3021) 3/8/1997, WRD. (*63) Angler’s Country Park (SE3715) 10/5/2000, MLD. According to Morris (1990) this species is locally abundant on Malva species as far north as Durham, although there are only three Yorkshire records. Protapion dissimile (Germar) - Notable B - (#63) Blaxton Common (SE6801) 8/7/2000, RJM det MGM. A species apparently restricted to Trifolium arvense. Ischnopterapion mode stum (Germar) - (#63) Gypsy Marsh (SE4102) 15/8/1995, JDC teste MGM. A species associated with Lotus uliginosus. Protopirapion atratulum (Germar) - (*65) Catterick Garrison (SE1697) 19/5/2001, WRD. Most records are from the south of the county. Hemitrichapion reflexum (Gyllenhal) - Notable A - (#62) Yedingham (SE8979) 2/1951, EFG. (*63) Potteric Carr (SE6000) 24/5/1968, PS. A species feeding on Sainfoin Onobrychis viciifolia. CURCULIONIDAE Sitona cambricus Stephens - (#64) Bishopwood (SE5633) 19/6/1981, RC. (*63) Hatfield Moor (SE6906) 1992, PS. Hatfield Moor (SE6906) 9/9/2000, PS. A scarce Sitona in Yorkshire, associated with Lotus uliginosus. These are the only records. Limobius borealis (Paykull) - Notable A - (*61) Howden (SE7528) 28/7/1987, PK. (64) High Batts (SE3076) 21/6/1992 (MLD), and the first VC64 record for 60 years. A declining species with very few recent records, on various Geranium species. Dorytomus hirtipennis Bedel - Notable A - (*61) Wheldrake (SE6944) 4/9/1997, MLD. (#63) Thorpe in Balne (SE591 1) 6/7/1976, PS. Associated with Salix alba, this species may be declining nationally. Sirocalodes mixtu s (Mulsant & Rey) - Notable B - (#61) North Cliffe Wood (SE8637) 1 1/7/1998, WRD teste MGM. A very local, southern insect on Fumaria officinalis. Neophytobius quadrinodosus (Gyllenhal) - Notable A - (#62) Deepdale (SE9291) 16/5/1998, LA det CHCL. The first authenticated county record - there is an old, unsubstantiated record from Scarborough around 1900 (RL). Anthonomus bitub erculatus Thomson - (*61) Hollym Carrs (TA3224) 20/5/1995, WRD. (*64) Bishop Monkton (SE3465) 1 4/6/1997, RJM. Very few county records exist for this species owing to confusion with other members of the genus. Morris (1977) clarified identification within the group. This species develops in flower buds of Crataegus monogyna. 79 Coleoptera Report for 1995-2001 Curculio rubidus (Gyllenhal) - Notable B - (.*63) Hatfield Moor (SE6906) 9/9/2000, PS. Very scarce nationally, associated with Betula and only the second county record, previously from Goathland in 7/1939 (RRUK). Sibinia primitus (Herbst) - Notable B - (#63) Hatfield Moor (SE6906) 9/9/2000, PS. Associated with Spergularia rubra and possibly others of the genus. The foodplant is quite widely distributed in the south of the county, in sandy areas, but this is the only known county occurrence of the weevil. Miarus graminis (Gyllenhal) - Notable B - (*63) Hatfield Moor (SE70) 1992, PS. The only other county record is that from Pannal Ash dated 5/1936 (RRUK). Scolytus rugulosus (Muller, P. W. J.,) - (#61) Oak Hill (SE7122) 16/6/1987, PK. Elstronwick (TA2332) 3/8/1996, WRD. (*64) Thorpe Arch (SE4446) 25/10/1999, JD. High Batts (SE3076) 24/6/2000, WRD. These remain the only county records. The species is subcortical, mainly on old fruit trees. SCOLYTIDAE Phloephthorus rhododactylus (Marsham) - (63) Lindrick (SK58) 1986, EJS. Hatfield Moor (SE6906) 9/9/2000, PS. These records are the first for the county since 1935. References Allen, A. A. (1952) Lathridius norvegicus Strand (Col., Lathridiidae) rediscovered: an addition to the British list. Entomologist’s Mon. Mag. 88: 282-283. Allen, A. A. (1993) Elodes tricuspis Nyholm (Col., Scirtidae), an addition to the beetles of Windsor Forest. Entomologist’s Rec. J. Var. 105: 90. Allen, A. J. and Owen, J. A. (1997). Philonthus spinipes Sharp (Staphylinidae) in Dorset - new to Britain. Coleopterist 6: 81-83. Coldwell, J. D. (1999) The Insects of the Barnsley Area. Sorby Natural History Society, Sheffield. Cooter, J. (1996) Annotated keys to the British Leiodinae (Col., Leiodidae). Entomologist’s. Mon. Mag. 132: 205-272. Cox, M. L. (1995) Identification of the Oulema ‘melanopus’ species group (Chrysomelidae). Coleopterist 4: 33-36. Denton, M. L. (1995) The Beetles of Spurn Peninsula. Yorkshire Museum, York. Foster, G. N. (1982) Another Enochrus. Balfour-Browne Club Newsletter 24: 1-2. Gonget, H. (1997) The Brentidae ( Coleoptera ) of Northern Europe. Fauna Entomologica Scandinavica, Vol. 34. Brill, Eeiden. Jessop, F. (1986) Handbooks for the Identification of British Insects V part II. Dung Beetles and Chafers. Coleoptera: Scarabaeoidea. Royal Entomological Society, Eondon. JNCC (1992) A Review of the Scarce and Threatened Coleoptera of Great Britain, Part 1. Joint Nature Conservation Committee, Peterborough. Johnson, C. (1987) Additions and corrections to the British List of Ptiliidae (Coleoptera). Entomologist’s Gazette 38: 117-122. Johnson, C. (1990) The Feather- wing Beetles of Yorkshire (Coleoptera: Ptiliidae). Naturalist, 115: 57-71. Johnson, C. (2001) Euryptilium gillmeisteri Flach (Coleoptera: Ptiliidae) new to Britain. Entomologist’s Gazette 52: 181-182. Kloet, G. S. and Hincks, W. D. (1977) Handbooks for the Identification of British Insects XI part 3. Coleoptera and Strepsiptera. Royal Entomological Society, London. Lane, S. A. and Mann, D. J. (1999) Philonthus spinipes Sharp (Staphylinidae) and other species of note from manure heaps in Warwickshire. Coleopterist 8: 92. Marsh, R. J. (1991) Coleoptera records for Yorkshire: a review and update. Naturalist 116: 101-106. Morris, M. G. (1977) The British species of Anthonomus Germar (Col., Curculionidae). Entomologist’s Mon. Mag. 112: 19-40. 80 Coleoptera Report for 1995-2001 Morris, M. G. (1990) Handbooks for the Identification of British Insects V part 16. Orthocerous weevils. Coleoptera Curculionoidea (Nemonychidae, Anthribidae, Urodontidae, Attelabidae and Apionidae). Royal Entomological Society, London. Morris, M. G. (1991) A taxonomic checklist of the British Ceutorhynchinae, with notes, particularly on host plant relationships (Coleoptera: Curculionidae). Entomologist’s Gazette 42: 255-265. Morris, M. G. (1993) 'British Orthocerous Weevils’: Corrections and New Information (Coleoptera: Curculionoidae). Entomologist’s Mon. Mag. 129: 23-29. Nyholm, T. (1984) The identity of Elodes elongata Tournier, 1868 (Col., Helodidae). Entomologica Scandinavica 15: 435-441. Skidmore, P. (1983) The Ecology of Sandall Beat Local Nature Reserve. Directorate of Educational Services, Doncaster Metropolitan Borough Council, Doncaster. Twinn P. F. G. and Harding, P. T. (1999) Provisional Atlas of the longhorn beetles ( Coleoptera , Cerambycidae) of Britain. Biological Records Centre, Institute of Terrestrial Ecology, Monks Wood. BOOK REVIEW Butterflies of Europe by Tom Tolman and Richard Lewington. Pp. 320, with 104 colour plates and numerous distribution maps. Princeton University Press, Princeton. 2002. $26.95, laminated covers. This book is a re-issue for the American market of the Collins Field Guide Butterflies of Britain and Europe, published by HarperCollins in 1997 as a successor to the classic work by Higgins and Riley, A Field Guide to the Butterflies of Britain and Europe, which was published in several editions. The author begins with a tribute to the memory of Lionel Higgins and Norman Riley which is followed by an introductory chapter outlining general butterfly anatomy, the geographical limits covered, life, history, behaviour and conservation of butterflies. The main part of the book follows the style adopted by Higgins and Riley but the text dealing with the description of each species is generally much reduced and even absent for some, much more reliance being placed on the illustrations to effect an identification. In particular no indication of the size of each species of butterfly is given in the text, the reader being informed in the introduction that “the best indication of approximate size is provided by the illustrations”. In contrast, the biological details are much more extensive and includes much useful information, particularly on larval foodplants. The book concludes with a check list of species, glossary and bibliography. The hybrid origin of this publication is emphasized by an interesting anomaly: although the use of the English spelling “colour” is consistent throughout the text, this becomes the North American equivalent “color” on the verso of the title page and back cover. The colour plates, from paintings by the renowned entomological illustrator Richard Lewington, are a delight and the book is well worth having for these alone, although the British reader may prefer the original HarperCollins edition. The underside illustrations are of butterflies depicted in a resting position rather than as “set” specimens and so will greatly aid identification of insects in the field. Despite some shortcomings this is currently the standard general identification guide to European butterflies, with updated distributional information and the inclusion of recently separated taxa, but the reviewer regrets the reduced descriptive content. HEB OBITUARY DR LEWIS LLOYD-EVANS (1916-2001) 81 Born on the 31st of March 1916 at Tenbury, Lewis Lloyd-Evans, ‘Lloyd’, was the son of a GP and grew up in rural Lincolnshire, in the Louth area, and enjoyed country pursuits. He studied medicine at Edinburgh in the years spanning the start of World War 2; he had a fine memory for factual detail, excelling in subjects like anatomy, at which he topped his class. Lloyd enjoyed the company of those who shared his interests and willingly passed on his expertise. He enjoyed his student years. One summer he took a trawler to Iceland to visit Lake Myvatn and on a trip to Dublin met his future wife, Maudie, Maudie, who predeceased him by 10 years, was the daughter of an Anglican minister in a rural parish in the Lough Corrib area of Co. Galway, where she too enjoyed a country childhood - riding, boating and fishing with her two brothers. She was working as a secretary at the Guinness Company in Dublin when they met. World War 2 started whilst he was at Edinburgh and when he qualified he was drafted into the Army Medical Corps. Here he trained in the then experimental technique of blood transfusion, which he put to use after D-Day as he moved across Europe with the advancing allied armies. Lloyd and Maudie married in neutral Eire whilst he was in the Army, and had to borrow friends as his own were not allowed to attend due to the fear of desertion. She moved around southern England with his various postings and whiled away spare time in the countryside enjoying its wildlife. After the war Lloyd became a GP in a practice in Ware, Hertfordshire, and their son Trevor was born. They lived over the surgery, in The Old House in the High Street; it was difficult to get away from work, so at every opportunity the family would go off into the countryside to explore its natural history. They joined the Hertfordshire Natural History Society, which brought them into contact with other enthusiastic naturalists, notably ‘Snaily’ Stratton, who triggered an interest in Mollusca, and John and Chris Dony whom they ferried about Hertfordshire doing fieldwork for the Naturalist 127 (2002) 82 Obituary botanical atlas of the county. This further developed Lloyd’s botanical expertise whilst birds, dragonflies and fungi were other special interests he developed about this time. In this period too, he wrote an authoritative paper on black rats in Hertfordshire {Trans. Herts. Nat. Hist. Soc 25: 4-6). Summer holidays about this time were usually taken camping in Wales, where they added to the list of Welsh dragonflies. In the early 1960s the family ‘discovered’ the recently formed bird-ringing group at Rye Meads and bird-ringing at Rye Meads became a major preoccupation of the family and led son Trevor to become a professional ornithologist. During the Rye Meads years almost all free time was spent there and when birds were scarce attention was turned to the other wildlife of the area. After a long day in the field, we would all be invited back to The Old House for warming coffee, sherry and conversation round the fire and then Maudie cheerfully produced dinner for the assembled company. Often Lloyd enjoyed supervising the preparation of some special dish for dinner and colourful fungi often featured on the menu. During the same period Lloyd took up the study of bird parasites, especially the Mallophaga, Fleas, and Ticks on birds caught for ringing. At home this involved a ‘production line’ of specimens being processed through a succession of clarifying agents, dyes, and mountants in preparation for examination under the microscope. This fitted in well with surgery hours and, in later years, the interest in microscopy led on to the dissection of critical species of Mollusca; he also commenced his study of plant rusts. In the 1960s he also adopted Heteroptera, helping Dr Bernard Nau to map the aquatic Heteroptera of Herts. Another major interest in the 1960s was fieldwork for the Conchological Society’s 1976 Mollusc atlas. He became a member of the Society in 1965. The family remained in Ware until June 1968, when they moved to Holmfirth in Yorks, when Lloyd took up a post in the local schools medical service. They soon became involved with the Yorkshire Naturalists’ Union and, later, Sheffield’s Sorby Natural History Society, regularly attending their field meetings of these societies. This led to new interests in yet more groups, notably plant galls and rusts, and renewed interest in others. His Presidential Address to YNU in 1974 was entitled ‘The biogeography of snails in Yorkshire’. He was also a very active member, and ex-President, of the Yorkshire Conchological Society. Normally, however, he studiously avoided formalities and purely social events. Summer holidays at this time were often taken camping in France and the Spanish Pyrenees, enjoying the many unfamiliar species. Then, in the early 1970s, his son Trevor took up an ornithological post in Massachusetts, USA, which led to a series of holidays exploring the birds and wildlife of various regions of that country. Over the ten years following Maudie’s death, poor circulation and then deteriorating vision gradually restricted his fieldwork; then a stroke, from which he largely recovered, prompted him to move to a nursing home near his sister in Caterham on the Hill in Surrey. By then he could not manage a walk of more than a few hundred yards but he still enjoyed car trips with visitors into the countryside, visiting such sites as the Ashdown Forest, Stodmarch, and the Isle of Sheppey. To the end he kept his mind exercised by completing the daily large crossword puzzle in the Guardian newspaper. He was looking forward to a forthcoming family holiday in Cornwall when he died in his sleep on 14th May 2001. Lloyd was elected to the Union at the AGM in Sheffield held on the 6th December 1969, and I met him and his wife Maudie in the early autumn of the same year, having had advance notice of his intended semi-retirement to Yorkshire. One of the very last of the all round naturalists, Lloyd will be missed by all who knew him. I personally have lost not just a friend in Lloyd but a second family in which Maudie took centre stage, and one in which I always remember the hospitality and kindness given to me over the years. I would like to thank Dr Bernard Nau for much of the above biographical information and Lloyd’s son Trevor for forwarding the picture of Lloyd and Maudie on one of their trips to the USA. A. Norris Obituary 83 Selected Bibliography Lloyd-Evans L. (1956) Ena montana (Draparnaud) in Hertfordshire. J. Conch. 24 (3): 102. Lloyd-Evans L. (1956) Acroloxus lacustris (L.) in Merionethshire. J. Conch. 24 (3): 104. Lloyd-Evans L. (1957) Vertigo substriata (Jeffreys) in Hertfordshire. J. Conch. 24 (6): 208. Lloyd-Evans L. (1958) Acanthinula lamellata (Jeffreys) in Pembrokeshire. J. Conch. 24 (7): 252. Colville, B., Lloyd-Evans, L. and Norris, A. (1973) The occurence of Boettgerilla pollens simroth, a slug new to Britain, in the English Lake District. Naturalist 98: 70. Colville, B., Lloyd-Evans, L. and Norris, A. (1973) Notes on the occurence of Boettgerilla pallens Simroth in the English Lake District (New to Britain). Concholo gists’ Newsletter 44 (March): 305-306. Lloyd-Evans, L. and Nore, B., (1973) Y.N.U. Excursion Report for Entomology, Wilton Woods and Teesmouth (VC62) 26-28 May 1973. Naturalist 98: 139. Lloyd-Evans, L. (1974) Y.N.U. Excursion Report for Mollusca, Boroughbridge (VC64) 29- 30 June 1974. Naturalist 99: 152. Lloyd-Evans, L. (1975) The biogeography of snails in Yorkshire. Naturalist 100: 1-12. Colville, B., Lloyd-Evans, L. and Norris, A. (1974) Boettgerilla pallens Simroth a new British species. J. Conch. 28: 203-207. Lloyd-Evans, L. (1980) Y.N.U. Excursion Report for Mollusca, Painsthorpe and Uncleby (VC61)21 June 1981. Naturalist 106: 188. Lloyd-Evans, L. (1980) Y.N.U. Excursion report for Plant Galls, Painsthorpe and Uncleby (VC61) 21 June 1981. Naturalist 106: 120. Lloyd-Evans, L. (1980) Y.N.U. Excursion Report for Mollusca, Beckermonds and Oughtershaw (VC64) 19-20 July 1980. Naturalist 106: 125. Lloyd-Evans, L. (1980) Y.N.U. Excursion Report for Mollusca, Garsdale (VC65) 5 July 1980. Naturalist 106: 128. Lloyd-Evans, L. (1982) Y.N.U. Excursion Report for Mollusca, Scaling Dam (VC62) lb- 17 May 1981. Naturalist 107: 103. Lloyd-Evans, L. (1983) Y.N.U. Excursion Report for Mollusca, Rise Park (VC61) 12 June 1982. Naturalist 108: 113. Lloyd-Evans, L. (1983) Y.N.U. Excursion Report for Mollusca, May Moss (VC62) 19 June 1982. Naturalist 108: 116. Lloyd-Evans, L. (1987) Y.N.U. Excursion Report for Plant Galls, Potter Brompton (VC61) 21 June 1986. Naturalist 112: 150. Lloyd-Evans, L. (1987) Y.N.U. Excursion Report for Mollusca, Goathland (VC62) 28-29 June 1986. Naturalist 112: 151 . Lloyd-Evans, L. (1987) Y.N.U. Excursion Report for Plant Galls, Goatland (VC62) 28-29 June 1986. Naturalist 112: 152. Lloyd-Evans, L. (1987) Y.N.U. Excursion Report for Plant Galls, Sunk Island (VC61) 17 August 1986. Naturalist 112: 158. Lloyd-Evans, L. (1988) Y.N.U. Excursion Report for Mollusca, Austwick and Lackland Mosses (VC64) 30 May 1987. Naturalist 113: 152. Lloyd-Evans, L. (1988) Y.N.U. Excursion Report for Plant Galls, Downholme Park (VC65) 27 July 1987. Naturalist 113: 163. Lloyd-Evans, L. (1989) Y.N.U. Excursion Report for Plant Galls, Bridestones (VC62) 223 August 1987. Naturalist 114: 76. Lloyd-Evans, L. (1989) Y.N.U. Excursion Report for Plant Galls, Netherby Dale (VC62) 28 May 1988. Naturalist 114: 144. Lloyd-Evans, L. (1990) Obituary: Fred Stubbs. Naturalist 115: 29. Lloyd-Evans, L. (1995) Y.N.U. Excursion Report for Plant Galls, Skipwith Common 17 July 1993. Naturalist 120: 78-79. Lloyd-Evans, L. (1995) Y.N.U. Excursion Report for Plant Galls, Malham Tarn 6 and 7 June 1993. Naturalist 120: 86. 84 BOOK REVIEW Pennak’s Freshwater Invertebrates of the United States. Porifera to Crustacea. Fourth edition by Douglas Grant Smith, x + 638 pp. New York. Wiley. 2001. £85.95 hardback. New editions of Pennak’s ‘Freshwater Invertebrates of the United States’ 1952 appeared in 1978 and 1989. The third omitted the insects. This fourth edition, revised by D. G. Smith, omits the Protozoa, but is still a substantial volume. A somewhat idiosyncratic but useful introduction deals with freshwaters in general, and their invertebrates, embraces a miscellany of topics, and includes much interesting information and a few errors. Here, although there is a separate section on distinctive larval stages, an accompanying table omits the unique larvae of the branchiuran Chonopelds and the various remarkable larvae of mutelid bivalve molluscs. The anomopod branchiopod labelled as Streblocerus is in fact Lathonura , and a table listing anabiotic stages omits the remarkable African mutelids of the genera Aspatharia and Spathopsis. The latter can firmly close and hermetically seal the shell valves, and regularly withstand complete desiccation of the habitat, and high temperatures throughout the dry season. Unionids may survive drought for a time, as mentioned in the relevant section, but are not specialised for this, and sometimes suffer mass mortality when the habitat dries out. The bulk of the work deals with the many groups of freshwater invertebrates and provides keys to the genera recorded in the U.S. A general, often substantial, account helpful to students everywhere introduces each group: the keys are specific to the U.S. It should hardly be necessary to say that these should not be used to identify animals elsewhere, but N. American works have been so used in the past, especially when it was believed that many small freshwater animals were cosmopolitan. This leads to confusion. The generously illustrated keys strikingly reveal how much richer is the N. American freshwater fauna than our own; for example, the U.S. boasts no fewer than 81 and 57 genera of gastropod and bivalve molluscs respectively, 1 2 genera of crayfishes that include more than 300 species, about 130 isopods, and 150 amphipods. It is often unfair to refer to omissions as the author may have made a conscious decision to omit some particular point, but one such is surprising. No mention is made of the amazing imitation fish formed from the mantle and marsupium of some bivalve molluscs of the genus Lampsilus that not only look like a fish, complete with eye, but undulate like one. This attracts real fishes that receive from the mollusc a discharge of glochidia larvae which, if they can, attach themselves and become parasitic. The key does say that the female “displays from a partially buried position” and a plate, one of a set new to this edition, actually shows an imitation fish (not a particularly good example) but makes no mention of this amazing mimicry. There are occasional mis-spelled or incorrect names, the ostracods Heterocypris incongruens and Cyprinotus incongruens are the same animal, and an update which tells us that in the 1990s crayfish in Louisiana occupied about 46,000 hectares is immediately followed by a statement that in that state they occupy about 7,000 hectares. English readers will know what is meant when they read that ostracod valves are “united on the dorsal margin by an elastic band” but the phraseology is unfortunate. Illustrations are abundant, almost always clear and helpful, and now include photographs taken via scanning electron microscopy, but sometimes give no indication of size. The book is beautifully produced, will be a boon to North American users, and a means of broadening the knowledge of students everywhere. GF BIRDS ON THE SPURN PENINSULA by Ralph Chislett Parts I and II (1996), edited by Michael Densley. Hardback 218 pages with coloured dust jacket, coloured frontispiece and twelve photographic illustrations in black and white. This enlarged edition is the first time that Part II has appeared in print. Not surprisingly this book virtually sold-out remarkably quickly, and only recently the printers have discovered a small quantity of the remaining books. Before releasing them on the open market, the publisher has offered this limited stock to YNU members for the same price as when it was published in 1996 - £14.95 per copy post free. Owing to a slight mix-up at the time, the newly released book has not previously been advertised in The Naturalist - thus members now have a chance to purchase a copy. The James Reckitt Charity of Hull and other individual benefactors have made a substantial contribution towards the publishing cost in order to keep the selling price relatively low. Apply to: Clive Varty, 61 St Chads Court, St. Chads Road, Leeds LS16 5QX Titus Wilson Kent Works • Burneside Road • Kendal • Cumbria • LA9 4RL Tel. 01539 720244 Specialist printers /binders of Academic Journals , Catalogues and Private Publications Our service includes attending to worldwide distribution Binding Why not have your copies of The Naturalist bound into volumes? One year’s issues per volume, or alternatively two years in one volume at less cost than binding as two separate volumes. We are also experienced in binding thesis and experts in the re-binding and repairing of all books. Spink & Thackray Broomfield Binder Back Broomfield Crescent Telephone: 0113 2780353 LEEDS LS6 3BP Fax 0113 2304155 www . spinkandthackray .co.uk Printed in Great Britain by Titus Wilson & Son, Kendal ISSN 0028-0771 Latest publication of the Yorkshire Naturalists’ Union ALIEN PLANTS OF YORKSHIRE GEOFFREY WILMORE The first comprehensive modem publication dealing with the alien plants of any major British county, detailing the flora (almost 1300 species) of Yorkshire’s wool waste dumps, railway sidings, sewage works, shoddy fields, industrial wasteland, dockland and oil mill sidings. Pp. 316, including 15 line drawings; published January 2000 Normal price: £15.00 (plus £2.00 per copy p.&p.) Reduced price for members of the YNU: £12.00 (plus £2.00 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Mr John A. Newbold, Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS THE FRESHWATER CRUSTACEA OF YORKSHIRE a faunistic & ecological survey GEOFFREY FRYER The crustacean fauna of Yorkshire reflects the great physiographic diversity of the region. Adopting an ecological approach, this book considers the Yorkshire fauna in relation to climate, topography, geology, soils and water chemistry, always keeping in mind that it is dealing with living organisms whose habits, requirements and physiological limitations determine exactly where they live. Matters covered include the ecological background; faunal assemblages and their regional attributes; an analysis of the factors that determine distribution patterns, many of which are mapped; wide geographical aspects; and conservation. Large areas, such as the Pennines, Howgill Fells, North Eastern uplands and the lowland plains are surveyed. So too are localised regions including Whernside, the Malham area, lowland heaths, and the largest lakes, as well as habitats such as upland tarns, seepages, cold springs, small lowland ponds, inland saline waters. Notes are given on every species recorded, including parasitic forms. Price £8.00 (plus £2.00 p.&p.) Special offer to members of the Yorkshire Naturalists’ Union £6.00 (plus £2.00 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Professor M. R. D. Seaward, Department of Environ- mental Science, University of Bradford, Bradford BD7 1DP. July-September 2002 Number 1042 A QUARTERLY JOURNAL OL NATURAL HISTORY LOR THE NORTH OL ENGLAND Finding Fungi - Gillian M. Brand Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat Felis catus Lin. in Yorkshire - Colin A. Howes Published by the Yorkshire Naturalists’ Union Editor M. R. D. Seaward MSc, PhD, DSc, ELS, The University, Bradford BD7 1DP Notice to Contributors to ‘The Naturalist’ Manuscripts (two copies if possible), typed double-spaced on one side of the paper only with margins at top and left-hand at least 2.5cm wide, should be submitted. Latin names of genera and species, but nothing else, should be underlined. S.I. Units should be used wherever possible. Authors must ensure that their references are accurately cited, and that the titles of the journals are correctly abbreviated. Volumes of The Naturalist for the years 1886 to 1975 have been retrospectively numbered 11 to 100 to accord with numbering before and after this period (see YNU Bulletin no. 3, pp. 21-22 1985); please cite these volume numbers in all references. Table and text-figures should be prepared on separate sheets of paper. Drawings and graphs, drawn about twice the linear size they are to appear, should be in jet-black Indian ink, and legends should not be written on the figures. Publishable manuscripts not conforming to the above requirements will be returned for alteration. Photographic Plates Readers of The Naturalist will have noticed that the number of photographic illustrations has increased in recent years. Good clear photographs, suitably captioned, to accompany articles or as independent features are always welcome. To encourage this development, a long-standing member of the YNU, who wishes to remain anonymous, has most generously offered to make a donation, the income from which would finance the publication of a plate or equivalent illustration in future issues whenever possible. The editor, on behalf of the YNU, wishes to record this deep appreciation of this imaginative gesture. © Yorkshire Naturalists’ Union — 2002 Single copies may be made of single articles in this journal provided that due acknow- ledgement is made and the copies are for non-profit making educational or private use. Copying of more than one article or multiple copying of a single article is forbidden unless special permission has been obtained from the Yorkshire Naturalists’ Union. Permission is granted for the use of brief quotations in published work provided that acknowledgement of the source is clearly stated, but the use of substantial sections of text and any illustrative matter requires the express permission of the Yorkshire Naturalists’ Union. Registered Office (for all enquiries, correspondence and payments): Mr John A. Newbould, Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS (tel: 01305-837384; email: janewbould@aol.com) The Naturalist is issued free to individual members of the Yorkshire Naturalists’ Union and to Affiliated Societies. Institutions and Subscribers £24.00 Registered Charity No. 224018 FINDING FUNGI GILLIAN M. BRAND 85 Presidential address presented to the Yorkshire Naturalists’ Union at Ripon, 1st December 2001 Yorkshire has an outstanding and continuing record of achievement in the study of its fungi. The pioneer studies of James Bolton in the second half of the 18th century preceded the formation of the Yorkshire Naturalists’ Union and, at a meeting in Selby in 1896, the founding of the British Mycological Society. Over the years, the YNU has sponsored books on Yorkshire fungi by Massee and Crossland (1905), Mason and Grainger (1937), and Bramley (1985). A key to the continuity of mycological studies in the YNU has been the handing on of the torch of knowledge from generation to generation right up to the present day. Although born and brought up in Yorkshire, I have spent much of my life in exile in Warwickshire and have been involved in the Warwickshire Fungus Survey since its conception in 1965. My aim here is to describe some experiences in trying to find and record fungi at the levels of vice county, site, habitat and species and to extract from these some of the special features of the natural history of fungi. Vice County The Warwickshire Fungus Survey was set up at the time of completion of the field work for mapping the distribution of flowering plants in the county, which led to the pioneer use of computer technology for map production (Cadbury et al., 1971). The initial aim was to map the distribution of fungi similarly. At that time, and with some notable exceptions including in Yorkshire, fungal studies tended to be neglected by local naturalists. One major reason for this neglect was the lack of comprehensive identification books for almost all groups of fungi. Taxonomic work continues and there is much more literature now (Brand et al., 2001). Recently, mostly in the 1990s and still continuing, there has been a national explosion in the formation of local fungus recording groups. Together with already existing groups a large part of the UK is now covered by at least 36 groups. Most of these are connected with the development by the British Mycological Society of a national database of fungal records. This has huge potential for increasing our understanding of fungal distribution but at present its national coverage is patchy, due not only to absence of records from some areas but also to logistic problems in database entry. The basic ground rule of the Warwickshire Fungus Survey was that every new county record in the survey must be supported by a herbarium specimen with field and descriptive notes. These specimens were scrutinized by a local panel and difficult species were sent to national experts. Identification of species was, and continues to be a major problem and the help of experts, at the national botanic gardens at Kew and Edinburgh and elsewhere, has been crucially invaluable. The introductory courses revealed a hard core of more than a dozen enthusiasts who stayed with fungal recording and together covered a good range of taxonomic groups. Table 1 shows the numbers of species of non-lichenized fungi recorded after 12 years, when preparation for publication of The Fungus Flora of Warwickshire ( ed . Clark, 1980) began. In the absence of comprehensive and up to date national check lists for most groups of fungi the best evaluation of coverage is comparison with well-recorded regions. In comparison with S.E. England, the favoured collecting area of workers at Kew, and Yorkshire, the numbers of species in most taxonomic groups were substantial but there was a weakness in recording microfungi. This was largely attributable to the absence of accessible identification literature until the publication of Microfungi on Land Plants (Ellis & Ellis, 1985). Despite the limited range of ecosystems in an inland county with no land above 260 m, further work in Warwickshire since the 1980 publication has resulted in a continued increase in the number of species recorded, with an overall increase of nearly 20% up to the year 2000 (Table 1 , records of the Warwickshire Fungus Survey). Naturalist 127 (2002) 86 Finding Fungi Class Teliomycetes: Urediniomycetes (rust fungi) and Ustilaginomycetes (smut fungi). Microfungi: a non-taxonomic aggregation of the anamorphic fungi (Hyphomycetes and Coelomycetes) with the Plasmodiophoromycetes, Mastigomycotina and Zygomycotina. Finding Fungi 87 FIGURE 1 Mean number of species of Discomycetes and agarics (Agaricales, Boletales, Russulales and Cantharellales) recorded per day by Malcolm Clark in 1976 (open columns) and in 1977 (hatched columns). There were no recording days in weeks 24, 25, 28 and 35 in either year. Limiting factors in regional recording include the problem of surveying a large area, and searching at the correct time of year, when the fungi are fruiting. Traditionally most fungus forays were held in the autumn but sample data from the late Malcolm Clark’s mycological diaries for 1976 and 1977 illustrate the number of species that he found and identified microscopically throughout the year (Figure 1). His special interest was Discomycetes but he also recorded other ascomycetes and agarics. He looked for fungi on an average of 2.5 days every week except when abroad on holiday. There is a marked difference between the all-year-round occurrence of Discomycetes and the sporadic, mainly autumnal fruiting of agarics in the litter habitats in which he searched. This pattern was evident in both years although they had very different rainfall profiles. 1977 was overall a wet year with 688 mm rain, 42% more than in 1976, but with two dry months on either side of torrential rain in September. This sort of experience has stimulated much more year-round recording (Legg, 1989) and is especially important for species which do not fruit in the autumn season. Regional studies can be combined to look at the national distribution of individual species and Figure 2 shows the post- 1965 distribution of two species at the level of vice county. The British Mycological Society’s Fungus Record Database (BMSFRD) shows only positive records and lack of a record could be due either to deficiencies in recording or to absence of the fungus. Geopora (= Sepultaria ) sumneriana is a large, spring-fruiting Discomycete which is associated, possibly mycorrhizal, with Cedrus. Fruit bodies occur particularly on bare loose soil under Cedrus and we have found it at the same locations in Warwickshire in successive years. The original version of Figure 2 suggested that this is a southern species with its northern limit in the Warwickshire area. However, after this address was given, Alan Legg told me that the fungus has occurred annually since 1981 in Darlington West Cemetery, Co. Durham (Legg, 1995). Further enquiries at the national herbaria at Kew and Edinburgh have confirmed that they have no specimens from Scotland or Wales and that the Darlington site is the only one in the north of England. This Finding Fungi Post- 1965 records of Geopora sumneriana and Fomes fomentarius in vice counties, based on the BMSFRD, with additions from Tom Hering, Alan Legg and Kerry Robinson. Darlington find greatly extends the known distribution of G. sumneriana and suggests that it would be well worth looking for it in Yorkshire, under cedar trees in the springtime. Fomes fomentarius is a polypore which grows on upright and fallen trunks, especially of Betula and Fagus. For recording it has the advantage that the perennial fruit bodies can be seen at all times of year. It is widely distributed in northern Britain and records from midland and southern regions are mainly in the east (Figure 2). It occurs on both major hosts throughout its range but is more common on Betula in highland Scotland and on Fagus in south-east England. In Yorkshire it occurs at widely scattered sites and has been recorded from prehistoric sites (Watling, 1978). However, searching in Warwickshire and nearby counties to the west (Herefordshire, Shropshire, Worcestershire, personal communications with members of local fungus groups) has failed to find it, suggesting a puzzling south-westerly limitation on the occurrence of this fungus. Single Site Earlswood, Warwickshire is a large site (c.35 ha) on acid soil of Quercus woodland with Ilex and Betula and containing streams and marshy areas with Alnus, Betula and Corylus. There are also mature planted Fagus and Castanea. This has been one of the favourite sites for the Warwickshire Fungus Survey’s annual weekend courses tutored by national experts, with a visit in late September or early October in almost every year. The procedure was for Finding Fungi 89 No. of years for which species have been recorded FIGURE 3 Records of fungi from Earlswood 1977-1999: (a) accumulation of species records, including recognized varieties, and (b) yearly frequency of Basidiomycete species. Solid circles, all species; hollow circles, Basidiomycetes. individuals to roam freely, searching for the diversity of fungal species (‘wandering sheep technique’) for 2-3 h and then to spend several hours on identification in the laboratory. Although there had been 12 forays before publication, subsequently there was a continuing increase in the number of species recorded from the site, with a 45% increase between 1980 and 1999, of which approximately two-thirds were new species of Basidiomycetes, the most intensively studied taxonomic group (Figure 3a). Inspection of the species and participant lists provides anecdotal evidence of possible explanations for this continuing increase. There are human factors, including different search routes. Bovista plumbea was restricted to the cindery ground of railway property through which the site was entered in the early years. It was recorded in most of these years but not afterwards, when the site was entered from another side. Most people, with a few notable exceptions, have difficulty in seeing certain fungi such as the small pale stalk-like Macrotyphula junceus. The only year in which this was recorded at Earlswood was that in which our specialist spotter of this species was amongst the participants. New literature for identification progressively becomes available. Thus four species in the Armillaria mellea complex have been distinguished at the site since publication of the Warwickshire Flora. New management activities can introduce new species. Wood chippings imported for path maintenance provided the substrate in 1996 for Stropharia aurantiaca , one of the species characteristic of this material (Shaw & Kibby, 2001). The most important human factor was the presence of the expert tutor. Although the taxonomic group selected for special study on each course was rarely present in abundance at the particular time, many species in this and other groups were confirmed by the experts. Fungal factors in the continuing increase in species numbers over time, e.g. rarity, are more difficult to discern. The large fruit bodies of Xerocomus (= Boletus ) parasiticus are readily seen but have rarely been recorded in either Warwickshire or Yorkshire. The species is unmistakable since its fruit bodies are always found attached to those of Scleroderma citrinum. Although S. citrinum was recorded at Earlswood on 10 out of 17 post-publication visits, X. parasiticus has only been found in one year. A more objective way of describing frequency is to present the data in terms of the number of years in which each species has been recorded at the site. Over the 17 years since 1978 in which forays have been held at Earlswood, the frequency of Basidiomycete 90 Finding Fungi species takes the form of a hollow curve (Figure 3b) in which a large proportion, i.e. 46% of the total of 338 species, were found only once and less than 2% were found in three- quarters or more of the years. These common species were Piptoporus betulinus, Mycena galericulata, Russula ochroleuca, Amanita rubescens, Paxillus involutus and Lactarius quietus. Although the Earlswood data are subject to limitations in species identification as well as to sampling deficiencies, similar results have been obtained in other investigations. After 21 autumn seasons of recording by means of walk surveys in a Caledonian pinewood by an expert taxonomist, the total species accumulation curve over successive years was still continuing to increase, with no indication of levelling off (Tofts & Orton, 1998). Out of 502 species of agarics and boleti recorded, 34% were found only once and less than 3% were found on at least three-quarters of the annual visits. Straatsma et al. (2001) worked in five fixed plots, each of 300m2, in a mixed broad-leaved and conifer forest in Switzerland and recorded every fruit body of every species of ground-living macromycete every week from May to December (weeks 21-50) for 21 years. This rigorous weekly sampling of the plots yielded a similarly shaped annual frequency curve in which 37% of the 408 species were only recorded once. However, the weekly samples resulted in a greater proportion of species (10%) being found in at least three-quarters of the years. Eight species were recorded every year, i.e. Lactarius blennius , Russula cyanoxantha, R. fellea, R. fageticola, R. ochroleuca , Xerocomus badius, Collybia butyracea and C. dryophila. The immediate conclusion is that it takes many years to record the total mycota of a site, or, to put it another way, there is endless fascination in looking at the diversity of fungi at one site. This conclusion exposes the problems involved in mapping species distribution and in determining the conservation value of a site from one or a small number of visits. The inconstancy of fruiting of many larger fungi explains the experience of the Warwickshire Fungus Survey, and others, that trying to map fungi by intensive recording during a single visit to each km square is unrewarding because of the low number of species ‘hits’ per visit. For site assessment and comparison adequate measurement of ecological diversity is important (Magurran, 1988) but is difficult for fungi. However, it is now recognised that fungi are crucial to the functioning of ecosystems. The first report on important fungus areas in the UK (Evans et al., 2001) uses four criteria for selecting areas, i.e. the occurrence and abundance of rare species, the recorded species richness, the importance for fungi of the habitat type and views on the special potential of nominated under-recorded sites. There is currently much debate amongst mycologists about the most efficient method of evaluating the fungal diversity of a site. Single site data raise questions about the complex meaning of rarity in fungi. The first problem is that the vegetative part, the mycelium, within the substratum is difficult to see. The actual substratum may be difficult to pinpoint and the extent of individuals in both space and time is rarely observable. Molecular methods have been used to solve this problem in a few economically important species. In Armillaria bulbosa (= A. gallica) and A. ostoyae, which attack roots of trees and spread underground as rhizomorphs, clonal individuals have been shown to extend over large areas of ground, up to 15 ha (Smith et al., 1992; Dettman & van der Kamp, 2001). From studies of rhizomorph extension rates it has been estimated that the largest individual is 1500 years old. On the other hand, experimental studies have shown that Trametes (= Coriolus) versicolor colonizes stumps and branches by airborne basidiospores, each successful colonizer being a new sexually generated individual, often with a distinctive colour pattern on its fruit bodies (Rayner & Todd, 1978; Williams et al, 1984). The second problem concerns the occurrence of fruiting on mycelia. How long does it take a growing mycelium to reach a fruiting state? How often and for how long is it capable of producing fruit bodies? How long do individual fruit bodies last? What is the effect of environment on all these phases? The detailed data of Straatsma et al. (2001) show that both the number of species fruiting and the abundance of fruit bodies of all species varied dramatically from year to year. There was a ten-fold difference between the lowest and highest number of species fruiting per year (mean 101, s.d. 52) and the range of Finding Fungi 9 1 fruit body abundance per year was almost 50 times (mean 3392, s.d. 2528). There is much to find out about the reproductive strategies of individual species. Single Habitat Grassland supports a different range of fungi from woodland. Bond (1981) details the occurrences of 83 species of macrofungi on mown grassland in his garden. A distinctive assemblage of species, including the often brightly coloured waxcaps i.e. Hygrocybe spp., occurs on unimproved grassland, i.e. that which has not been ploughed and to which no fertilizers, other than faeces of grazing animals, has been added. There is good evidence that many species of unimproved grassland are lost on addition of fertilizers; for example, Arnolds (1989) showed that annual addition of an artificial fertilizer containing NPK and Ca(NC>3)2 and also liquid manure to a study site resulted in an immediate and permanent loss of 10 species of larger fungi, whereas some other species increased in abundance. This loss of certain fungal species occurred earlier than the loss of flowering plant species. In Nordic countries these sensitive fungi have been used as indicators of the conservation value of grassland sites on the basis of two criteria, firstly the number of species of Hygrocybe s.l. and secondly the range of indicator species in the taxa Hygrocybe s.l., Clavariaceae, Leptonia (now part of Entoloma ) and Geoglossaceae (Raid and Nitare respectively, both cited in Boertmann, 1995). During and since World War 2 there has been a massive loss of unimproved grassland in Britain, especially in the south and in 1996 a UK “waxcap” survey was begun (Rotheroe et al., 1996). Preliminary information from this survey indicates that Britain is relatively rich in sites of high conservation value (Holden, 2000; Rotheroe, 2001). Good sites include not only upland hay meadows and pastures but also long established mown grassland e.g. at some National Trust properties, and in graveyards (Fortey, 2000). The distinctive pink species, Hygrocybe calyptriformis, was placed on the provisional UK red data list of fungi (Ing, 1992) as ‘vulnerable’. We now know that it is widespread, especially in northern Britain. The species is very rare in much of Europe (Henrici, 2001) and Britain is a major repository of it. It appears to fruit sporadically so that information on the times and abundance of fruiting in relation to site management will be particularly useful for conservation. The grassland in our garden is a much more modest site, the cumulative total of six species of Hygrocybe s.l., including one recognized variety, placing it in the ‘of local interest’ category on Raid’s scale. However, the laying of turf on some former borders in 1992 has enabled a comparison to be made between the fungi occurring on newer and older grassland areas. The garden was established on former arable land on heavy clay soil overlying Liassic deposits and grass seed was sown between 1955 and 1958. Until about 1970 treatments (fertiliser, weedkiller, lawnsand etc.) were applied to maintain a high standard of garden lawn. At that time all lawn treatments ceased and subsequently the grassland has been mown lightly and infrequently with a rotary mower and the mowings collected. This old grassland consists of separate north (180 m2, including a large bush of Juniperus communis ) and south (64 m2) areas adjacent to the house. The borders, which were turfed in 1992, are located adjoining two sides of the north area (each 20 m2) and as a separate east area (58 m2). The bought-in turf was initially herb- and moss-free, suggesting that it had been treated with chemicals, including weedkiller, but after ten years under the same mowing regime it resembles the older grassland areas in consisting of a moss- and herb-rich grassland community. The pH of the surface soil is 6.4 and that of the imported turf was initially 7.2 and by now is not appreciably different from that of the older grassland. The areas have been scanned for fungal species richness during routine garden surveys throughout the year, with more frequent recording in the tenth year. The older grassland contains a diversity of fungal species (Table 2), including in 2001 six species in Hygrocybe s.l. and five other indicator species. The most abundant Hygrocybe was H. psittacinus , with a maximum per recording day in 2001 of 90 (north) and 29 (south) fruit bodies. Many of the other species are characteristic of grassland. 92 Finding Fungi TABLE 2 List of basidiomycete species in different ecological categories recorded in old and turfed grassland in north (N), south (S) and east (E) locations in a garden 1992-2001 (nr: location not recorded) Category Old grassland Location Turfed grassland Location Indicator species Hygrocybe s.l. Dermoloma cuneifolium N S Hygrocybe chlorophana N H. conica N S H. psittacinus N s H. virgineus N s H. virgineus var. ochraceopallida s Clavariaceae Clavaria acuta N s Clavaria acuta E C. incarnata N C. rosea var. subglobispora N Clavulinopsis fusiformis nr C. helvola N C. laeticolor N Ramariopsis kunzei N Entoloma ( Leptonia ) Entoloma sericellum N s Other species associated with grassland Arrhenia retiruge N Conocybe rickeniana N E Bolbitius vitellinus nr Coprinus auricomis E Calocybe carnea N Camarophyllopsis schultzeri N Clitocybe fragrans N C. rivulosa N Conocybe rickeniana N s C. tenera s C. vexans nr Coprinus friesii s C. leiocephalus N s C. plicatilis s C. rhombisporus s C. xantholepis s Crinipellis stipitarius N Entoloma sericeum N s Galerina clavata N s Marasmius graminum N Mycena aetites N s M. filopes N s M. flavoalba N s M. leptocephala N s Panaeolus foenisecii s P. olivaceus s Rickenella fibula s R. setipes s Vascellum pratense N s Species with other associations Hebeloma crustuliniforme N Armillaria mellea E Tubaria confragosa N Coprinus atramentarius E T. conspersa N C. comatus E T. furfuracea N Hebeloma crustuliniforme N Telamonia sp. N 93 Finding Fungi TABLE 3 Numbers of species in different categories (see text) occurring on old and turfed grassland in successive years Years Old grassland Turfed grassland All Indicator Hygrocybe All Indicator Hygrocybe 1992 11 6 4 0 0 0 1993 6 4 1 1 0 0 1994 10 2 2 0 0 0 1995 5 3 2 1 0 0 1996 4 1 1 1 0 0 1997 18 8 4 2 0 0 1998 14 8 3 2 0 0 1999 13 6 3 2 0 0 2000 10 5 2 2 0 0 2001 28 11 6 6 1 0 occurring on humose soil, grass and other herbaceous debris, amongst bryophytes or on dung fragments (Arnolds, 1989). A few are associated with trees, either mycorrhizal or growing on woody debris (Table 2). On the older grassland there is little if any evidence of progressive change in the occurrence of indicator species over the ten years. On the other hand, the turfed areas have been virtual mycological deserts for ten years (Table 3). All of the earlier-occurring species on these areas are not true grassland species. Coprinus comatus is characteristic of disturbed ground and C. atramentarius grows from buried wood. Hebeloma crustuliniforme is a mycorrhizal species, which has fruited since 1997 as an expanding ‘tethered’ ring (Gregory, 1982) in association with the outer margin of the root system of a tree of Tilia cordata which was planted near the edge of the grassland in the late 1980s. Of the species first recorded in the tenth year, Telamonia is also mycorrhizal and Armillaria mellea was growing in the roots of a tree stump, whereas Conocybe rickeniana and Clavaria acuta are grassland species and also occur on the older grassland. C. acuta is a member of the Clavariaceae, one of the groups indicating unimproved grassland. So far it has only been found on the east turfed section, which is not in physical contact with any other grassland. There are serious design deficiencies if this is regarded as an experiment since the plot sizes and positions are not consistent between the two treatments. However, the species differences between the older grassland and the turf, both that adjoining the north grassland and the separated east patch, are very large and consistent over years. These observations indicate the long time span taken for grass turf to acquire a diverse fungus community. The fruiting of mycorrhizal and wood decomposing species suggests that the turf is not toxic to fungi and focuses attention on the nature and availability in this community of minerals and substrates for these sensitive fungi. Management policy is a major challenge for those responsible for wildlife reserves. For woodland, Kirby (1998) advocates monitoring the results of procedures such as coppicing in order to improve our understanding and decision making. Since many fungi grow on plant and other debris, there is a need for recording the effects of management procedures on the fungal component of different communities. Single Species Studies of the occurrence of fruit bodies of an individual species are useful both for understanding their mode of life and as a prelude to mapping their distribution. There is much variation between species, from the perennially-growing brackets of e.g. Ganoderma 94 Finding Fungi adspersum to the very short-lived toadstools of Coprinus friesii, which expand overnight from the button stage, shed their spores and are beginning to collapse by the following morning. These features are not always obvious to casual observation. The apparently fragile fruit bodies of Sarcoscypha austriaca have been observed to grow from the same fallen 40 mm diameter branch of Populus every year for, so far, seven years. Each year the same individual fruit bodies persist for three months (late December to March). The species selected for special consideration here is a rust fungus, a member of the Urediniomycetes. The rust fungi are parasitic on living tissue of green plants and are unusual amongst fungi in that individuals, i.e. individual infections, can often be seen and counted, which is useful for analysis. The YNU has a long tradition of vice county field meetings bringing together people interested in all types of living organisms. In Warwickshire, ‘Rings’, a network for all naturalists, was set up only recently. It was at the very first joint field meeting of ‘Rings’ on 5 July 1997 that a fungus growing on living leaves and fruits of Crataegus was drawn to our attention by John Robbins, a plant gall expert. This fungus was new to us. The infections could be seen on the upper sides of leaves as red spots up to 5 mm across each with a yellow margin and gradually blackening from the centre, where tiny dark swellings could be seen with a hand lens. From the underside of leaf infections and on developing fruits there were finger-like projections more than 1 mm long. The first problem with a strange fungus is identification. The size and wall ornamentation of the spores in the projections showed that it was a rust fungus. In Britain this group of fungi is well documented (Henderson, 2000). They often have complex life cycles forming up to five successive spore types which in many species alternate between two different types of host plant. The finger-like projections on the Warwickshire find are characteristic of the aecia of the genus Gymnosporangium (Table 4). According to Wilson and Henderson (1966) two species of this genus occur on Crataegus in the Britain. G. clavariiforme is TABLE 4 Host ranges of Gymnosporangium species (Henderson, 2001) Species Aecial host Telial host G. clavariiforme Crataegus spp. Juniperus communis G. confusum Crataegus spp. Cydonia oblonga Mespilus germanicus J. sabina cult. G. cornutum Sorbus aucuparia S. sargentiana S. x thuringiaca J. communis G. sabinae Pyrus communis J. sabina cult. frequent and completes its life cycle by forming telia on Juniperus communis , whereas the alternate host of the rare G. confusum is the garden plant Juniperus sabina. Our specimen had diagonal ridges of wall ornamentation on the peridial cells enclosing the aecia, a diagnostic feature for G. confusum. Plowright described G. confusum in 1889 after carrying out infection experiments in which inoculations with germinating teliospores from J. sabina resulted in aecial infections on Crataegus monogyna, C. laevigata , Cydonia oblonga and Mespilus germanica. He did not report finding the aecial stage in nature. There are a very few authenticated records from Britain, mainly from the south of England, and the most recent previous record was in 1944. Moore (1945) described the aecia of G. confusum on Mespilus and telial infections on J. sabina in 1943 and 1944 at a location in East Mailing Research Station from which 95 Finding Fungi Wormald had observed the fungus on Mespilus in 1928. In Yorkshire there are recent records of G. clavariiforme and G. cornutum on both their aecial and telial hosts (Yeates and Legg, personal communications). However, the only records of G. confusum and G. sabinae (= G. fuscunri) date back to Massee at the beginning of the 20th century. Massee recorded the aecial stage of G. confusum on Crataegus in Sedbergh cemetery in 1912 (Yeates, pers. comm.), having previously recorded the other species, G. clavariiforme, on Crataegus before 1905 in the Scarborough area (Massee & Crosland, 1905). After the initial discovery in Warwickshire, G. confusum was found in the same year on Crataegus in our garden hedge in Stratford-upon-Avon and also near Rugby, where it had been photographed in 1994 by Peter Cook. Infections on Crataegus have been found during early summer in every subsequent year and were abundant in the year 2000, when a distribution survey was carried out. In Warwickshire Crataegus is widespread in older mixed hedgerows as well as in recently planted quickthorn hedges and fragments of older hedges persist around the town of Stratford-upon-Avon. The infections were best seen on long unclipped flowering shoots at the time when the fruits were swelling and such shoots were the basis of the sampling procedure. The survey aimed to cover all km squares containing flowering Crataegus in the urban area of Stratford-upon-Avon and more distant sites in different compass directions from the town. The individual infections are conspicuous, visible at a distance of 0.5 m, and at each sample site, usually a hedge, the plants were inspected from a maximum of ten positions beside the bushes. Failure to see any infections in that sample would have given a negative score. In fact there were no • • • N t •• • • • • 1 • * • 4 < 4 > • > %• •• /•l 3 i ... '-..W • • j # • FIGURE 4 Distribution of the aecial stage of Gymnosporangium confusum on Crataegus in and around the urban area of Stratford-upon-Avon in 2000. The urban area is enclosed by a dotted line and star symbols mark ‘hot spots’. Scale: 1 square = 1 km x 1 km. 96 Finding Fungi negative scores and the fungus was found at every one of 32 sites situated in the 16 one km squares encompassing the urban area of Stratford-upon-Avon (Figure 4) as well as at 48 more distant sites in Warwickshire and in the borders of neighbouring counties (Oxfordshire, Staffordshire and Worcestershire). Infections were found on ornamental garden Crataegus and also on medlar ( Mespilus germanica ), which conveniently grows in a public place in Stratford-upon-Avon, the Shakespeare tree garden. This widespread distribution emphasised the need to find the source of the infections on Crataegus. The aeciospores cannot re-infect Crataegus and instead infect the alternate host, Juniperus sabina. The telia are said to be formed in the spring but searches on accessible J. sabina in our own garden and elsewhere had failed to find them. Mapping the density of infections, which is a reflection of the density of spores deposited on host plants, can be used to indicate the likely locations of the spore source on the alternate host. The pattern of dispersal and deposition of airborne spores, including those of rust fungi, has been much studied and in a more or less uniform topography characteristically takes the form of a hollow curve, with the majority of spores being deposited close to the source (Ingold, 1971). If the source is distant there will be very little difference in infection frequency between nearby sites, although the density of infections could be very high if the distant source is massive. However if the major source is local there will be a steep gradient in infection frequency between nearby susceptible hosts. During the Warwickshire survey it was observed that individual infections seemed to be much more abundant at six locations (described as ‘hot spots’) than elsewhere. At one of these, Bishopton Lane, infrequently clipped remnants of the old field hedge containing hawthorn still exist whilst 33 prostrate conifer bushes, possibly J. sabina, were discernible through the hedge in adjacent gardens and amenity plantings. The number of infections per 300 mm length of overwintered hawthorn shoot, including all leaves and fruits growing from this length and its new shoots was counted in June. There were sometimes significant differences in the abundance of infections on adjacent bushes, but these were small in comparison with the differences between more distant bushes (Figure 5). The gradient of change in infection frequency was steep near the position of maximum score (the ‘hot spot’) and fell to scarcely discernible at greater distances. Such analysis enabled likely FIGURE. 5 Mean number of infections of Gymnosporangium confusum on shoots of Crataegus at different positions near ‘hot spots’ at Bishopton Lane and Shottery. The vertical bar shows the s.e.m. of the mean for five bushes in the hedge (with five shoot lengths per bush), except at the two positions of high density at Shottery, each of which was a single large bush. 97 Finding Fungi positions of source J. sabina to be pinpointed. Permission to search was obtained for four out of the 6 ‘hot spots’ and in every case telia were found in April 2001. After rain the telia appeared as triangular gelatinous masses up to 1cm long protruding from spindle-shaped swellings on the twigs. These telia readily dropped off, or in dry conditions dried to tiny horny black masses. One hot spot was initially puzzling, in a garden-free industrial estate in the centre of Stratford-upon-Avon, until it was realized that a large container planting consisted of J. sabina , which was infected. It thus seems that the fungus is completing its life cycle locally and that the local climate has been suitable for its persistence for at least four and probably seven or more years. Last spring we appealed in the journal Field Mycology (Brand & Brand, 2001) for people to look out for it elsewhere, but we have not heard of any other finds. The species is said to be introduced in Britain (Henderson, 2000) and the origin of its current occurrence in Warwickshire is puzzling. It occurs widely in Europe, extending as far south as north Africa and eastwards into Asia (from Iran to India) and has become established in California (Laundon, 1977). This suggests that England is at the northern limit of its range. There is no obvious connection between the current occurrence in Warwickshire and the last recorded finding in the south of England more than 50 years ago. The prostrate forms of J. sabina have been popular garden plants since World War 2, being advertized as slow-growing ground cover plants. With diminishing garden sizes and changing fashions this rather aggressively spreading conifer is seen less in gardens but continues to be used in amenity plantings. There has also been widespread destruction of old hedgerows. The suburban parts of Stratford-upon-Avon may be unusual in the close proximity between remnants of infrequently clipped hawthorn and J. sabina, which permits successful completion of the life cycle of the fungus. The size of some of the spindle- shaped swellings on the stems of J. sabina suggests that they have persisted for at least several years. Henderson and Bennell (1979) report that Gymnosporangium sabinae, another uncommon introduced species, was found in an Edinburgh garden in 1974 on a plant of J. sabina which had been acquired from a nursery in the south of England. It is tempting to speculate that the Warwickshire population of G. confusum similarly had its origin in infected nursery stock of J. sabina. Conclusion These examples show that there is so much to find out about the natural history of fungi, both about the diversity of fungi in regions or sites and about the activities of individual species. Interesting fungi can be found in urban as well as rural areas (Legg, 1995; Yeates, 1999). Your past President and professional mycologist, Professor Roy Watling, has written recently about the role of amateurs (Watling, 1998). This has always been important. However, today, with the progressively diminishing number of professionals, the activity of amateurs is becoming crucial. A theme throughout this address has been the value of continuing studies over years. Most professional contracts are for a maximum of three years so that the ability of amateurs to continue observations over longer periods is important. Long may the YNU maintain the continuity of mycological studies in Yorkshire. Acknowledgments I am indebted to all past and present members of the Warwickshire Fungus Survey team, and especially the late Malcolm Clark, for their records. I am grateful to staff at the Royal Botanic Gardens at Edinburgh and Kew and to officials and members of the British Mycological Society, the Yorkshire Naturalists’ Union and local fungus recording groups for much helpful information. I thank my husband Bert Brand for his unfailing encouragement, and all the many people who have shared with us their interest in fungi. References Arnolds, E. (1989) The influence of increased fertilization on the macrofungi of a sheep meadow in Drenthe, the Netherlands. Opera Bot. 100: 7-21. 98 Finding Fungi Boertmann, D. (1995) The Genus Hygrocybe. Fungi of Northern Europe 1. Svampetryk, Greve. Bond, T. E. T. (1981) Macro-fungi on a garden lawn, 1971-1978. Bull. Brit. My col. Soc. 15: 99-138. Bramley, W. G. (1985) A Fungus Flora of Yorkshire 1985. Mycological Section, Yorkshire Naturalists’ Union, Leeds. Brand, A. W. and Brand, G. M. (2001) Have you seen this? A Warwickshire oddity. Fid My col. 2: 66-67 . Brand, A. W., Henrici, A. and Leonard, P. (2001) Guide to the Fiterature for the Identification of British Basidiomycetes. British Mycological Society, Kew. Cadbury, D. A., Hawkes, J. G. and Readett, R. C. (1971) A Computer-mapped Flora: a study of the County of Warwickshire. Academic Press, London. Clark, M. C. (ed.) (1980) A Fungus Flora of Warwickshire . British Mycological Society, London. Dettman, J. R. and van der Kamp, B. J. (2001) The population structure of Armillaria ostoyae and Armillaria sinapina in the central interior of British Columbia. Can. J. Bot. 79:600-611. Ellis, M. B. and Ellis, J. P. (1985) Microfungi on Land Plants. Croom Helm, London. Evans, S., Marren, P. and Harper, M. (2001) Important Fungus Areas: a provisional assessment of the best sites for fungi in the United Kingdom. Plantlife, London. Fortey, R. (2000) Old churchyards as fungal conservation areas. Fid Mycol. 1: 121-123. Gregory, P. H. (1982) Fairy rings; free and tethered. Bull. Brit. Mycol. Soc. 16: 161-163. Henderson, D. M. (2000) A Checklist of the Rust Fungi of the British Isles. British Mycological Society, Kew. Henderson, D. M. and Bennell, A. P. (1979) British rust fungi: additions and corrections. Notes R. Bot. Gdn Edinb. 37: 475-501. Henrici, A. (2001) Notes and Records. Fid Mycol. 2: 15-16. Holden, L. (2000) Conservation Comer: Scottish Waxcap Survey. Fid Mycol. 1: 77. Ing, B. (1992) A provisional Red Data List of British Fungi. Mycologist 6: 124-128. Ingold, C. T. (1971) Fungal Spores: their liberation and dispersal. Clarendon, Oxford. Kirby, K. (1998) Judging woodland management by tradition or by results. In: M. A. Atherden and R. A. Butlin (eds) Woodland in the Landscape: Past and Future Perspectives, pp. 43-59. Leeds University Press, Leeds. Kirk, P. M., Cannon, P. F., David, J. C. and Stalpers, J. A. (eds.) (2001) Ainsworth and Bisby’s Dictionary of the Fungi 9th ed. CAB International, Wallingford. Laundon, G. (1977) Gymnosporangium confusum. CMI Descriptions of Pathogenic Fungi and Bacteria no. 544. Commonwealth Mycological Institute, London. Legg, A. (1989) Collecting throughout the year. Mycologist 3: 139-141. Legg, A. (1995) Urban fungi - a few words of encouragement. Mycologist 9: 50-51. Magurran, A. E. (1988) Ecological Diversity and its Measurement. Croom Helm, London. Mason, F. A. and Grainger, J. (1937) A Catalogue of Yorkshire Fungi. A. Brown, Hull. Massee, G. and Crossland, C. (1905) The Fungus Flora of Yorkshire. A. Brown, Hull. Moore, M. H. (1945) A note on medlar cluster-cup rust ( Gymnosporangium confusum Plowr.) in Kent in 1943 and 1944. Trans. Brit. Mycol. Soc. 28: 13-15 and Plate I. Plowright, C. B. (1889) A Monograph of the British Uredineae and U stilagineae . Keegan Paul Trench, London. Rayner, A. D. M. and Todd, N. K. (1978) Polymorphism in Coriolus versicolor and its relation to interfertility and intraspecific antagonism. Trans. Brit. Mycol. Soc. 71: 99- 106. Rotheroe, M. (2001) A preliminary survey of waxcap grassland indicator species in South Wales. In: D. Moore, M. M. Nauta, S. E. Evans & M. Rotheroe (eds) Fungal Conservation: issues and solutions, pp. 120- 135. Cambridge University Press, Cambridge. Finding Fungi 99 Rotheroe, M., Newton, A., Evans, S. and Feehan, J. (1996) Waxcap-grassland survey. Mycologist 10: 23-25. Shaw, R J. A. and Kibby, G. (2001) Aliens in the flowerbeds: the fungal biodiversity of ornamental woodchips. Fid My col. 2: 6-11. Smith, M. L., Bruhn, J. N. and Anderson, J. B. (1992) The fungus Armillaria bulbosa is amongst the largest and oldest living organisms. Nature 356: 428-431. Straatsma, G., Ayer, F. and Egli, S. (2001) Species richness, abundance, and phenology of fungal fruit bodies over 21 years in a Swiss forest plot. Mycol. Res. 105: 515-523. Tofts, R. J. and Orton, P. D. (1998) The species accumulation curve for agarics and boleti from a Caledonian pinewood. Mycologist 12: 98-102. Watling, R. (1978) The distribution of larger fungi in Yorkshire. Naturalist 103: 39-57. Wading, R. (1998) The role of the amateur in mycology - what would we do without them! Mycoscience 39: 513-522. Williams, E. N. D., Todd, N. K. and Rayner, A. D. M. (1984) Characterization of the spore rain of Coriolus versicolor and its ecological significance. Trans. Brit. Mycol. Soc. 82: 323-326. Wilson, M. and Henderson, D. M. (1966) British Rust Fungi. Cambridge University Press, Cambridge. Yeates, C. S. V. (1999) Park and garden mycology. Bull. Yorks. Nats. Un. 31: 15-17. BOOK REVIEWS Dictionary of the Fungi edited by P. M. Kirk, P. F. Cannon, J. C. David and J. A. Stalpere. Pp. xii + 655, inch 40 figures (line drawings). CAB International, Wallingford. 9th edition. 2001. £49.95 hardback. Although this latest edition has only 39 more pages more than the 8th edition published in 1995, it contains about 500 new entries and many revisions to keep pace with the rapidly expanding knowledge of these organisms; these include a refined classification of fungal phyla based on the latest molecular evidence, including a major revision of the Ascomycota, complete integration of anamorphic genera in the classification, and a revised synopsis of the proposed classification. Entries provide the most complete listing available of generic names of and terms used to describe fungi. Extended entries (occasionally supported by line drawings) are provided for a wide range of topics, such as conservation, lichens, literature, media, molecular biology, mycorrhizae, numbers of fungi (currently given as 80060 spp.), phylogeny, soil fungi and spore discharge. Most entries cite bibliographical sources for further information. An essential handbook for all those who work on fungi, lichens and slime moulds. MRDS Fungal Conservation. Issues and Solutions edited by D. Moore, M. M. Nauta, S. E. Evans and M. Rotheroe. Pp. x + 262. Cambridge University Press. 2001. £65.00 hardback. This is an interesting collection of papers on a group of organisms, excluding lichens (which are not covered in this book), which have hitherto received little attention in terms of their conservation. The book has a breadth of coverage, but, as with most multi-author works, the overall structure and direction are not clear; furthermore, although it purports to provide ‘constructive guidance for the management of nature in ways beneficial to fungi’, recommendations for such are hard to find, and even practical conservation doesn’t figure highly in most contributions - so all-in-all, the content does not reflect the title. The work 100 Book Reviews is supported by a rather weak index. However, there is a much to be gained from this work, and although much is based on European experiences, contributions on Cuba, Kenya, Ukraine and China provide a global context, in some cases highlighting the decline in wild fungal populations due to the increase in commercial harvesting. Those employed in conservation will require something more practical for guidance, and those interested in conservation, who should be encouraged to read this book, will be appalled at the price. MRDS The Lichen Flora of Devon by B. Benfield. Pp. 102, inch numerous maps & 1 p. colour plates. Privately published. 2001. £10.00 paperback (A4 format), plus £1.00 postage, available from Glyn Benfield, Plymtree, Cullompton, Devon XE15 2JY. This is a welcome contribution to our knowledge of the lichen flora of Devon, especially since the last comprehensive list for this large and lichenologically rich county appeared as long ago as 1883. The present flora covers more than 900 species, those with interesting distributions complemented by computer generated maps; notes on changes in occurrence are also provided for some species. Introductory matter includes sections on the history of lichenology in Devon, its climate, geology and topography, as well information on major habitats, the dyeing industry, and lists of red data and ecological continuity lichens. Maps (including 8 national distributional ones of rare species), a page of colour plates and a gazetteer add to the usefulness of the volume. The work concludes with a list of references, but the citations are incomplete. MRDS Protocols in Lichenology edited by I. Kranner, R. F. Beckett and A. K. Varma. Pp. xvi + 580, inch numerous figures. Springer- Verlag, Berlin. 2002. £74.00 softcover. This practical guide pulls together a corpus of useful methological material from a wide variety of sources. Thirty-two chapters, contributed by specialists in their fields, provide detailed descriptions of one or more protocols necessary for the experimental lichenologist. The contributions are of variable quality and mainly for laboratory-based work. The sub- title to this work, “culturing, biochemistry, ecophysiology and use of biomonitoring”, is patchily addressed: the first two subject areas are very well covered, and the third reasonably well (although only one chapter is devoted to on-site measurements, and the pioneering work of Otto Lange is given scant attention), but the fourth is poorly covered. Two of the three chapters on biomonitoring are related to radionuclide and heavy metal pollution, while the third, on sulphur dioxide pollution, is rather out-dated ( vide the references provided) and misses recent developments, more particularly in respect of decreasing levels of this pollutant. One or more chapters on monitoring other pollutants which for some countries are more influential than sulphur dioxide would have been a valuable addition. (To redress this, the recent Monitoring with Lichens - Monitoring Lichens edited by P. L. Nimis et al. published in the NATO Science Series by Kluwer should be consulted.) On the other hand, there are some very useful chapters, not implicit in the sub-title, on macrophotography, herbarium management, computer-aided identification systems and on-line documentation of lichen biodiversity. An unfortunately large number of typographical errors, particularly in bibliographical citations, have been noted, and the short index does not do justice to the contents. Despite these misgivings, the volume under review contains a wealth of practical advice (in terms of materials, procedures and trouble-shooting, often supported by extensive lists of references), some of which will have wider relevance beyond the lichenologist’s requirements. Although only in paperback, the flexi-covers are reasonably robust and the book conveniently opens flat for laboratory use. Unfortunately, at this price, there will be few copies dedicated to this use and using copies from libraries (which are strongly recommended to purchase this work) will be inconvenient. MRDS 101 RED IN TOOTH AND CLAW: 2. STUDIES ON THE NATURAL HISTORY OF THE DOMESTIC CAT FELIS CATUS LIN. IN YORKSHIRE COLIN A. HOWES Doncaster Museum & Art Gallery DN1 2AE Part 2 of the Presidential Address to the Yorkshire Naturalists' Union at Doncaster, 3rd December, 2000 Preface The massed ranks of our household pets have become a hugely significant though as yet little studied ecological influence in our man-made environment. Pet bedding and regular sleeping places can harbour a range of ectoparasites, the ubiquitous cat flea ( Ctenocephalides felis ) perhaps being the most obtrusive. Dried pet foods in pellet or biscuit form can attract a menagerie of invertebrate ‘pests’ of stored foods (Munro 1960). Exotic seeds imported for the pet and wild bird food market have given rise to a fascinating flora of aliens and adventives (Wilmore 2000). The daily product of tonnes of surface deposited dog or shallowly buried cat faeces is the focus of coprophagous beetle and fly faunas and attracts an interesting fungal mycota. Endoparasites such as the tape worms (cestodes) Taenia taeniaeformis and Dipylidium canium occur in 3% and 2% of cats respectively, the former introduced via a rodent element in the diet (Lewis 1987) and the latter via its intermediate host the cat flea. Roundworms (nematodes) Toxocara cati and Toxiascaris leonina occur in the intestines of 20% and 1% of cats respectively (Tabor 1983). Eggs picked up and ingested from handled faeces develop into larval stages which can migrate to organs such as the eyes or brain of a human host. The protozoan Toxoplasma gondii which in Britain occurs in common rat ( Rattus norvegicus), wood mouse ( Apodemus sylvaticus ) and bank vole ( Clethrionomys glareolus) and whose final host is the domestic cat (Cox 1987) is the causal agent of the condition in humans known as Toxoplasmosis. Oocysts excreted in cat faeces can be transmitted to those of us who potter in gardens where cats defecate; the Communicable Diseases Surveillance Centre monitored some 621 cases in 1978 (Southam 1981). There is also the matter of ‘what the cat brought in’, the succession of small mammals, song birds and other vertebrates harvested from local habitats when the domestic cat’s latent behavioural drive to hunt is aroused. Under these circumstances, even the neighbour’s goldfish and the family hamster can fall as prey. Cats in domesticated form ( Felis catus Lin.) are with us in very substantial numbers (see Table 3) as household pets or companion animals, as semi-domestic ‘working’ cats in farms, factories etc., and as free-living feral cats, famously around dockyards (Dards 1981), hospitals and other public institutions (Rees 1981, Howes 1985). Curious to know something of the natural history, population structure and predator-prey relationships of free-range cats in Yorkshire, a series of questionnaire surveys were designed to investigate domestic cats on farms and in town centre, suburban and rural situations. The first to be presented here concerns an investigation into Yorkshire’s farm cats. A Survey of Farm Cats in Yorkshire “We rat catchers might raise our fees, Sole guardians of a nation’s cheese!” The Rat-catcher and Cats John Gray (1733) Methods With the enthusiastic assistance of members of the Yorkshire and Humberside Federations of Young Farmers’ Clubs, a survey of farm cat populations throughout the Yorkshire regions was conducted from September to December 1979. Using a standard questionnaire Naturalist 127 (2002) 102 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat sheet, volunteers recorded the type and size of the farm, the number of cats kept there and, where known, their age, sex and whether they had been neutered. Methods of rodent control and reasons for keeping cats on farms were also recorded. Most of the farms surveyed were the homes of the volunteer surveyors or the homes of their friends or neighbours. In order to detect local variations in the size and structure of cat populations in such a large and geographically varied region as Yorkshire, the resultant data were divided for separate analysis into the following areas according to topographical and agricultural characteristics: Pennines, Vale of Mowbray, Southern Vale of York, Humberhead Levels, and the North York Moors, Vale of Pickering, Wolds and Holderness. Results Appendix 2 shows the numbers of farms surveyed, frequencies of various farming practices, farm acreages, number of cats, reasons for their presence, methods of rodent control and perceptions as to the effectiveness of cats for this. Appendix 1 provides statistics on gender and rates of neutering, and for those cats for which age data was provided, Appendix 3a classifies intact and neutered males and females into age classes. Frequency, size and density of farm cat populations Of the 239 farms surveyed, 226 (95%) had one or more cats. The numbers of cats per farm is shown in Figure 1 and varies from one to an alleged thirty. The most frequent colony sizes were from two to five cats (on 58% of farms), the county mean being 4.8. FIGURE 1 Cat colony sizes on 239 Yorkshire farms. Figure 2, which compares the mean colony sizes across Yorkshire farming regions, shows marked variations and indicates an intriguing progressive increment of colony size from south and west (3 on Pennine farms) to north and east (7 on the farms of the North York Moors). By comparing the numbers of cats with the declared farm acreages, a mean farm cat population density was shown to be one cat per 52.7 acres (21.3 ha). Again, Figure 2 shows marked variations, the smaller mixed or stock rearing farms of the North York Moors and the Pennines recording one cat to 21.3 acres (8.6 ha) and 40.5 acres (16.4 ha) respectively, compared with the situation on the huge arable farms of the Humberhead Levels and the Wolds recording one cat per 76.2 acres (30.8 ha) and 78 acres (31.6 ha) respectively. Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 103 FIGURE 2 Mean population sizes and population densities of cats on Yorkshire farms. Age ranges The ages of 558 cats (48% of the total sample) for which gender and neutering information was available are shown in Appendix 3a and form the basis of Figure 3a. The oldest individuals were two neutered females, both reportedly fifteen years old, though the most elderly farm cat in the survey, claimed to be aged seventeen years old, was omitted from this sample owing to lack of information on either gender or neutering status. Since these are essentially not pet cats in the sense of most domestic house cats, a detailed knowledge of cat age may be imprecise, particularly in the larger colonies and for the older cats. The accuracy of declared cat ages can only be regarded as approximate and the resultant mean and survivorship calculations should be regarded with caution. The most frequent claimed age was two years, making up 21% of the sample. Assuming cats of less than one year to be 6 months old, the mean age for the entire sample was 3.3 years. The numbers of intact and neutered males and females are given separately for each year age category in Appendix 3a and provide the basis of mean age calculations showing that for intact females this was 3.3 years, compared with 3.0 for intact males. For neutered cats, mean ages were substantially greater, with 5.0 years for females and 4.7 for males. The under-representation of cats under one year old is no doubt due to the absence from the survey of dependent unweaned kittens, though other anomalous depressions in the survivorship curve (Figure 3a) (at years one, seven and eleven) could be due to intermittently low recruitment into the population during epizootics of cat flu or feline enteritis (John Jack pers. comm.). Gender and neutering rates “Continual wars and wives are what Have tattered his ears and battered his head. ” Esther’s Tomcat Ted Hughes (1960) Statistics on the 971 cats for which gender and neutering data were available have been assembled into Appendix 2 from which % frequencies have been calculated for the regional sub-sets and for the county sample as a whole. Males made up 36.6% of this sample, a lower rate than for pet cats in rural, suburban and urban areas (see Figure 7) fitting in with the popular concept of intact adult ‘tom’ cats having ‘harems’ of females. 104 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat FIGURE 3a & 3b Survivorship pattern of (a) 588 cats on Yorkshire farms (b) 245 domestic house cats. 105 animals (10.8%) had been neutered, again a markedly lower rate than for pet cats, even those in rural areas (see Figure 6). Neutering rates differed between the sexes, accounting for 15.2% of males and only 8.3% of females. Although undertaken partly as a means of controlling colony size, the necessary veterinary expenses incurred, particularly in neutering female cats, may be regarded as unjustifiable in the context of the overall farm budget. The wide regional variations in neutering rates possibly provides evidence for this interpretation, with the small mixed farms of North York Moors recording a neutering rate of 3.8% whereas the huge farms from the arable (probably wealthier) regions recording 13% (Humberhead Levels), 14.8% (Holderness) and 18.7% (Wolds). The role of the farm cat In an attempt to evaluate the practical role of cats on farms, enquiries were made as to whether cats were kept for rodent control, as pets or for any other reasons (see Appendix 2). Cats were partially regarded as pets on 39% of farms but were kept for rodent control 105 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat on 86% of them. This compared favourably with other methods of control such as poisons 65%, traps 17%, firearms 6%, dogs 5% and ferrets 0.5%. Of the perceived effectiveness of cats in farm rodent control, they were regarded as playing an effective anti-rodent role on 90% of farms: indeed, on 21% of these, farm cats were claimed to be the only form of rodent control necessary. Attitudes and experience varied somewhat, with 10% of farmers regarding cats as of little or no use in rodent control and, according to annotations on returned forms, on 7.6% of farms they were positively not welcome, farmers claiming that their presence was due to an inability to get rid of them. Interestingly, footnotes on several returned forms maintained that the presence of cats on farms generally suppressed the build-up of rodent pests or kept populations below levels where it became necessary to employ commercial rodent control contractors. It was also felt that they prevented or delayed re-colonisation after major infestations had been dealt with. These impressions were borne out by experiments monitored by Elton (1953). Discussion and general observations Farm cat populations were found to be largely self-perpetuating, although colonies eradicated by cat flu (one colony of ten cats died out during the course of the survey) and in one case by road casualties, were re-established by introductions of young cats from neighbouring farms. 7.6% of farms harboured colonies which were claimed to be unwanted and untended. The tenacity of the colonies was evidenced by some farmers not sympathetically disposed towards cats, claiming that populations had become established through strays taking up residence (5% of colonies were of purely feral origin), or that colonies had persisted despite attempts to discourage them. Colony size was claimed to be partially under the control of the farmer in that unwanted kittens, if located early enough, were frequently destroyed or given away to either neighbouring farms or as domestic pets. Although neutering was employed as a means of population management, this practice was relatively marginal, affecting only 10.8% of the sample. At mean population densities such as one cat per 78 acres (31.6 ha) on the Wolds or even one cat per 21 acres (8.5 ha) in North-east Yorkshire, it may seem unlikely that farm cats would have any significant effect on small vertebrate populations in the wider countryside: indeed, Elton (1953) showed that cats had little effect on field rat populations. However, these densities seem high compared with some wild predators, Harris et al. (1995) estimating a post breeding density for stoats ( Mustela erminea ) in unimproved grassland and arable land as being 2 per 1 km2 or a maximum of 1 fox ( Vulpes vulpes) social group ( 1 adult male and a breeding female) per 1 km2. As cats, particularly females and immature males, generally remain in the vicinity of the farm buildings (Macdonald 1981), it is highly likely that here they could exercise a more focused and effective control over synanthropic species such as house mouse (. Mus musculus ) and common rat ( Ratus norvegicus). However, Elton (1953) demonstrated that even here they had little effect on well established rat infestations, though they were found to be very efficient at preventing rats (perhaps unfamiliar with the terrain) from re-colonising after premises had been cleared of rodents. Field observations on arable (root crop) farmland on the Hatfield Levels in the southern Vale of York showed that cats seasonally congregate for hunting around field-side root crop stores (clamps) established hundreds of metres from the nearest buildings. Foxes are also attracted to these field-side stores, their diets, studied from scat analysis (Howes in prep.) consisting largely of young common rats. It is reasonable to assume that the same quarry also attracts the cats. Cats are also known to move to newly mown hay meadows and fence lines overlooking recently dredged ditch networks, to prey on rodents made vulnerable by disturbance and removal of cover. Under such circumstances farm cats in the USA have been shown to compete successfully for prey against wild predators, Pearson (1964) showed that cats alone had reduced a rodent population by 88% and George (1974) gives an example of cats competing effectively for small mammals against birds of prey. Clearly, farm cats only represent one component of the total rural cat population, others 106 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat being feral animals not part of the farm-orientated populations and pet cats kept on adjacent domestic premises. Since the days of Elton’s (1953) studies, urban development in rural communities has proceeded apace and the effect of cat predation on small mammals and competition with other mammalian predators must be considerably greater than in the 1950s. Two characteristics of domestic (pet) house cat populations are that males (for whatever reason) tend to outnumber females (see Figure 7) and that the population is subject to a relatively higher rate of neutering (see Figures 6 and 8). Using data from Appendix 2 as an index of ‘petness’, Wolds farm cats exhibit the highest level of ‘petness’ in the Yorkshire region. Here, males, which make up 47.6% of the population (mean Yorks. = 36.6%) are more frequent than in any other farming region, also with 25.3% of the Wolds farm cat population being neutered (mean Yorks. = 10.8%), this is also the highest level in the region. The role of cats on Wold farms may be providing pointers towards changes in the status of farm cats generally. Perhaps the poetic idea of farm cats being purely guardians of a nation’s cheese could be changing, cats adding the therapeutic role of ‘companion animal’ to their repertoire of benefits. A Survey of ownership levels and the population structure of Domestic House Cats in Yorkshire “ For every house is incomplete without him ...” For I will consider my cat Jeoffry Christopher Smart (1760s) Questionnaire surveys undertaken by the YNU and Doncaster Museum were used to gather information on the population densities, sex ratios, neutering rates and age structure of pet cat populations in urban, suburban and rural areas of Yorkshire. Although preliminary data were used extensively and humorously in Roger Tabor’s excellent book The Wildlife of the Domestic Cat (Tabor 1983) and referred to in Ecology of Urban Habitats (Gilbert 1989), this occasion provides an opportunity to present the final results of these unpublished surveys. Methods With the indefatigable assistance of members of the YNU affiliated societies throughout Yorkshire and groups of school children, particularly in the Doncaster region, a questionnaire survey was conducted from September to December 1980. Surveyors recorded the numbers and, where known, the ages, sexes and neutering status of cats from ten domestic properties adjacent to (but not including) their own home. In addition, housing types were recorded, as were a range of habitat types and land-uses. The general nature of the sampling area was allocated to categories described as urban/town centre, suburban and rural. Results Three hundred and seventy-one survey sheets were completed, the resultant data on housing types, adjacent habitats and land-uses and cat ownership levels being summarised in Appendix 4. Statistics on gender and rates of neutering are summarised in Appendix 5 and for those cats for which age data was provided. Appendix 3b categorises intact and neutered males and females into age classes. Housing types and habitat availability How the housing types and the habitat and land-use availability varied across the three district categories is illustrated in Figures 4a-c and Figures 5a-c respectively. Of the housing type analysis (monitoring the relative frequencies of flats, terraces, semi-detached and detached properties) the most significant feature, which increased along with cat population density, was the detached properties with a surrounding garden ‘territory’, Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 107 FIGURE 4a-c Housing types in (a) Urban (b) Suburban and (c) Rural areas. which increased from 9% of properties in town centre and urban areas to 37% in suburban districts to 49% in rural areas. Of the habitat and land-use availability analysis, the relative frequencies of larger gardens of above 150 yd2 and access to adjacent arable and pasture land seemed to be most significant, large gardens being represented in 5%, 16% and 19% of urban, sub-urban and rural districts respectively and agricultural land being represented in 0%, 17% and 47% of urban, sub-urban and rural districts respectively. Curiously, the availability of parks, sports fields and allotments seemed to be negatively related to the presence of cats; thus it could be said that from a cat’s point of view, these municipal provisions are no compensation for the absence of rural open space and protected large gardens. Population densities of domestic cat populations “Not long ago this phenomenal Cat Produced seven kittens right out of a hat!” Mr Mistoffelees T. S. Eliot (1939) 108 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat FIGURE 5a-c Adjacent Habitat/Land-use types in (a) Urban (b) Suburban and (c) Rural. From the 3,710 properties surveyed some 1,401 cats were recorded, giving a mean population level of 37.76 cats per 100 households. The levels of ownership varied substantially according to housing region, the fewest, at 25.09 cats per hundred properties, being recorded in town centre and urban areas, rising to 33.76 in suburban districts, with the highest ownership level of 53.12 in rural areas. These statistics are of considerable significance when considering aspects of predatory pressure and may in some cases have a bearing on the sustainability of populations of certain prey species and of other native wild predators (see Table 4). Neutering rates and gender ratios Figure 7, based on the amalgamation of data from the farm cat study with that from the domestic cat gender and neutering survey (see Appendix 1 and 5), shows that sex ratios vary substantially according to rurality and housing regions. Male cats on farms and in rural properties represent 36% and 44% of their respective populations, whereas males in suburban and urban/town centre regions represent more than half their populations, making up 54% and 5 1 % respectively. Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 109 Farm (971) Rural (634) Suburban (323) Urban (201) Land-use category and population sample size FIGURE 6 Neutering rates of cat populations on farms and in rural, suburban and urban regions in Yorkshire. 70 65 60 FIGURE 7 Males as % of populations on farms and in rural, suburban and urban regions in Yorkshire. Two factors which may be at work here are (a) disproportionate mortality rates for males and (b) gender preference in domestic cat ownership. Taking mortality first, causes of mortality which may affect males to the extent that sex ratios are tipped in favour of females could be (a) intact males wandering in search of mates and therefore being subject to mortality through road traffic accidents or being killed or injured by dogs or in ‘vermin’ traps and (b) injury through fights with other intact males for access to mateable females. If wandering and fighting are characteristic behaviours of intact males and if these 110 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 80 70 60 1 50 jfi 40 1 | 30 20 10 0 20 30 40 50 60 Males as % of population FIGURE 8 Relationship between male neutering and males as % of population. behaviours are suppressed or absent in males which are neutered (pre-maturity), then the proportion of males in a population could be a function of the rate of male neutering within that population. Interestingly, Figure 8 shows that the % frequency of males in a population does indeed rise as male neutering rates increase. According to anecdotal input to this study, evidence for gender preference being a major influence in domestic cat population composition seems to be contradictory. On one hand there are owners who claim to prefer male cats on the basis that a males will not themselves become pregnant and are cheaper to have neutered; on the other hand there are those who, despite the higher cost of neutering, prefer females on the basis that they don’t ‘spray’ or ‘caterwaul’ and are less likely to wander and be killed. If cost of neutering was a significant factor affecting choice, then one would expect the less affluent urban and town centre areas to have a higher incidence of males and for the more affluent suburban sample to have a higher incidence of females. Figure 6 shows this is not the case. Statistics on gender ratios and neutering rates are available from some cat ecology and behavioural studies (Dards 1981, Macdonald 1981, Rees 1981, Tabor 1983, Churcher & Lawton 1987, Soulsby & Serpell 1988). However, since neutering levels are regarded as being central to the management of a range of cat-related animal welfare and public health issues (Hammond 1981, Jackson, 1981, Remfrey 1981) it is surprising how few surveys have been undertaken. The Yorkshire study appears to be unique in revealing patterns of variation across different land-use and housing type categories. Figure 6 shows that major differences occur in cat neutering levels according to land-use and housing areas, with below 15% on farms, below 45% in rural areas but over 70% in both suburban and urban areas. Although the farm and rural neutering levels are particularly low, even the Yorkshire suburban and urban figures are relatively low compared with the 82% of males and 90% of females recorded for the London suburb of Romford (Tabor 1983). Differences in neutering rates between the sexes however, show relatively little variation in the pet cat samples. The situation with the farm cat ratio is discussed above. Age structure The ages of 245 cats (17.5% of the total house cat sample for which gender and neutering information was available) are shown in Appendix 3b and form the basis of Figure 3b. The Suburban house cats 75.5% 0 Urban house cats 70.8% 0 Mean Yks. house cats 58.8% 0 Rural house cats 43.7% 0 Wolds farm cats 25.3% 0 Mean Yks. farm cats 15.2% 0 0 North York Moors farm cats 4.5% Ill Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat oldest individuals were a neutered male and female of 17 years and a neutered female of 19 years of age respectively. Since these animals were generally not the pets of the surveyors, the accuracy of declared cat ages can only be regarded as approximate and the resultant mean and survivorship calculations should be regarded with caution. The most frequent claimed age was three years, making up 20% of the sample. Assuming cats of less than one year to be 6 months old, the mean age for the entire sample was 4.3 years. The numbers of intact and neutered males and females are given separately for each year age category in Appendix 3b and provide the basis of mean age calculations, showing that for intact females this was 3.9 years, compared with 2.9 for intact males. For neutered cats mean ages were substantially greater, with 4.7 years for females and 4.8 for males. As with the farm cat study, the under- representation of cats of less than one year of age is no doubt due to the absence from the survey of dependent unweaned kittens, though other apparent anomalies in the survivorship curve (at years one, two, eight and ten) could, as was observed in the farm cat study, be due to intermittently low recruitment into the population during epizootics of cat flu or feline enteritis (John Jack pers. comm.). In that this census was undertaken in 1980, a year later than the farm cat survey, it is interesting to note that anomalies in the survivorship pattern in Figure 3a are shifted on a year in Figure 3b, indicating that the relatively unmanaged farm cat populations and the cosseted house cat populations are subject to the same epidemiological cycles and survivorship trends. What the Cat Brought In: a study of the predatory ecology of domestic cats in Yorkshire “. . . and tell me all thy frays Offish and mice and rats and tender chick. ” To Mrs Reynold’s Cat John Keats (1818) Introduction Amid the artificial garden and backyard eco-systems, there lurks the scourge of fish, fowl and flesh - the domestic cat. This obligate carnivore, which has little dietary need to hunt, still harbours a potent behavioural drive to ambush, attack and kill (Turner & Meister 1986). Anecdotal accounts of cat prey items (be they birds, mammals, herptiles or fish) gathered by various natural history societies have periodically thrown light on the taxonomic range, relative frequency and seasonality of prey taken by domestic cats. Mead’s (1982) analysis of cat-killed ringed birds submitted to the British Trust for Ornithology (BTO) bird ringing scheme perhaps provided the earliest critical insight into the hunting strategies of cats and indicated which bird species are most likely to fall victim. Investigations into the predatory activities of domestic cats in Britain received an unexpected boost during the compilation of An Atlas of Yorkshire Mammals (Howes 1983) when in 1978-79, after mammal trapping and bird pellet analysis had been exhausted as sources of data, a questionnaire survey entitled What the Cat Brought In was designed as a ploy to garner records of small mammals from unrecorded areas. This exercise, which began to reveal fascinating insights into the predatory behaviour and impact of domestic cats, was continued in order to provide further information on this little studied aspect of cat ecology. Initially launched through the YNU and Doncaster Museum it involved cat- owning members of YNU affiliated societies and ultimately the general public in monitoring for a year the prey items caught by their cats. Preliminary reviews of results were published in the Yorkshire Naturalists’ Union Newsletter (Howes 1979a, 1981a) which served to stimulate further interest. Following the success of the Yorkshire prototype, a series of promotional and feed-back articles in the Mammal Society’s Youth News (Howes 1979b, 1980a, 1981b) and the RSPB’s Young Ornithologists’ Club’s Bird Life (Howes 1980b, 1981c), assisted by 112 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat extensive press coverage, succeeded in developing the What the Cat Brought In project into a major national survey and placed on record some of its preliminary results. Together with substantial assistance from the Mammal Society, the RSPB, regional support groups of the Cat’s Protection League, RSPCA and a range of County Wildlife Trusts, the project succeeded in monitoring the annual predatory activities of c. 1000 domestic cats throughout Britain. The survey’s methodology provided a basis for the now celebrated work of Paul Churcher on the predatory impact of cats in a Bedfordshire village (Churcher & Lawton 1987). Batches of unprocessed prey records have been used in various undergraduate research projects and also by Professor Tim Birkhead and his student Ian Massie of Sheffield University for comparison with the predatory activities of magpies {Pica pica) (Birkhead 1991). Though light-heartedly summarised in Howes (1990, 1992), the full range of data has never been fully analysed; however, the raw data archive has been passed to Professor Stephen Harris and his team at Bristol University for comparison and merging with the Mammal Society’s recent (1997) re-run of the project (see http://www.abdn.ac.uk/mammal.catkillsl.htm). There is currently a vigorous worldwide interest in the food ecology and predatory impact of domestic cats, the now extensive literature (though not including the Yorkshire component) being reviewed in Fitzgerald (1986), Turner and Meister (1986), Bradshaw (1992), and Pearre and Maass (1998). This occasion provides an opportunity to present the results of the survey of the predatory activities of domestic cats in Yorkshire. Methods The What the Cat Brought In questionnaire survey was designed to monitor the prey items caught by individual domestic cats living in rural, suburban and town centre areas for a year. To increase the likelihood of prey items being correctly identified, the survey was targeted primarily at the cat owning members of local natural history and ornithological societies. In addition to recording the name, age, sex and neutering status of each cat, the questionnaire forms recorded the address of the surveyed cats, the prevailing housing types and nearby habitat and land-use types. The date of receipt and return of the form was noted, followed by the date and identification of each successive prey item. As an index of palatability, those items which were eaten or partially eaten were indicated. Results The predatory activities of 180 cats (84 from rural, 70 from sub-urban and 26 from urban and town centre areas of Yorkshire) were each monitored for a year. Monthly and annual totals of prey types are categorised separately for taxa of mammals (16), fish (3), herptiles (2), birds (35) and invertebrates (7) in Appendix 6. The numbers of prey types recorded varied substantially according to hunting areas, Figure 9 showing that biodiversity of prey taxa generally increased from the lowest level in urban/town centre areas to the highest in rural areas. Of the vertebrate prey items recorded in the overall sample, mammals made up 69.1%, birds 30.5% and other vertebrates (herptiles and fish) constituted 0.4, however, Table 1 reveals substantial regional variations. Capture rates Some 5,205 vertebrate prey items were recorded, giving a mean annual prey capture rate of 28.91 per cat. However, a wide variation in rates of predation was evident across the housing regions, Table 4 showing a mean annual rate per cat rising from 7.88 in town centre and urban areas to 37.51 in rural districts. Table 1 shows the relative compositions of birds, mammals and other vertebrates across the three hunting areas, revealing the general principal that birds constitute the highest proportion (72.2%) of vertebrate prey in urban areas, declining to the lowest level (22.3%) in rural areas, whereas mammals constitute the Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 113 Cat Hunting Zone FIGURE 9 Diversity of predated species in different hunting zones. highest proportion (77.6%) of vertebrate prey in rural areas, declining to the lowest level (27.8%) in urban areas. Interestingly, this demonstrates a similar predatory trend as observed in the analysis of tawny owls ( Stryx aluco) prey in Greater London where high levels of bird predation were linked with the inner London Boroughs and the higher levels of mammal predation were encountered in the outer more rural districts (Bevan 1965). TABLE 1 Recorded annual predation rates on birds, mammals and other vertebrates in different housing areas of Yorkshire Housing/ Cat Birds Mammals Other Total Land-use Sample size vertebrates vertebrates Type No. No. % Mean No. % Mean No. % Mean No. Town centre/ Urban 26 148 72.2 5.69 57 27.8 2.19 0 0.00 0.00 205 Suburban 70 735 39.8 10.50 1,095 59.3 15.62 17 0.92 0.24 1,847 Rural 84 703 22.3 8.37 2,448 77.6 29.14 2 0.06 0.02 3,153 TOTAL 180 1,586 30.5 8.81 3,600 69.1 19.98 19 0.36 0.10 5,205 * Mean = mean per cat per year. By linking the mean prey capture figures per cat in Table 1 with cat population density levels in Appendix 4 and Figure 4a-c, it becomes possible in Table 2 to estimate separately the predation levels of prey types per 100 households respectively in town centre, suburban and rural housing/landuse areas. Again, this emphasises an increase in predation levels the more rural the hunting habitat becomes. Interestingly, although relatively token, the rate of predation on herptiles and fish in the suburban sample is probably a reflection of the frequency of ornamental garden ponds. 114 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat TABLE 2. Estimated mean annual cat predation levels on birds, mammals and other vertebrates per 100 households in different housing areas of Yorkshire Housing/ Land-use type Birds Mammals Fish & Herptiles Town centre/Urban 142.8 54.9 0.00 Suburban 354.5 527.3 8.10 Rural 444.6 1,546.3 1.06 Seasonality Monthly totals of reported prey for all housing regions, provided in Appendix 6, have been used to generate Figure 10 which shows relative % composition of mammals and birds vertebrate prey for the entire sample throughout the year. This shows that mammals outnumber birds in each month, exceeding 60% of vertebrate prey in the ten months from July to April, and exceeding 80% in October and November. Predation on birds reaches a peak in the months of May and June when rates exceed 40% of vertebrate prey, followed by an isolated peak of above 35% in January. JAN Monthly variations in the percentages of 5,205 mammal and bird prey items caught by 180 cats. By separating the seasonal vertebrate prey data according to housing regions, Figure 1 1 a-c demonstrates the increasing significance of birds as prey items in the suburban and urban samples, emphasising their exploitation during breeding season, exceeding 40% of vertebrate prey from May to August in the suburban sample and exceeding 80% from May to September in the urban sample. The suburban and urban samples also emphasise the winter predation of birds that peaks in December and January. It is likely that this phenomenon is a consequence of birds being artificially attracted during periods of hard weather into urban cat territories to feed on well-stocked bird tables. Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 115 FIGURE 1 1 Monthly variations in vertebrate prey types taken by (a) 26 cats in Urban/Town Centre, (b) 70 cats in suburban and (c) 84 cats in Rural areas of Yorkshire. Mammal prey case studies Considerable variation in the composition of the mammalian prey faunas in the urban, suburban and rural hunting zones is illustrated in Figures 9 and 12, both of which show that taxonomic diversity increases from urban to rural hunting zones. Figure 12 shows that murids (mice and rats), which accounted for 97% of mammals in the urban sample and only 58% in the rural sample, were the most frequently taken mammal prey group across the regions. Although the incidence of voles, shrews and other mammals increased with rurality, it was interesting to note token evidence of both shrew and vole occurrence in urban areas. Figure 13, based on records in Appendix 6, shows specifically identified mammal species placed in order of frequency, with the wood mouse ( Apodemus sylvaticus ) accounting for 56% of the sample. The following review deals with mammals in family order. Only eight bats were represented in the survey, though their presence from June to October and the fact that 116 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 100 Urban Suburban ■ MURIDS ffi VOLES IB SHREWS S3 OTHERS Rural FIGURE 12 Variations in MAMMAL prey taken in different habitats. FIGURE 13 % frequency of identified prey types: Mammals. recorders described them in terms of ‘half inch’ and later ‘three quarter inch’ bats suggested that these were likely to be newly fledged young pipistrelle bats ( Pipistrellus pipistrellus s.l.). In addition, anecdotal information and WTCBI recording sheets submitted by correspondents elsewhere in Britain (Howes 1981a, 1981b) showed a pronounced peak of captures in May which coincided with the formation of summer nursery colonies. Since colonies are largely associated with domestic property, bats at this time are vulnerable to 117 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat cats attracted to windowsills, dormer roofs etc. by the activities and ultrasound emissions of the aggregating bats. None of the bats were reportedly eaten. The usually subterranean mole ( Talpa europaea ) is generally safe from cat predation; indeed, the 25 moles recorded in this survey represent only 0.7% of mammal prey items. The occurrence of moles above ground tends to vary according to prevailing soil moisture conditions, typically in times of flood or drought. Interestingly, the peak of 15 captures in June and July of 1979 coincided with one such drought period. None of the moles were reported to have been eaten. All three species of shrew were reported and although occurrences were monitored throughout the year, there was a pronounced peak in July and August (see Appendix 6). With 409 reported captures, shrews represented the third most frequent mammal prey group, though significantly none were reported as having been eaten. This may be associated to unpalatability, Macdonald (1977) demonstrating by feeding experiments that foxes ( Vulpes vulpes ) would only resort to eating shrews when particularly hungry. All three species of vole were reported with the 630 captures, most of which were eaten, representing the second most frequent mammal prey group. Of those specifically identified, 78% were field voles ( Microtus agrestris ), 21% were bank voles ( Clethrionymus glariolus ), and only 1% (2 specimens) were water voles (Arvicola terrestris). Although significant numbers are taken through the year, numbers rose markedly in July, remained high through the late summer and early autumn and peaked in October (see Appendix 6). The sudden rise in vole predation in July may have been a response to field voles made vulnerable after the removal of grass crops. The autumnal peak in predation is probably due to the aggregations of new generations of voles with breeding continuing through the summer and early autumn. “Such appetite he hath to eat a mous ” The Maunciple’s Tale Geoffrey Chaucer (14th century) Three species of mice were reported, with the 2,160 captures, most of which were eaten, representing the most frequent mammal prey group. Of those specifically identified, 93% were wood mice ( Apodemus sylvaticus), 6.5% were house mice ( Mus musculus ), and only 0.1% (1 specimen) was harvest mouse ( Micromys minutus). Seasonality of occurrence as prey was similar to that in the voles, with fewest captures in February, rising to a late summer peak in September, a month earlier than for the voles (see Appendix 6). “Cats I scorn, who sleek and fat, Shiver at a Norway Rat; ” A True Cat Anna Seward (1810) Although much discussed by correspondents, common rat (. Rattus norvegicus ) was recorded on only 103 occasions and represented 7% of identified mammal prey items. Since specimens were frequently described as ‘young’ or ‘small’, this would indicate that adult rats tend to be avoided. Like the other rodent groups there was a marked September and October peak, probably representing aggregations of new generations, with breeding continuing through the summer and early autumn. “I know a Cat, who makes a habit of eating nothing else but rabbit, ” The Ad-dressing of Cats T. S. Eliot (1939) Rabbits ( Oryctolagus cuniculus), often described as ‘young’, were recorded on 159 occasions and represented 10% of identified mammal prey, all kills being reported in rural areas. Since they were taken throughout the year, some of those caught would certainly have been fully-grown. A significant peak of kills recorded in August may have been in response to the appearance on the surface of newly independent young of the year or animals made vulnerable through the harvesting of cereal crops. 118 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat Grey squirrels ( Sciurus carolinensis) were caught on three occasions within the survey but cat owners in suburban areas adjacent to community woodlands anecdotally reported frequent kills. In addition, one suburban and four rural weasels ( Mustela nivalis ) were reported in the survey. Other mammals, which though caught in Yorkshire were caught outside the survey period, included brown hare ( Lepus europaeus ) and stoat ( Mustela erminea ). Notable historical cat kill records include one of the last red squirrel to be recorded in the Barnsley area, ambushed by two Siamese cats in March 1967 in the grounds of the Northern College, Stainborough (I. Mackerness pers. comm.), a black rat ( Rattus rattus ) from the premises of R. Wade & Co. Hull 10.2.1901 (Specimen in Hull Museum) and a dormouse ( Muscardinus avellanarius ) killed on the banks of the Hodder in Bowland during the 1860s (Anon. 1884). Bird prey case studies Birds featured significantly as prey items but despite extensive negative comments by correspondents on what was regarded as this unacceptable behaviour, the 1,586 kills represented only 30.5% of reported vertebrate prey items. Considerable variation in the composition of the avian prey faunas in the urban, suburban and rural hunting zones is illustrated in Figures 9 and 14, both of which show that taxonomic diversity increases with rurality. Figure 14 shows that house sparrow ( Passer domesticus ), which accounted for over 70% of birds in the urban sample and less than 30% in the rural sample, was the most frequently taken bird species across the regions. Curiously, like the house sparrow, the blackbird ( Turdus merula) also exhibited an urban orientation, declining slightly in the suburban and rural samples. By contrast, the starling (Sturnus vulgaris ) and song thrush ( Turdus philomelos) exhibited a suburban orientation, declining in both urban and rural samples. 100 90 80 I 70 i 60 i 50 i 40 ^ 30 20 10 0 Urban Suburban Rural Cat Hunting Zones FIGURE 14 Variations in BIRD prey taken in different habitats. Figure 15, based on records in Appendix 6, shows the ten most frequently identified bird species placed in order of frequency, with house sparrows accounting for 62% of the total sample, the next most frequently predated species being starling, blackbird and song thrush, representing 9, 7 and 5% respectively. ■ HOUSE SPARROW ffl BLACKBIRD ID SONG THRUSH 0 STARLING □ OTHERS Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 119 FIGURE 15 % frequency of identified prey types: Birds. Although 35 taxa of identifiable birds are indicated in Appendix 6, the ten most frequently taken species (see Figure 15) are generally similar to the cat-killed species of ringed birds identified by Mead (1982). All of the frequently taken prey species either regularly feed or breed in domestic gardens and are therefore likely to live in close proximity to cats. However, the most likely criterion of vulnerability is that house sparrow (. Passer domesticus ), starling ( Sturnus vulgaris), blackbird ( Turdus merula ), song thrush (Turdus philomelos ), robin ( Erithacus rubecula ), dunnock ( Prunella modularis) and wren (' Troglodytes troglodytes ) are essentially ground feeders, often feeding close to shrubs and vegetation which cats can use for purposes of ambush. Although blue tits ( Parus caeruleus) are normally canopy feeders and are therefore out of reach of cats, they are regular visitors to bird tables and garden feeding stations and frequently breed in nest boxes erected in sites vulnerable to house cats. Greenfinches (Carduelis chloris ) are also generally arboreal feeders, though Mead (1982) points to their habit of feeding terrestrially on food debris dislodged from bird tables or nut feeders, making them particularly vulnerable to cat attack. Seasonality of most prey species was closely associated with breeding period when adults were nest building and particularly when young were fledging. This is particularly noticeable in the blackbird with most casualties in May and June, song thrush with most casualties in May, June and July, robin with a peak in June and July and dunnock with a peak in July. The prolonged breeding period of the house sparrow is reflected in high levels of predation from May to August. With this slow flying species nesting almost exclusively under the same roofs, so to speak, as their house cat predators and in feeding terrestrially in gardens, frequently on household scraps, it is understandable why this species constituted over 60% of all identified bird prey. Of the migrant species, redwing ( Turdus iliacus ) (1 in January) was the only winter visitor, summer migrants being chiffchaff (1 in April), willow warbler (2 in July and 1 in August), house martin ( Delichon urbica ) (2 in June and 4 in July), and swallow ( Hirundo 120 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat rustica) (1 in July and 1 in September). Although the two hirundine species frequently inhabit the same premises as house cats, their frequency as prey species is minute, presumably reflecting their aerial feeding behaviour and aerobatic skills. However, anecdotal information from correspondents and evidence in Mead (1982) suggests that by waiting on windowsills or roofs, individual cats contrive to catch birds as they leave or return to their nests. Although the vast majority of prey species were small to medium sized passerines, larger species like black-headed gull ( Larus ridibundus) (1), moorhen ( Gallinula chloropus ) (3), collared dove ( Streptopelia decaocto ) (7), street pigeon ( Columba livia ) (3), wood pigeon (' Columba palumbus) (16) and even pheasant ( Phasianus colchicus) (1) were also taken, though most of those reported were rescued and liberated by the cat owner. Fish and herptile prey case studies Correspondents frequently referred to incidents where cats had regularly removed fish from garden ponds; however, actual evidence in the survey was sparse, producing records of six specimens of three species (see Appendix 6). Similarly, stories of cats catching large numbers of frogs during spawning season, were not corroborated, the survey producing records of only eleven frogs and two newts. Small though this sample was, it identified an early springtime (March) seasonality in the suburban sample (see Figure lib). Although not included in this survey, additional anecdotal records include a grass snake ( Natrix natrix ) caught in Harthill in July 1975 (W. Ely pers. comm.) and a viviparous lizard (Lacerta vivipara) caught at the Crown and Anchor, Kilnsea in July 1976 (J. Harrup pers comm.). Invertebrate prey According to correspondence and annotations on the recording forms, invertebrates were substantially under-recorded for reasons of difficulty with identification and, more particularly, in monitoring this activity from which there would be few if any remains and which did not seem to engender the behaviour of ‘presenting’ the prey. However, of the 116 items recorded, Appendix 6 shows a peak of activity from June to October. Anecdotally, recorders mentioned the indoor autumnal activity of their cats catching and eating the large Tegenaria spiders, the mature males of which nocturnally wander around the ground floors of domestic property in search of occupied female webs. Of the butterflies, large white (. Pieris brassicae), green-veined white ( Artogeia napi), red admiral ( Vanessa atalanta) small tortoiseshell (. Aglais urticae) and peacock ( Inachis io) were listed, the latter two featuring in early spring when warm days in March and April brought specimens which had hibernated indoors fluttering to sunlit windows where they were easily caught by sunbathing cats. The same strategy also led to the demise of larger muscid flies and bumble bees ( Bombus spp.) in summer and, in late summer, the ‘Drone flies’ ( Eristalis spp.). In addition there were anecdotes of cats catching large Aeschna dragonflies and bringing home the larvae of the elephant hawk moth ( Deilephila elpenor). Predatory rate according to age “He was a mighty hunter in his youth ” The White Cat ofTrenarren A. L. Rowse (1967) In counting the numbers of vertebrate prey items recorded for cats in each age group it has been possible to assemble Figure 16, which reveals that predatory activity rapidly builds to a peak of 43.5 vertebrate prey items per year at two years of age, numbers of prey items caught declining thereafter. Broadly, cats up to four years old catch a mean of 3 1 vertebrate prey items per year. This falls to a mean of 20 for cats between five and nine years of age and only 15 for cats of ten years and above. That cats up to four years of age catch prey in excess of the sample mean of 28.91 may be related to the natural life expectancy of cats as Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 121 ! 's I 1 1 FIGURE 16 Changes in levels of predatory actvity with age: illustrated by mean annual number of vertebrate prey items caught per cat (5,205 prey items caught by 180 cats). a species, the survivorship graphs of both farm and house cats (Figures 3a & 3b) showing substantial diminution in the proportion of the population of cats over four years of age. Discussion Being generally well fed, the domestic cat is probably fitter to go hunting than its untended feral counterpart. In receiving a food source independent of the productivity of their ‘wild’ environment, domestic cat populations, unlike those of wild predators, are not controlled by prey availability. They are therefore potentially in a position to a) severely deplete or eradicate wild prey populations and b) unfairly compete with native terrestrial carnivores. These aspects together with the sheer number of cats in Britain, could be having detrimental effects on vertebrate faunas around our towns, suburbs and rural villages; indeed, where localised or rare prey species are involved, this could be of conservation significance. Soulsby and Serpell (1988) showed that the domestic cat population in Britain increased from 4 million in 1963 to 6.2 million in 1988, and the JNCC review of British mammal populations (Harris et al. 1995) estimated that this would rise to around 8 million by the year 2000. In addition, Harris et al. (1995) considered that to this could be added an estimated 813.000 feral cats. Table 3 reveals that these numbers are extremely high compared with estimated totals of all other British terrestrial mammalian predators, domestic cats outnumbering the entire estimated population of wild terrestrial carnivores by 2.6 to 1. The survey found that on average each cat brought in some 29 prey items a year which by simple extrapolation would suggest that Britain’s eight million domestic cats could be killing some 232 million vertebrates per year. Since the largest sample of cats (46.6%) in the survey was from the predatorily successful rural sample (see Table 1), this blanket figure gives a misleadingly high impression. Having organised the Yorkshire cat prey and cat population surveys on the basis of distinct housing, habitat and land-use regions, this very crude global figure has been refined in Table 4 to predict a potential annual predatory pressure by domestic cats per 100 households, giving figures in the order of 1,992, 890 and 197 in rural, suburban and town centre areas respectively. 122 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat TABLE 3 Population estimates of terrestrial and riparian carnivores in Britain compared with domestic and feral cat Felis catus (data from Harris et al. 1995) Hedgehog 1,555,000 Stoat 462,000 Weasel 450,000 Badger 250,000 Fox 240,000 American Mink 110,000 Polecat 5,000 Otter 7,350 Pine Marten 3,650 Wild Cat 3,500 Feral Ferret 2,500 Total 3,099,000 Domestic Cat 8,000,000 Feral Cat 813,000 Total 8,313,000 TABLE 4. Estimated mean annual vertebrate predation rates by cats in different housing areas of Yorkshire Housing type +Mean predation *Cat density Estimated mean predation rate per cat per 100 rate of cats per 100 households households Town centre/Urban 7.88 25.09 197.7 Suburban 26.37 33.76 890.3 Rural 37.51 53.12 1,992.5 +based on Table 1; *based on data in Appendix 4 and Figure 6a-c. Cat Predation Management In providing a statistical rather than a purely anecdotal review, this study usefully reveals that the predatory activities of cats vary substantially with: a) Age - predatory activity declining substantially after year 4 (see Figure 16). b) Housing/land-use/habitat districts - annual predatory success (or perhaps opportunity) declining from 37.5 vertebrate prey item per cat in rural areas to 7.9 in town centre/urban areas (see Table 1). c) Season - the majority of garden birds are taken in winter (December and January), presumably when attracted to bird tables, and during the nesting season (notably in May and June) when fledglings are vulnerable (see Figures 10 & lla-c). The restriction of nocturnal and crepuscular activities of the more numerous and more predatory younger cats would therefore achieve significant reductions in small mammal and bird predation and in turn would reduce competitive pressures on wild carnivore populations, particularly if this prohibition were exercised on rural and suburban cats. If excessive predation on garden birds is perceived to be a problem, then the restriction of diurnal access to gardens during peak winter feeding and summer breeding periods would certainly reduce this effect. Whether the fitting of cats with a sonic (e.g. collar and bell) or optical (e.g. reflective collar) warning device could significantly reduce predatory success, 123 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat would require experimental verification. Similarly, an examination of the camouflage/ predatory efficiency of cats may show that certain coat colour/pattern morphs may be less efficient as predators and therefore more desirable as ‘animal welfare-friendly’ pets. In our reportedly stress dominated and progressively dysfunctional lives, where cats evidently play an important role as therapeutic companion animals (Soulsby & Serpell 1988), the simple reduction of pet cat populations as a means of controlling predatory pressure would not seem to be a straightforward option. Intriguingly, a progressive substitution of domesticated carnivores by domesticated herbivores as companion animals may overcome a range of perceived animal welfare and predator-prey difficulties, but would set in train a range of quite different dynamic trends in our urban ecology. Yorkshire cats and their owners have contributed substantially to our knowledge of the population structure and ecology of the domestic cat in Britain. The studies have given cat owners added interest in their free-range charges but with the various discoveries, revelations and conservation implications, the project has indicated the need for more fieldwork on the ‘wild life’ of the domestic cat. Acknowledgements I would like to thank the club secretaries and members of the Yorkshire and Humberside Federations of Young Farmers’ Clubs for their support in undertaking the farm cat survey. Thanks are also due to the numerous school groups and natural history societies throughout Yorkshire for assistance with the cat population surveys. For the year-long monitoring of the predatory activities of their pet cats I would like to thank P. Abbott, L. Ablitt, V. Acaster, A. Adams, J. P. Allen, K. A. Appleby, R. S. Atkinson, C. Backhouse, S. Baines, C. Ball, L. Barber, N. Barrowcliffe, K. M. Bates, C, Blackstone, R. Blair, G. M. Boothroyd, M. Boothroyd, B. Bradshaw, J. M. Bradshaw, C. Bryant, P. Bullock, A. Cappleman, H. Chapman, K. Constable, M. Crookes, H. Cunningham, C. Davies, M, Davies, R. Dinsdale, S. Dobson, W. J. Emmott, A. M. Evans, D. Faulkner, R. Foster, M. Fraser, S. Gamson, I. Green, J. Grimes, T. Guest, M. E. Hampshire, D. Hancock, J. Hancox, I. Hare, A. Henderson, A. Hill, K. Hill, S. Hirst, M. C. Hockenhull, M. Holgate, R. Holloway, J. M. Hudson, A. Hughes, J. Hunt, J. Huntington, A. J. Ingham, M. Jackson, P. Jackson, M. Jenkins, P. A. Jobson, M. Johnson, P. Jones, G. Leeming, J. M. Ling, S. P. Lister, L. Liversidge, E. Lucas, S. Lunn, M. Macdonald, S. C. Madge, A. Marrison, M. Marrison, C. Marsden, S. Marsden, Mrs. Maw, J. McConnell, A. M. Mead, K. Miles, C. Miller, E. Morley, P. Morley, M. Moss, M. N. Newton, E. S. Noble, C. Oxford, E. A. Parkinson, D. E. Pearce, P. A. Pennington-George, J. D. Pottas, H. M. Roberts, D. Robinson, S. C. Rylatt, E. J. Scott, G. Sedgwick, B. Shail, M. Sharrock, R. Sharrock, K. Smith, L. Smith, M. Smith, S. P. Smith, B. Summerscales, J. E. Sowden, R. Spencer, R. Staley, L. Stather, M. A. Stevenson, D. Stewart, C. A. Street, C. Thoday, A. Thompson, G. M. Tutt, G. U. Tyerman, J. Walsh, R. A. Weed, B. Wetherby, C. R. Wheway, J. Whitfield, C. Wilkinson, G. Woodcock, T. Yardley, C. L. Yates and K. Yates. Throughout this and other mammal studies in the Yorkshire region I have received enthusiastic support from members of the Yorkshire Naturalists’ Union and its network of affiliated societies. The breadth of expertise encountered through the activities and publications of the YNU has added immeasurably to my understanding and appreciation of Yorkshire’s wildlife, and the friendship, foibles, hospitality and humour of Yorkshire naturalists is treasured indeed. I felt greatly honoured to be elected YNU President in the year 2000. References Bevan, G. (1965) The food of tawny owls in London. London Bird Report 29: 56-72. Birkhead, T. R. (1991) The Magpies. T. & A. D. Poyser, London. Bradshaw, J. W. S. (1992) The Behaviour of the Domestic Cat. C.A.B. International, Oxford. 124 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat Chaucer, G. (14th c.) The Manciple’s Tale. In Skeat, W. W. (ed.) (1947) The Complete Works of Geoffrey Chaucer. Oxford University Press, London. Churcher, P. B. and Lawton, J. H. (1987) Predation by domestic cats in an English village. Journal of Zoology, London, 212: 439-455. Cox, F. E. G. (1987) Protozoan parasites of British small rodents. Mammal Rev. 17 (2/3): 1-66. Dards, J. (1981) Habitat utilisation by feral cats in Portsmouth Dockyard. In The Ecology and Control of Feral Cats, pp. 68-72. UFAW, Potters Bar, London. Eliot, T. S. (1939) Old Possum’s Book of Practical Cats. Faber and Faber, London. Elton C. S. (1953) The use of cats in farm rat control. British Journal of Animal Behaviour 1: 151-155. Ely, W. (1975) pers. comm. Fitzgerald, B. M. (1986) Diet of domestic cats and their impact on prey populations. In Turner, D.C. and Bateson, P. (eds.) The Domestic Cat: the Biology of its Behaviour, pp. 123-144. Cambridge University Press, Cambridge. George, W. G. (1974) Domestic cats as predators and factors in winter shortages of raptor prey. Wilson Bull. 86: 384-396. Gilbert, O. L. (1989) The Ecology of Urban Habitats. Chapman & Hall, London. Gray, J. (1733) Fable sXXI: The Rat-catcher and Cats. In Ward, J. (ed.) (1990) The Oxford Anthology of English Poetry. Oxford University Press, Oxford. Hammond, C. (1981) Long term management of feral cat colonies. In The Ecology and Control of Feral Cats, pp. 89-91. UFAW, Potters Bar, London. Harris, S., Morris, R, Wray, S. and Yalden, D. (1995) A Review of British Mammals. JNCC, Peterborough. Howes, C. A. (1979a) The ecology of the Yorkshire Mog. Yorkshire Naturalists’ Union Newsletter 36: 2-3. Howes, C. A. (1979b) What the cat brought in. Mammal Society Youth News 7: 1-2. Howes, C. A. (1980a) The great British cat survey. Mammal Society Youth News 12: 2. Howes, C. A. (1980b) What the cat brought in. Bird Life Jan./Feb. [1980]: 2. Howes, C. A. (1981a) The ecology of the Yorkshire Mog. Yorkshire Naturalists’ Union Newsletter 40: 3. Howes, C. A. (1981b) Acro-bat-ic cats. Mammal Society Youth News 15: 2. Howes, C. A. (1981c) What the cats are bringing in. Bird Life March/ April [1981]: 18-19. Howes, C. A. (1983) An atlas of Yorkshire mammals. Naturalist 108: 41-82. Howes, C. A (1985) Feral cat Felis catus. in Delany, M. J. (ed.) Yorkshire Mammals, pp. 175-176 University of Bradford, Bradford. Howes, C. A. (1990) Red in tooth and claw. Natural World 28: 50. Howes, C. A. (1992) The backyard jungle. Yorkshire Naturalists’ Union Bulletin 18: 13-16. Howes, C. A. (in prep) The seasonal diet of foxes Vulpes vulpes on Thorne Moors. Hughes, T. (1960) Lupercal. Faber & Faber, London. Jack, J. (1980) pers. comm. Jackson, O. (1981) Problems identified by the feral cat working party. In The Ecology and Control of Feral Cats, pp. 81-82. UFAW, Potters Bar, London. Keats, J. (1818) To Mrs. Reynold’s Cat. In Gitting, R. (ed.) (1966) Selected poems and letters of John Keats. Heinemann, London. Lane, J. (1976) pers. comm. Lewis, J. W. (1987) Helminth parasites of the British rodents and insectivores. Mammal Rev. 17(2/3): 81-93. Macdonald, D. (1977) On food preference in the red fox. Mammal Rev. 7(1): 7-23. Macdonald, D. (1981) The behaviour and ecology of farm cats. In The Ecology and Control of Feral Cats, pp. 23-29. UFAW, Potters Bar, London. Mackemess, I. (1979) pers. comm. Mead, C. J. (1982) Ringed birds killed by cats. Mammal Review 12 (4): 183-186. Munro, J. W. (1960) Pests of Stored Food Products. Hutchinson, London. Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat 125 Pearre, S. and Maass, R. (1998) Trends in prey size-based trophic niches of feral and house cats Felis catus L. Mammal Review 28 (3): 125-139. Pearson, O. P. (1964) Carnivore-mouse predation: an example of its intensity and bioenergetics. Journal of Mammalogy 45: 177. Rees, P. (1981) The ecological distribution of feral cats and the effects of neutering a hospital colony. In The Ecology and Control of Feral Cats, pp. 12-22. UFAW, Potters Bar, London. Remfrey, J. (1981) Strategies for control. In The Ecology and Control of Feral Cats, pp. 73-79 UFAW, Potters Bar, London. Rouse, A. L. (1967) The White Cat ofTrenarren. Faber and Faber. London. Seward, A. (1810) A true cat. In Scott, W. (ed.) Collected Poetical works of Anna Seward. Ballantyne and Co. Edinburgh. Smart, C. (1760s) For I will consider my cat Jeoffry. In Stead, W. F. (ed.) (1939) Rejoice in the Lamb. Jonathon Cape, London. Soulsby, L. and Serpell, J. (1988) Companion Animals in Society. Council for Science and Society, Oxford University Press, Oxford. Southam, N. (1981) Health hazards in hospital areas. In The Ecology and Control of Feral Cats pp. 63-61. UFAW, Potters Bar, London. Tabor, R. (1983) Wildlife of the Domestic Cat. Arrow Books, London. Turner, D. C. and Meister, O. (1986) Hunting Behaviour of the domestic cat. In Turner, D. C. and Bateson, P. (eds.) The Domestic Cat the Biology of its Behaviour, pp. 111-121. Cambridge University Press, Cambridge. Wilmore, G. T. D. (2000) Alien Plants of Yorkshire. Yorkshire Naturalists’ Union, Leeds. APPENDIX 1: NEUTERING RATES OF FARM CATS IN YORKSHIRE ALL CATS FEMALES MALES YORKSHIRE REGION Number of Cats* Recorded as Neutered % neutered (all sexes) FEMALE Intact Female Neutered Female % females neutered MALES Intact Male Neuterd Male % males neutered VALE OF MOWBRAY 119 5 4.2 83 80 3 3.6 36 34 ' 2 5.6 VALE OF YORK 127 9 7.1 101 94 7 6.9 26 24 2 7.7 HUMBERHEAD LEVELS 100 13 13.0 57 49 8 14.0 43 38 5 11.6 VALE OF PICKERING 20 0 0.0 13 13 0 0.0 7 7 0 0.0 PENNINES 332 37 11.1 212 194 18 8.5 120 101 19 15.8 WOLDS 166 31 18.7 87 76 11 12.6 79 59 20 25.3 HOLDERNESS 54 8 14.8 31 28 3 9.7 23 18 5 21.7 NORTH YORK MOORS 53 2 3.8 31 30 1 3.2 22 21 1 4.5 TOTAL 971 105 10.8 615 564 51 8.3 356 302 54 15.2 * of the 1,169 cats, gender and neutering data was only provided for 971. APPENDIX 2: YORKSHIRE FARM CAT SURVEY 126 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat ^ W o g s* S§OQ sun0 sdBJi SUOSIOJ OOO—IOOOO O— — ^040c00inv0 04 Oh^i'incn'ca ggSS < h S 2 wwgg “£“o 0AIP9JJ9UI 3AIP0JJ3 OOO^OO^H-H coino400inr-"co - GO 04 04 < IBJ3J z o gg|H w ^ 5 a 3C IOJJU03 juapofl SJ9J N h ^ h OO ^ ff) - S1V3 TO ON S}B3 jo ^r^(N(NONON^m hOO\OH(Nimo\ ^ Tt — aovanov s in co ■ VO OO ; Os 04 )" -h" VO oo h m m VD 04 ^ 04 VO CO © 04 co '-h '-h in Aiqnoj S9SJ0JJ SJBOO o o o o Tf 00 O Tj- O CO o o o o o o o daaqs S§IJ 3W«D ajqBjy rHMoo\roo\|-oin >n -h 04 VDOlO'tO^lNOO 'sO^h'sD'sDOCNcOCO ,-H O' 04 ^ — CO ^ ^ 1 —< CO VO O' I 04 04 CO Q3A3AMS SH3WdV3 30 ON i-3 c n W £ > © Bag £ X P £ e* ix a g g a S Q“hz i-J § ftS 7; ogog ffi ffi Z CU ^ O 3g « 2 PQ S ggo S Oh >* fa (Jh ooog w a a j J j j o £ £ £ § 127 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat APPENDIX 3a: AGE STRUCTURE OF CAT POPULATIONS ON YORKSHIRE FARMS AGE Total Population Intact Female i rX 3 z Intact Male Neutred Male <12 months 62 32 28 2 1 65 37 3 22 3 2 117 68 8 30 11 3 84 45 6 29 4 4 70 42 7 17 4 5 51 17 3 25 6 6 48 16 10 9 13 7 15 9 4 2 8 16 8 1 6 1 9 8 4 1 1 2 10 8 6 2 11 3 3 12 8 2 3 3 13 14 1 1 15 2 2 16 17 18 19 20 Totals 558 290 44 171 53 APPENDIX 3b: AGE STRUCTURE OF DOMESTIC CAT POPULATIONS AGE Total Population Intact Female Neutered Female Intact Male Neutred Male <12 months 27 9 7 8 3 1 21 4 6 4 7 2 35 4 11 8 12 3 49 15 14 7 13 4 23 4 7 6 6 5 24 5 7 7 5 6 16 4 6 3 3 7 14 4 4 6 8 8 2 2 4 9 6 1 1 1 3 10 5 2 2 1 11 4 2 2 12 5 2 3 13 4 1 1 2 14 1 1 15 16 1 1 17 1 1 18 19 1 1 20 Totals 245 55 75 44 71 128 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat APPENDIX 4: DOMESTIC CAT REGIONAL HABITAT AND OWNERSHIP SURVEY District Category Housing Type Habitats Cat Population SAMPLE SURVEYS (Each sample consisted of 10 adjacent properties) Terrace Flats Semi-detached Detached Garden to 50y2 Garden from 50-1502 Garden above 150y2 Allotments Parks & Sports Fields Woodland Arable Pasture PROPERTIES CENSUSED Number of Cats Mean Cats per 100 household Urban 108 31 5 64 10 53 51 7 9 36 2 0 0 1,080 271 25.09 Sub-Urban 138 14 0 84 58 21 91 45 12 51 27 23 27 1,380 466 33.76 Rural 125 23 0 50 71 21 46 56 7 12 21 56 82 1,250 664 53.12 Totals 371 68 5 198 139 95 188 108 28 71 50 81 109 3,710 1,401 37.76 APPENDIX 5: CAT GENDER AND NEUTERED RATES Housing Category All Cats Females Males Number of Cats* Recorded as Neutered % neutered (both sexes) FEMALE Intact Female Neutered Female % females neutered MALE Intact Male Neutered Male % males neutered Urban Sub-Urban Rural 201 323 634 146 237 275 72.6 73.4 43.4 98 147 355 25 43 202 73 104 153 74.5 70.7 43.1 103 176 279 30 43 157 73 133 122 70.8 75.9 34.7 Totals 1,158 658 56.8 600 270 330 55.0 558 230 328 58.8 * of the 1,401 cats, gender and neutering data was only provided for 1,158. 129 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat APPENDIX 6: RANGE OF PREY AND SEASONALITY IN PREDATORY ACTIVITY OF DOMESTIC CATS IN YORKSHIRE (ALL AREAS) SAMPLE SIZE: 180 Cats; Total vertebrae prey items: 5,205; Mean vertebrate prey items per cat: 28.91 PREY JAN FEB MARCH APRIL MAY JUNE 1 AUG SEPT OCT NOV DEC TOTAL INVERTEBRATES Bee/Wasp 1 1 1 1 3 7 Beetle 3 1 4 Butterfly 1 4 3 7 7 19 8 49 Fly 2 3 1 5 2 5 3 1 22 Moth 1 1 1 2 2 5 2 4 1 1 20 Spider 1 1 1 2 1 3 9 Worm 1 1 3 5 Sub-total (invertebrates) 4 2 6 9 13 17 29 19 14 2 1 116 MAMMALS (indet) 4 5 3 9 8 8 12 7 18 14 4 5 97 BAT sp. (indet) 2 2 1 2 1 8 Mole 1 1 1 1 10 5 2 3 1 25 SHREW sp. (indet) 3 7 11 15 33 24 71 47 56 27 19 6 319 Common Shrew 1 2 8 7 10 14 2 2 2 3 51 Pygmy Shrew 1 5 14 11 5 1 37 Water Shrew 2 2 VOLE sp. (indet) 25 18 30 23 30 20 46 37 52 52 44 33 410 Bank Vole 1 1 1 2 5 5 3 3 12 7 6 46 Field Vole 19 12 3 3 3 8 32 11 27 22 21 11 172 Water Vole 1 1 2 MOUSE sp. (indet) 69 47 63 98 79 92 141 121 174 452 121 96 1,253 Harvest Mouse 1 1 House Mouse 6 4 4 9 3 2 5 5 5 7 6 3 59 Wood Mouse 30 44 59 65 56 87 85 81 151 111 52 26 847 Brown Rat 2 2 4 5 5 7 4 8 25 26 12 3 103 Grey Squirrel 2 1 3 Hamster 1 1 Rabbit 2 3 13 19 27 23 42 15 10 3 2 159 Weasel 1 2 2 5 Sub-total (mammals) 160 143 185 244 250 306 456 392 537 441 294 195 3,600 FISH Goldfish 1 1 1 3 Roach 1 1 2 Stickleback 1 1 HERPTILES Frog 3 1 1 3 1 2 11 Newt sp. (indet) 1 1 2 Sub-total (herptiles & fish) 1 5 1 1 4 3 2 2 19 130 Red in Tooth and Claw: 2. Studies on the Natural History of the Domestic Cat PREY JAN FEB MARCH APRIL MAY JUNE JULY AUG SEPT OCT NOV DEC TOTAL BIRD sp. (indet) 15 6 10 30 67 83 62 71 44 25 19 27 459 Blackbird 6 5 4 2 16 19 7 5 4 1 1 4 74 Black-headed Gull 1 1 Blue Tit 3 2 1 2 3 6 4 2 1 6 5 4 39 Budgerigar 1 1 2 Bullfinch 1 1 2 Canada Goose (gosling) 1 1 Canary 1 1 Chaffinch 1 1 1 2 5 Chicken (chick) 1 1 Chiff Chaff 1 1 Collared Dove 1 1 3 1 1 7 Dunnock 2 1 1 1 3 9 6 23 Goldcrest 3 3 Goldfinch 2 5 1 1 1 10 Great Tit 2 2 4 Greenfinch 1 2 2 1 1 3 1 1 2 1 15 House Martin 2 4 6 House Sparrow 46 20 32 34 87 129 116 86 67 35 20 23 695 Linnet 1 1 Mallard (duckling) 2 2 Meadow Pipit 1 1 Moorhen 1 1 1 3 Pheasant 1 1 Pied Wagtail 2 2 Redpoll 1 1 Redwing 1 1 Robin 2 3 2 2 2 6 7 5 3 1 2 35 Song Thrush 3 1 2 7 18 9 5 1 5 3 54 Starling 2 5 10 12 28 8 9 7 4 3 2 7 97 Street Pigeon 1 1 1 3 Swallow 1 1 2 Willow Warbler 2 1 3 Wood Pigeon 3 1 1 3 2 1 1 3 1 16 Wren 1 3 2 3 1 1 1 12 Yellow Hammer 1 1 1 3 Sub-total (birds) 88 44 66 88 220 292 245 193 137 84 51 78 1,586 Sub-total (mammals) 160 143 182 244 250 306 456 392 537 441 294 232 3,600 Sub-total (herptiles & fish) 1 5 1 1 4 3 2 2 19 Sub-total (birds) 88 44 66 88 220 292 245 193 137 84 51 78 1,586 Total Vertebrates 248 188 253 333 471 602 704 587 676 525 345 310 5,205 131 BOOK REVIEWS Mammals of Europe by David W. Macdonald and Priscilla Barrett, Pp. 312, with over 600 colour illustrations & 200 b/w line drawings. Princeton University Press, Princeton. 2002. $24.95 softback. First compiled by F. H. van den Brink under the familiar title Collins Field Guide to the Mammals of Britain and Europe, its English translation by Hans Kruuk and H. N. Southern was published by HarperCollins in 1967. Van den Brink’s text and Paul Barruel’s colour illustrations gave stout service to generations of naturalists, helping in no small way to encourage the subsequent revolution in European mammalian studies. This new fund of information, expertly marshalled by Professor Macdonald, and the fresh portfolio of artwork by Priscilla Barrett, has achieved a brilliant overhaul of this valued and trusted volume. The 64 colour plates include over 600 of Barrett’s excellent paintings of the animals and sketches of their tracks, nests, burrows, field signs, teeth and young. Over 200 distribution maps are significantly updated from the 1967 edition. This is an impressively comprehensive contribution to the Princeton Field Guide series, reviewing and comparing the descriptions, habitats, breeding, behaviour and field signs of more than 200 mammals occurring in Europe and its surrounding seas. Having delectably wallowed through Professor Macdonald’s 4.5 kg New Encyclopaedia of Mammals, I can report that this handy pocket-sized volume (130 x 195 x 25 mm) is a masterpiece of precis. An advantage of compression is that series of similar species are illustrated together, enabling subtle differences to be highlighted. I particularly draw attention to the excellent series of bat illustrations, prepared with advice from Dr Bob Stebbings, which alone justifies the acquisition of this book. Interesting additions are tables comparing the hydrophore call, clicks and moans of cetaceans and bat detector ultrasound pulses of bats. The acknowledgements to numerous contributing experts, including M. J. A Thompson, past President of the YNU, testify to this book’s technical rigour. CAH Birds of the Harrogate District by John R. Mather. Pp. 310. 90 b/w line drawings Harrogate and District Naturalists’ Society, Harrogate. 2001. £10 softback, available from: Mrs J. McClean, 6 Rossett Park Road, Harrogate HG2 9NP. With an enthusiastic forword by Ian [D.I.M.] Wallace, this book forms a complete but concise historical review of some 288 bird species shown to have occurred in the Harrogate area from the 1 8th century to the present. Line drawings in the form of miniature vignettes, contributed by Ray G. Hawley, D. I. M. Wallace, M. Whorley and the author himself help to punctuate the systematic list. Also included is a fascinating account of the geography, topography and habitats of the Harrogate region, a potted history of the Harrogate and District Naturalists’ Society (the local ornithological recording body) and a history of ornithologists and their investigations of the local avifauna. There is a glossary of ornithological terms, an extensive bibliography and a useful map of the Harrogate recording area. The compact (A5) format of this tome conceals a mighty work, corpulent with carefully researched and critically evaluated detail and laced with the phraseology and personality of the author: you’ve heard Johnny Mather lecture, you’ll be able to hear him performing the script as you read. Out of the Thomas Nelson mould of ornithological writers, every historical source has been tracked down and every record woven into a meaningful review by an author steeped in local knowledge born of a lifetime of enthusiastic hands-on involvement in natural history. John Mather’s book is an essential work of reference for Harrogate naturalists in general and Yorkshire ornithologists in particular, and many will enjoy and benefit from this 132 Book Reviews historic compilation, but more than this, it stands as a masterly and collectable example of a faunal review, the calibre of which few have been capable since the time of the Victorian and Edwardian naturalists. The investment of effort, dedication and craftsmanship necessary to create a work of this eminence is impressive and the acquisition of two copies, one to use and one to archive is certainly justifiable here. Hopefully, the advent of the new millennium will encourage the production of equivalent biodiversity reviews around the Yorkshire region. Birds of the Harrogate District will certainly stand as a model in this respect, though I suspect that in our abbreviated email, text message and internet world of communication, skilfully used words are likely to give way to skilful statistics and computer graphics. CAH Alfred Russel Wallace: a Life by Peter Raby. Pp. xii + 340, inch 2 maps & numerous illustrations, plus 16 pp. of b/w plates. Chatto & Windus, London/Princeton University Press. 2001. £20.00 hardback; Pimlico. 2002. £12.50 paperback. At long last Wallace, to judge by the number of books recently or in the course of being published on him, is receiving the due attention he so rightly deserves as co-founder of the theory of evolution. This latest contribution is both readable and well researched, tracing Wallace’s modest background, early career, self-education, his passion for biology realized by his exploration of the Amazon and Malay Archipelago, and the circumstances surrounding the presentation of his and Darwin’s papers on natural selection at the famous Linnean Society meeting in 1858. Wallace never begrudged Darwin’s pre-eminent role in the dissemination of the theory, and according to Raby . . seemed genuinely relieved that he had not been compelled to write . . .” The Origin of Species. Upon his return to England in 1862, Wallace, despite ill health and financial problems, commenced work on The Malay Archipelago , a milestone in natural history published seven years later. Following this, his interests greatly diversified, and some of his enthusiasms, such as his defence of spiritualism, were heavily criticized. Nevertheless, his Island Life , published in 1880, received due acclaim, and he continued to be honoured in terms of a Civil List pension, medals, degrees, Order of Merit, etc. In later life his mind remained ever active, particularly in respect of social and political compaigns, and his published output never ceased, with books and articles on a wide variety of subjects (including anti-vaccination, land-laws, state endowment of education) continuing right up to his death in 1913. According to Raby, “Wallace, in the twentieth century, was not only the popular exponent of the previous century’s scientific achievement, but represented the living embodiment of its changes and developments. His steady stream of books, and his articles and letters in the intellectual and popular press, kept his name in the public eye”. Of particular interest to Yorkshire readers will be the numerous references to Richard Spruce: it is remarkable to think that three of the greatest naturalists of their time, Bates, Spruce and Wallace, were working on the same relatively short stretch of the Amazon. Wallace was undoubtedly influenced by the character and work of Spruce, his closest scientific friend, whose extensive journals (and correspondence) he painstakingly edited - a true act of homage; these were subsequently published in 1908 as Notes of a Botanist on the Amazon and Andes. Although Raby shows his non-scientific background, he is to be congratulated on his rich and powerful portrayal of Wallace and for justifiably raising the profile of a truly remarkable man. MRDS BIRDS ON THE SPURN PENINSULA by Ralph Chislett Parts I and II (1996), edited by Michael Densley. Hardback 218 pages with coloured dust jacket, coloured frontispiece and twelve photographic illustrations in black and white. This enlarged edition is the first time that Part II has appeared in print. Not surprisingly this book virtually sold-out remarkably quickly; however the printers have discovered a small quantity of the remaining books. Before releasing them on the open market, the publisher has offered this limited stock to YNU members for the same price as when it was published in 1996 - £14.95 per copy post free. Owing to a slight mix-up at the time, the newly released book has not previously been advertised in The Naturalist - thus members now have a chance to purchase a copy. The James Reckitt Charity of Hull and other individual benefactors have made a substantial contribution towards the publishing cost in order to keep the selling price relatively low. Apply to: Clive Varty, 61 St Chads Court, St. Chads Road, Leeds LS16 5QX Titus Wilson Kent Works • Burneside Road • Kendal • Cumbria • LA9 4RL Tel. 01539 720244 Specialist printers /binders of Academic Journals , Catalogues and Private Publications Our service includes attending to worldwide distribution Binding Why not have your copies of The Naturalist bound into volumes? One year’s issues per volume, or alternatively two years in one volume at less cost than binding as two separate volumes. We are also experienced in binding thesis and experts in the re-binding and repairing of all books. Spink & Thackray Broomfield Binder Back Broomfield Crescent Telephone: 0113 2780353 LEEDS LS6 3BP Fax 01 13 2304155 www.spinkandthackray.co.uk Printed in Great Britain by Titus Wilson & Son, Kendal ISSN 0028-0771 Latest publication of the Yorkshire Naturalists’ Union ALIEN PLANTS OF YORKSHIRE GEOFFREY WILMORE The first comprehensive modem publication dealing with the alien plants of any major British county, detailing the flora (almost 1300 species) of Yorkshire’s wool waste dumps, railway sidings, sewage works, shoddy fields, industrial wasteland, dockland and oil mill sidings. Pp. 316, including 15 line drawings; published January 2000 Normal price: £15.00 (plus £2.00 per copy p.&p.) Reduced price for members of the YNU: £12.00 (plus £2.00 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Mr John A. Newbold, Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS THE FRESHWATER CRUSTACEA OF YORKSHIRE a faunistic & ecological survey GEOFFREY FRYER The crustacean fauna of Yorkshire reflects the great physiographic diversity of the region. Adopting an ecological approach, this book considers the Yorkshire fauna in relation to climate, topography, geology, soils and water chemistry, always keeping in mind that it is dealing with living organisms whose habits, requirements and physiological limitations determine exactly where they live. Matters covered include the ecological background; faunal assemblages and their regional attributes; an analysis of the factors that determine distribution patterns, many of which are mapped; wide geographical aspects; and conservation. Large areas, such as the Pennines, Howgill Fells, North Eastern uplands and the lowland plains are surveyed. So too are localised regions including Whemside, the Malham area, lowland heaths, and the largest lakes, as well as habitats such as upland tarns, seepages, cold springs, small lowland ponds, inland saline waters. Notes are given on every species recorded, including parasitic forms. Price £8.00 (plus £2.00 p.&p.) Special offer to members of the Yorkshire Naturalists’ Union £5.00 (plus £2.00 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Professor M. R. D. Seaward, Department of Environ- mental Science, University of Bradford, Bradford BD7 1DP. October-December 2002 Number 1043 A QUARTERLY JOURNAL OL NATURAL HISTORY LOR THE NORTH OL ENGLAND Barn Owls (Tyto alba) in Churchwardens’ Accounts of the 17th and 18th Centuries: An Indication of Past Abundance in East Yorkshire - C. A. Howes The Dawn Chorus and How Its Singers are Conducted - Geoffrey F.ryer Y.N.U. Bryological Section: Annual Report 2000-2001 - T. L. Blockeel and J. M. Blackburn Yorkshire Naturalists’ Union Excursions in 2000 - A. Henderson and Janetta Lambert Published by the Yorkshire Naturalists’ Union Editor M. R. D. Seaward MSc, PhD, DSc, FLS, The University, Bradford BD7 1DP Notice to Contributors to ‘The Naturalist’ Manuscripts (two copies if possible), typed double-spaced on one side of the paper only with margins at top and left-hand at least 2.5cm wide, should be submitted. Latin names of genera and species, but nothing else, should be underlined. S.I. Units should be used wherever possible. Authors must ensure that their references are accurately cited, and that the titles of the journals are correctly abbreviated. Volumes of The Naturalist for the years 1886 to 1975 have been retrospectively numbered 11 to 100 to accord with numbering before and after this period (see YNU Bulletin no. 3, pp. 21-22 1985); please cite these volume numbers in all references. Table and text-figures should be prepared on separate sheets of paper. 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The editor, on behalf of the YNU, wishes to record this deep appreciation of this imaginative gesture. © Yorkshire Naturalists’ Union — 2002 Single copies may be made of single articles in this journal provided that due acknow- ledgement is made and the copies are for non-profit making educational or private use. Copying of more than one article or multiple copying of a single article is forbidden unless special permission has been obtained from the Yorkshire Naturalists’ Union. Permission is granted for the use of brief quotations in published work provided that acknowledgement of the source is clearly stated, but the use of substantial sections of text and any illustrative matter requires the express permission of the Yorkshire Naturalists’ Union. Registered Office (for all enquiries, correspondence and payments): Mr John A. Newbould, Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS (tel: 01305-837384; email: janewbould@aol.com) The Naturalist is issued free to individual members of the Yorkshire Naturalists’ Union and to Affiliated Societies. Institutions and Subscribers £24.00 Registered Charity No. 224018 133 BARN OWLS ( TYTOALBA ) IN CHURCHWARDENS’ ACCOUNTS OF THE 17th AND 18th CENTURIES: AN INDICATION OF PAST ABUNDANCE IN EAST YORKSHIRE C. A. HOWES Doncaster Museum & Art Gallery DN1 2AE . . . and it shall be an habitation of dragons, and a court for owls. Isaiah 34: 11-15 Introduction Investigation into long term distribution and status changes of Britain’s vertebrate fauna is severely hampered by the paucity or absence of accessible data sources earlier than the 19th and 20th centuries. A particularly fruitful but seldom used source of documentary evidence, however, is the series of ‘vermin’ bounty payments contained in the yearly accounts of churchwardens and other parish or township officials relating to the period from the late 16th to the early 19th centuries. These documents, hand- written and often on vellum, are generally housed in Local Authority Archives or Records Offices. Though somewhat laborious to locate, decipher and extract, the particular value of these sources lies in their meticulous, statistically based and long term nature. Beverley Churchwarderns’ Accounts An examination of the churchwardens’ accounts for the parish of St. Mary’s, Beverley, which date from 1593 to 1831, show that between 1642 and 1736 six pence per head was paid for at least eighty-five owls (see Appendix 1 ). Figure 1, which gives the yearly distribution of owl bounty payments, shows that from one to seven owls were killed per year. Figure 2, which examines the number of owls killed at any one time, shows that although on most (thirty-one) occasions only single owls were killed, on seventeen occasions between two and five owls were killed. The Identity of ‘Church Owls’ There are no precise descriptions in the Beverley archive as to which species of owl was involved and why they were being killed; however, clues to both are to be found in account entries (see Appendix 1). On many occasions bounties for the killing of owls were linked with payments for work which was probably done within the church, for example mending locks in 1663, making locks in 1670 and for supplying candles in 1674. The washing of clerics or chorister’s surplices and church linen featured with killing owls in 1674, 1680, 1687 and twice in 1693. References to the killing of ‘birds’ specifically within the church come in two 1711 entries for ‘For powder and shot for killing birds in the church’. Specific mention of owls being killed within the church again comes from 1711 and 1712 entries. It may be significant that these entries follow those for the payments for powder and shot. Circumstantial evidence of owls being encountered in the bell tower comes in entries for 1674, 1675, 1678 and 1679 when payments were also made for work on the bell chamber or the bells themselves. Since the owl most closely associated with buildings is the barn owl and from the 17th century the term ‘church owl’ was an accepted vernacular synonym for the barn owl (Nelson 1907), the target species is therefore likely to have been the barn owl Tyto alba. Cultural Attitudes and Possible Reasons for Killing Owls Geoffrey Chaucer in The Parlement of Foules (written between 1372 and 1382) refers to The oule, that of deth the bode bringeth and is perhaps one of the earliest of a long series of literary figures to dwell on a perceived sinister and unearthly quality of owls. Naturalist 127 (2002) 134 Barn Owls (Tyto alba) in Churchwardens’ Accounts of the 17th and 18th Centuries FIGURE 1 Annual records of bounty payments made for owls in the Parish of St Mary’s, Beverley during the 17th and 18th Centuries. 1 2 3 4 5 NUMBERS OF OWLS FIGURE 2 Numbers of Owls presented for bounty payments on individual occasions. The various biblical references, notably in the books of Isaiah (14: 20-21 and 34:11-15) and Leviticus (11: 13-17), imply that owls were ‘an abomination’ and ‘unclean’ and were associated with destruction, desolation and ruination. This authoritative doctrine, together with the owl’s eerie nocturnal hoots and shrieks was probably the basis of a widespread superstitious fear and folkloric prejudice against these birds. Despite this weight of cultural negativity, in a rural agricultural context, notably in grain Barn Owls (Tyto alba) in Churchwardens’ Accounts of the 17 th and 18th Centuries 135 producing regions, owls were sympathetically regarded due to their rodent control capacities. Farm buildings of the 18th and 19th centuries were frequently constructed with ‘owl holes’ in their gable apexes to enable Barn owls to enter and breed (Jones 1972). Quite why the public purse in the form of parish rates should have promoted the killing of owls is not immediately clear, though the examination of the numbers and periodicity of kills at Beverley and a comparison with evidence from parishes elsewhere in England and Wales provide an interpretation. References to bounties being paid for killing owls are very rare and one may conclude that the rodent pest controlling virtues of owls were probably widely recognised. Decisions as to which species were to be regarded as ‘vermin’, and the levels of bounty tariff were parochial prerogatives. In practice, the vicar and churchwardens decided in the light of prevailing circumstances, and records of such decisions can occasionally be found in vestry minutes. The frequency and levels of payments for given ‘vermin’ species can therefore reflect a species’ perceived ‘pest’ status within a given parish at a given time. Significantly there is no mention of owls in the extensive surveys of churchwardens’ accounts from seventy-nine Bedfordshire parishes (Elliott 1936) or from twenty-two Hertfordshire parishes (Oldham 1931a). In Yorkshire, the accounts of some eighty-eight parishes studied to date (Howes in prep.) have only produced evidence from Beverley. Generally, the owls for which bounties were paid were those which were actually inhabiting the church itself. Ticehurst (1935) in reviewing the copious numbers of birds brought to the churchwardens of Tenterden, Kent between 1626 and 1712 only encountered one reference to owls where, in 1646-7 a shilling was paid for killing two ‘owles which doe use in the churche’. Similarly, Oldham (1931b), investigating the bounty payments made by the churchwardens of Llanynys in the Vale of Clwyd in the years 1635-63, 1738-69 and 1788-1838, only encountered a single entry, for sixpence paid in 1769 for the killing of an owl in church. In 1759 two pence was paid for bird lime to catch owls in the church of St John’s, Chester. Preventative methods were also used by church officials; hence in 1711 at Redenhall, Norfolk, three shillings and six pence was paid for ‘work and stuffe and nails in stopping out ye owles at ye church’ (Bunn et al. 1982). In St. Mary’s, Beverley, the groupings of records, often in two to four year periods (see Figure 1), indicate successive concerted attempts to remove owls. The frequency of multiple specimens (from two to five) being collected at any one time (see Figure 2), strongly suggests that the clearing of occupied owl nesting sites could well have been the purpose of these actions. The considerable debris of droppings, pellets, discarded corpses of prey items etc., particularly in the vicinity of an occupied nesting site, could understandably have been deemed unacceptable in a holy place, evoking the passage in Isaiah 14, 20-21 ‘. . . their houses shall be full of doleful creatures; and owls shall dwell there Earliest Yorkshire Records and an indication of former abundance In the absence of palaeontological or archaeological evidence, the earliest indication of the barn owl occurring in Yorkshire has long been held to be from Willoughby’s Ornithology of 1678 and refers to Mr Ralph Johnson alluding to the ‘Church Owl’. Since Johnson corresponded with Willoughby from Brignall near Greta Bridge (then in Yorkshire), this is taken as evidence that the allusion referred to Yorkshire (Nelson 1907). The Beverley sample, dating from 1642, provides a slightly earlier set of dates and is from a guaranteed provenance. More importantly, the Beverley accounts, itemising at least eighty-five owls between 1642 and 1736 with up to seven being recorded per year, is unparalleled in parishes elsewhere in Britain. This study therefore provides evidence of a persistent and evidently robust barn owl population in the Hull valley from the mid- 17th to the mid- 18th centuries. Interestingly this period pre-dates the widespread drainage of the Hull valley region which commenced in the 1760s (Sheppard 1958). Thus, in addition to the managed post-medieval mosaic of pasture, meadow and arable fields, the owls would have had access to extensive areas of wetland to hunt over. 136 Barn Owls (Tyto alba) in Churchwardens’ Accounts of the 17th and 18th Centuries Acknowledgement Thanks are due to Mr Ian Mason of the East Riding of Yorkshire Council Archives and Records Service, for permission to publish items from Beverley churchwardens’ accounts (Archive Ref. PE 1/51 to 126). References Bunn, D. S., Warburton, A. B. and Wilson, R. D. S. (1982) The Barn Owl. T. & A. D. Poyser, Calton. Elliott, J. S. (1936) Bedfordshire ‘Vermin’ Payments. Luton Museum, Luton. Howes, C. A. (1992) Historical notes on the starling Sturnus vulgaris in Yorkshire. Naturalist 117: 109-111. Jones, E. L. (1972) The bird pests of British agriculture in recent centuries. Agricultural History Review 20 : 107-125. Nelson, T. (1907) The Birds of Yorkshire. Brown, Hull. Oldham, C. (1931a) Payments for ‘vermin’ by some Hertfordshire churchwardens. Transactions of the East Hertfordshire Natural History Society and Field Club 19 (2): 79-112 Oldham, C. (1931b) Payments for ‘vermin’ and for dog-tongs. North Western Naturalist 6 (1): 13-17. Ray, J. (1678) The Ornithology of Francis Willughby of Middleton in the County of Warwick. Royal Society, London. Sheppard, J. A. (1958) The draining of the Hull Valley. Local History Series No. 8. East Yorkshire Local History Society, Hull. Ticehurst, N. F. (1935) Rewards for vermin killing paid by the churchwardens of Tenterden, 1625 to 1712. Hastings and East Sussex Naturalist 5 (2): 69-82. APPENDIX 1 Details of owl bounty payments in the Churchwardens’ accounts of the Parish of St. Mary’s, Beverley 1593 to 1831 (East Riding County Records Office Ref. PE 1/51 to 126) Year Entry Bounty payment £. s. d. 1642 Item paid . . . Johnson for killing 3 Owls in the Wood Hall Closes which he did testify to belonging to the church -. 1. 6 1663 Paid for killing 5 Owls -. 2. 6 1663 Item paid for mending locks . . . and killing 1 Owl -. 2. - 1663 Item paid for working on Mondays and Tuesdays and killing 1 Owl -. 2. - 1667 Paid for killing 5 Owls -. 2. 6 1667 Pd. for killing of old Fox and cub & 1 Owl -. 1. 4 1670 Item paid for work and making locks and for killing 1 Owl -. 4. 6 1671 Pd. for mending lock and killing 1 Owl -. 1. - 1671 Pd. for killing 3 Owls -. 1. 6 1671 Pd. for killing 1 Owl -. -. 6 1672 Pd. for killing 1 Owl 6 1672 Pd. for two baskets and killing 1 Owl -. 1. 6 1672 Pd. for killing 1 Owl 6 1672 Pd. for killing 1 Owl 6 1674 Item for . . . and staples for the bells and killing 2 Owls -. 3. 6 1674 Item to . . . nails and killing 1 Owl -. 1. 6 Barn Owls (Tyto alba) in Churchwardens’ Accounts of the 17th and 18th Centuries 137 Year Entry Bounty payment £. s. d. 1674 Item to Robert Thompson for your work about the church and killing 1 Owl 2. 1674 Pd. for candles used on November 14th and washing surplice and killing 1 Owl _ 2. 8 1675 Item for 7 yds of matting for bell chamber . . . and for killing 3 Owls 3 10 1676 Pd. for 1000 of . . . 1 bushells of coal and for killing 3 Owls 5. 2 1678 For work about the bells and killing 4 Owls 5. - 1679 For . . . and killing 1 Owl 2.10 1679 For killing [2] Owls . . . bells and other work 5. 6 1680 For washing surplices and killing 1 Owl 2. 6 1687 For killing 2 Owls and working 1. 4 1687 For washing surplices and killing 1 Owl 2. 6 1690 Spent at . . . ground and for killing 4 Owls 3. - 1691 For sack and killing 3 Owls 2. 372 1692 To John Tuting for killing [4] Owls 2. - 1692 To Richard Tuting for killing 1 Owl -. 6 1693 To washing linen and killing 4 Owls 4. - 1693 To washing surplices and killing 1 Owl 1. 6 1695 To John Tuting for an Owl -. 6 1697 For killing 1 Owl 6 1710 Pd. to John Tuting for 1 Owl killing 6 1710 Pd. 1 Owl killing 6 1711 For powder and shot for killing birds in the Church 4 1711 For powder and shot for killing birds in the Church 4 1711 To John Tuting for killing 1 Owl in the Church 6 1712 For killing 1 Owl in the Church 6 1714 For [1] Owl killing 6 1714 For 1 Owl killing 6 1714 Pd. to John Tuting for 1 Owl 6 1718 For 2 Owls killing 1. - 1719 To John Tuting for killing 1 Owl 6 1722 An Owl killing in the Church 6 1723 Pd. John Tuting for killing an Owl 6 1723 Pd. for sand 6d. and Pd. John Tuting for 3 Owls killing 2. - 1723 Pd. John Tuting for killing 1 Owl -. 6 1736 Pd. for killing [2] Owls -• 1. - BOOK REVIEWS A Field Guide to the Mammals of Australia by Peter Menkhorst, illustrated by Frank Knight. Pp. x + 269, with 116 colour plates, numerous line drawings & 380 maps. Oxford University Press, South Melbourne, Australia. 2001. £15.95 softback. This guide covers all 379 mammal species, indigenous and introduced, living in Australia and its adjoining seas. The introductory chapters include details of how to use the guide, field techniques, a checklist and extensive identification keys. Throughout these chapters the reader is supplied with numerous simple line drawings to assist with field observation. The main habitat types are described through the use of twelve colour plates. The main account follows. Here the left page provides details of each species - its description, range, distribution map, habitat, status, voice, behaviour etc - and on the 138 Book Reviews opposite page a high quality colour illustration. By using a small, but very readable print, the text includes a remarkable amount of useful ecological information, often supplemented with small drawings highlighting diagnostic identification characteristics. The book is compact and readily transportable in the field, with a format that makes for quick and easy identification of the more conspicuous mammals. The guide will certainly enable identification of the smaller species such as dunnarts, bats and rodents but inevitably these will need to be close at hand. This is an excellent, inexpensive field guide, essential for any naturalist visiting Australia. MJD The Nature of Mediterranean Europe: an ecological history by A. T. Grove and Oliver Rackham. Pp. 384, with 123 b/w illus. + 245 colour plates. Yale University Press, 2001. £45.00 hardback. This substantial book represents the fruits of several research programmes funded by the EU over the past two decades. It is based on fieldwork by the authors and their collaborators in many parts of Mediterranean Europe and includes a large number of detailed case studies. The coverage is very wide, from geology and climate to erosion and water resources. The first thirteen chapters cover the physical geography, ecology and ecological history of the region. The last seven chapters focus on the specific issues of erosion and desertification. The major theme is that many previous writers have assumed Mediterranean landscapes to be the result of human mismanagement - the ‘Ruined Landscape Theory’. Much of the book is aimed at showing this theory to be unfounded and revealing the intricate balance between geology, climate, vegetation, soils and land-use. The authors find little evidence to support the notion that the area is currently undergoing desertification or suffers from a major problem of soil erosion. They discuss the role of deluges, both in historical times and at the present day, and show these unpredictable events to be a major force in shaping the landscape. The book is written in an accessible style with a minimum of technical jargon. The bibliography is impressive, including many historical sources, and there are good footnotes throughout the book. It is lavishly illustrated with superb photographs and numerous useful maps and diagrams. The standard of. presentation is very high. This is a weighty tome and on the pricey side, but it is a mine of fascinating information for naturalists, geographers and historians. The authors have, perhaps, overstated their case against ‘Ruined Landscape Theory’, references to which occur throughout the book with tedious frequency. However, the refreshing, field-based approach makes for stimulating reading, even though at times it leads to sweeping statements and over-generalisation. Not everyone will agree with the authors’ conclusions but this is certainly a book that should be read by all serious scholars of Mediterranean Europe. MAA Scotland’s Nature & Wildlife by Kenny Taylor. Pp. 224, with full colour illus. 2002. Lomond Books, Edinburgh. £15.00 hardback. Lovers of nature will find much to interest and instruct them in this book. Scotland is a country with many different types of habitat and a rich diversity of wildlife. Kenny Taylor, with decades of experience behind him, first discusses what to look for, covering plants, birds, mammals, insects and amphibians, when to find each species seasonally, and where you might best expect to see them. The wildlife of eight regions of Scotland is described individually, and the text is supported by clear maps and, where appropriate, further notes on sites of special significance. While not a field guide, it is one of the most comprehensive popular guides to Scotland’s wildlife. The book is lavishly illustrated throughout with stunning photographs by some of the country’s leading wildlife photographers. It is an attractive book to browse through, and an instructive reference and travelling companion. DAC 139 THE DAWN CHORUS AND HOW ITS SINGERS ARE CONDUCTED GEOFFREY FRYER Elleray Cottage, Windermere, Cumbria LA23 1AW That the species of birds which contribute to the dawn chorus do not all begin to sing at the same time has long been known, and the sequence in which they do so has been referred to in various places. An example is provided by the work of the Leeds Naturalists’ Club, some of whose members spent a night in the Washburn Valley in 1935 recording the times at which different species began to sing or call. The results of their observations were reported in The Naturalist by Grist (1935). It was particularly noted that the chorus was not made up of all species singing simultaneously but that, for the most part, the various species replaced each other sequentially. Moreover, as Armstrong (1963) noted in his remarkable book A Study of Bird Song, in spite of the inevitable variations and anomalies, “the order in which different species sing in the morning remains remarkably constant”. The Sky Lark, Alauda arvensis, which of course is a contributor only in suitable habitats, is an early riser (as the Leeds Naturalists found) and Common Redstart, Phoenicurus phoenicurus, Blackbird, Turdus merula, Song Thrush, T. philomelas, Wood Pigeon, Columba palumbus, Robin, Erithacus rubecula , Wren, Troglodytes troglodytes, House Sparrow, Passer domesticus, Common Bullfinch, Pyrrhula nesa, and Common Chaffinch, Fringilla coelebs, generally begin to sing in that sequence. Armstrong drew attention to the fact that vermivorous and insectivorous species were early risers (was the coiner of the saying ‘the early bird catches the worm’ aware of this?) and, apart from pigeons, grain- eaters tend to be late risers. The explanation involves the acuity of vision and the behaviour of the prey of the early risers, some components of which are most readily available early in the morning. In essence, it implies that the order in which species begin to sing is related to differences in their ability to see at low light intensities. There is no point in a bird becoming active before it can see well enough to perform essential tasks effectively, though it may be advantageous to begin to sing while light intensity is still too low to enable it to forage profitably. By so doing it may be able to advertise, or indulge in vocal defence of, its territory for a short period without impinging on the time devoted to those vital activities for which vision is essential. Armstrong noted that “the times when birds arise and roost, and consequently are first heard in the morning and last heard in the evening, are related to their foraging behaviour as well as to light intensity”. He then went on to make the remarkably perceptive observation that “So closely related is the relative size of some birds’ eyes to the length of daily activity that in some instances one might make a fair estimate by looking at its eyes as to whether the bird sings early or late in the dawn chorus”. He cited as examples that the early rising Common Redstart, Black Redstart, Phoenicurus ochruros, and Robin have larger eyes than the late rising House Sparrow, and noted that these early risers are insectivorous and adapted for foraging in dim light. The Sky Lark is to some extent an exception in that it eats both invertebrates, especially in summer, and vegetable matter. As Armstrong says in a different context, “There are few principles of bird behaviour to which some species do not conform”! Thomas et al. (2002) have now tested the relationship between the times at which different species begin to sing and visual capability at low light intensities - which is related to eye size - and have confirmed the validity of Armstrong’s belief. Both visual sensitivity (the ability to detect light at low intensities) and visual resolution (the ability to distinguish detail at a given light intensity) can be enhanced by an increase in pupil aperture, in combination with other changes that need not concern us here. An increase in pupil aperture can be achieved by an increase in eye size. The larger an eye, the larger can be the diameter of its pupil. Thomas et al. (2002) confined their study to passerine birds. The times at which different species began to sing was ascertained from 40 sets of observations on bird communities at Naturalist 127 (2002) 140 The Dawn Chorus and how its Singers are Conducted seven sites, five in England or Wales, one in Portugal, and one in Switzerland. The number of species involved at a site ranged from 8 to 23. At two sites ambient light intensity was measured throughout the singing period. Maximum pupil diameter was estimated by measuring the diameter of the exposed eye surfaces, which could be done by dial callipers to an accuracy of 0. 1 mm. As the relationship between eye size and body size is important, body mass was also recorded by simple weighing. Although the sequence in which the species began to sing was not reported for any site, it was found that, as Armstrong had observed, species with large eyes began to sing at lower light intensities, i.e. earlier, than those with small eyes. The relationship was stronger when differences in body size were allowed for by statistical procedures. As the authors say in the precise but arid language of science, the results “provide robust support for the hypothesis that visual capability at low light levels influences the time at which birds start to sing at dawn”. To the naturalist, their comment that “our results do not necessarily imply that large eyes have evolved to allow a bird to start singing earlier at dawn” seems self-evident, and unnecessary. Known examples of echo location (the Oil bird and Cave Swiftlets) notwithstanding, vision is the key sense in birds, without which they would be helpless. The time at which they begin to sing at dawn, while probably of some selective significance, must surely be a secondary exploitation of more important advantages of a particular eye size that operate throughout every day of the bird’s life. Moreover, singing is predominantly a male activity. The contrast between Armstrong’s observation and its testing almost 40 years later throws into relief some of the differences between inspired, well-informed, natural history and modern science. He used no technical apparatus, made no measurements, conducted no experiments, and was able to make his deductions single-handedly because of his intimate familiarity with the animals concerned; the recent study, which is in no way denigrated, was carried out by seven collaborators from five institutions, who set out to test a specific hypothesis, used technical apparatus, and had recourse to statistical tests involving such things as regression coefficients - and confirmed what Armstrong had already concluded. The latter’s observational skills and insight were both impeccable. This does not mean that testing them was not necessary. It does, however, show that, in essentials, insight and the approach of the naturalist can sometimes be just as revealing as elaborate scientific tests. Thomas et al. (2002) revealed the interesting fact that small birds begin to sing at lower light intensities than large birds of equivalent eye size. Among possible explanations is that an early start to the day’s activities in small species may reduce the risk of night starvation. Small birds often lose 5-10% of their body weight overnight, and even more during long, cold, winter nights, though they are not usually singing much at this time, and require larger energy reserves, relative to body mass, than do larger birds. The need to devote the maximum amount of time to foraging in order to build up fat reserves to carry them through the following night is therefore particularly important in small species, and is helped by early rising. As a footnote to the timing of the dawn chorus it might be noted that, not surprisingly (though it has interesting physiological and evolutionary implications), nocturnal birds react to light intensity in the opposite way to diurnal species. As long ago as 1769, Gilbert White, writing to Thomas Pennant (in a letter included in The Natural History of Selborne) referred to the European Nightjar, Caprimulgus europaeus, as being “most punctual in beginning its song exactly at the close of day; so exactly that I have known it strike up more than once or twice just at the report of the Portsmouth evening gun, which we can hear when the weather is still”. Although the time at which the Europaean Nightjar begins its song in fact varies considerably according to the intensity of the light, beginning later on moonlight nights (Ashmore, 1935), this was a perceptive observation. Was it the earliest relating to the matter? Edward Armstrong (1900-1978) was a splendid example of the scientifically minded amateur naturalist, and indeed one of the long line of parson-naturalists. In 1942, while ministering in parishes in industrial Yorkshire, he produced a book on bird display, of which an enlarged edition appeared five years later under the title Bird Display and 141 The Dawn Chorus and how its Singers are Conducted Behaviour. This was sufficiently appreciated to be re-published with revisions and a new foreword as a Dover edition in 1965. This and his book on bird song were but two of his many contributions to ornithology. Although he reproduces his father’s portrait in a recent book on parson-naturalists, his son is modest about his achievements (Armstrong, 2000). Edward Armstrong was, however, a first rate naturalist and an outstanding student of birds, whose understanding of what determines the sequence of singers in the dawn chorus was but one example of his abilities. References Armstrong, E. A. (1963) A Study of Bird Song. Oxford University Press. London. Armstrong, P. (2000) The English Parson-Naturalist. Gracewing. Leominster. Ashmore, S. E. (1935) Time of singing of the Goatsucker. Brit. Birds 28: 259-260. Grist, W. R. (1935) Bird song at dawn. Naturalist 61: 168. Thomas, R. J., Szekely, T., Cuthill, I. C., Harper, D. G. C., Newson, S. E., Frayling, T. D., and Wallis, P. D. (2002) Eye size in birds and the timing of song at dawn. Proc. R. Soc. Lond. B. 269: 831-837. Y.N.U. BRYOLOGICAL SECTION: ANNUAL REPORT 2000-2001 T. L. BLOCKEEL 9 Ashfurlong Close, Dore, Sheffield S17 3NN AND J. M. BLACKBURN 6 By lands Grove, Fairfield, Stockton-on-Tees TS19 7BG Excursions The spring meeting in 2001, which had been planned for Upper Teesdale, had to be cancelled because of the outbreak of foot and mouth disease. The remaining sectional meetings were held as follows: Spring 2000: Roulston Scar and Caydale (VC62), 6 May (JMB) A party of five assembled at the White Horse car park on a fine sunny day. The morning and early afternoon were spent on Roulston Scar with the agreement of Forest Enterprise, and in the afternoon the upper end of Caydale was visited, also with the permission of the owner, to whom we extend our thanks. Roulston Scar. The purpose of the visit was to examine the steep slopes below the limestone cliffs. The area was visited by the British Bryological Society in 1967 and it was hoped to refind many of the species seen on that occasion. We were soon examining the rocks by the forestry track below the scar. Besides common species such as Bryum capillare and Mnium hornum, the calcicoles Ctenidium molluscum, Eurhynchium pumilum, Hypnum lacunosum var. lacunosum and Neckera complanata were present. The tiny moss Seligeria recurvata was growing in sizeable patches here with its distinctive curved seta. The slopes were accessed from the path which rises diagonally from the forestry track. The vegetation was well advanced and we were hampered in places by bramble thickets. The slopes had an excellent covering of bryophytes with large patches of Ctenidium molluscum, Dicranum majus and D. scoparium, Eurhynchium striatum, Thamnobryum alopecurum, Thuidium tamariscinum, and the three species of Rhytidiadelphus, R. loreus, R. squarrosus and R. triquetrus. Mnium stellare and Tortella tortuosa were also present. Naturalist 127 (2002) 142 Y.N. U. Bryological Section: Annual Report 2000-2001 Shaded limestone high up on the slopes produced the best finds. Tom Blocked spotted Leiocolea alpestris, a liverwort recorded in only two sites before in North East Yorkshire, in the Osmotherley and Kepwick areas in 1967. Close by Tritomaria quinquedentata was seen in small quantity on rocks, only the second sighting in the vice-county. Some time was spent comparing Neckera complanata with N. crispa. Eucladium verticillatum was growing on damp limestone and the liverworts Aneura pinguis, Apometzgeria pubescens, Frullania tamarisci, Porella platyphylla, Riccardia chamedryfolia and Scapania aspera were seen. The few elders on the slopes had Orthotrichum affine, O. diaphanum and Zygodon viridissimus var. viridissimus. The area should repay a further visit, perhaps in late winter before the vegetation has developed. Caydale. The afternoon was spent in the upper part of Caydale. This is also a limestone area with a stream flowing along the valley bottom fed from springs in Limperdale and King Spring Gills. In and by the stream three varieties of Paulstriella commutata (Cratoneuron commutatum) were seen, var. commutata, var. falcata and var. virescens, the last of these being only the fourth sighting in the vice-county. The marshy areas higher up had Plagiomnium datum, Bryam pseudotriquetrum and Drepanocladus cossonii. King Spring at the western end of the valley was very rewarding, the limestone rocks and tufa producing Bryam pallens, Didymodon fallax, Eucladium verticillatum, Leiocolea badensis and Preissia quadrata. This was a most rewarding day, with a total of 123 species recorded. Autumn 2000: Brockadale, Wentbridge (VC63), 14th October (TLB) Brockadale is one of the premier botanical sites in South-west Yorkshire, and a number of interesting bryophyte records were made here in the 1970s. The aim of the present visit was to assess the current status of the flora. The visit began from the Leys Lane car park. By the footbridge over the R. Went, a large waterside tree produced Syntrichia (Tortula) latifolia and Leskea polycarpa, two characteristic riparian species growing in the flood zone. Damp woodland nearby was interesting for its epiphytic species on Salix. These included plentiful Orthotrichum affine and O. diaphanum , and smaller amounts of Ulota bruchii and U. phyllantha. The Ulota species were unknown in this district in the 1970s, and the Orthotricha very rare. They have clearly recovered in recent years with improvements in air quality. Also noted as epiphytes were Dicranum tauricum and Bryum laevifilum (B. flaccidum). Some time was spent examining the limestone outcrops in the wooded parts of the valley. Among the species recorded were Jungermannia atrovirens, Plagiochila asplenioides, Fissidens gracilifolius, Tortella tortuosa, Trichostomum brachydontium, Gymnostomum calcareum, Tortula marginata, Mnium marginatum, M. stellare, Plagiomnium cuspidatum, Eurhynchium pumilum and Taxiphyllum wissgrillii. Several of these species are regionally rare, and the Gymnostomum is nationally scarce. On the north side of the valley, bare earth in the grassland near Brockadale Plantation produced some tiny ephemerals, including the delightful Microbryum rectum (Pottia recta) and M. davallianum (P. starckeana ssp. conica). The exposed south-facing crags in the Plantation are noted as a station for the tiny moss Gymnostomum viridulum, a southern species growing here near its northernmost limit in Europe. It was refound in rather small quantity on thin soil on rock slabs. Aloina abides was associated with it. Other species on the rocky slope included plentiful Trichostomum crispulum and some Pseudocrossidium revolutum (Barbula revoluta). 78 species were recorded within the YWT Reserve, and it was pleasing to find that many of the noteworthy bryophytes of the valley are still present. Although some of the previously known species were not seen, this is hardly surprising in such an extensive area and it would be premature to conclude that they have disappeared. Y.N. U. Bryological Section: Annual Report 2000-2001 143 Autumn 2001: Morton Wood and Reynard Clough, Hepworth (VC63), 13th October (TLB) The excursion to these sites near Hepworth were prompted by two notable discoveries made here recently by Mr R. A. (Bob) Finch. Morton Wood is a linear stretch of woodland along a stream bank on shale and grit. The flora is calcifuge, and the richest places were by the stream. Among the species growing in and near the water were Racomitrium aciculare, Brachythecium plumosum and Hyocomium armoricum. Small quantities of Heterocladium heteropterum and Scapania nemorea were present on one boulder. In places there are low cliffs of shale/grit, and here we found Pohlia annotina and Jungermannia pumila. Following Bob Finch’s directions, we were easily able to locate the two boulders which support Hygrobiella laxifolia, a very slender liverwort, growing here in a alga-like form on the moist surface of the grit rock. When found here in 1999, it was a new species for the vice-county. It is known from a few places in the High Peak of Derbyshire, but may be overlooked elsewhere in the South Pennines. We recorded in the wood until lunchtime, noting 43 species. Reynard Clough is situated on the moorland edge, 3 km south-west of Hepworth. It is bordered on both sides by conifer plantations, and in the absence of grazing the vegetation is becoming rather overgrown. Jungermannia sphaerocarpa was by the stream, and a tuft of Orthotrichum pulchellum was seen growing on a sallow bush. The north-facing sides of the clough are damp, and have cushions of Sphagnum rossowii and S. fimbriatum. There were several good patches of Calliergon stramineum mixed in with the grass and dwarf shrubs, and some Dicranum majus. An unexpected find in this habitat was a small patch of Rhytidiadelphus loreus, a very rare species in the South Pennines. Barbilophozia atlantica and Pogonatum urnigerum were noted on thin soil on an open shale bank. In the upper part of the clough, near the footbridge, we searched for Lepidozia pearsonii, the second of Bob Finch’s notable discoveries. We found several patches, on damp ground under heather and bilberry. L. pearsonii is a more slender plant than the common L. reptans, and it may be passed over as an etiolated form of that species. However it is distinguished by the presence of male inflorescences on lateral branches, and these were subsequently confirmed in the material that we found. L. pearsonii has a strongly ‘Atlantic’ distribution, and before its discovery at Reynard Clough it had not been seen in the South Pennines for more than a century. It is likely that it has been overlooked and that it will be found in further sites. We recorded 56 species in the clough and on the walls at its foot. Records There has been an increasing influx of records, especially from South Yorkshire, providing further evidence for the recovery of epiphytic species following reductions in the levels of SO2 pollution. Particularly notable in VC63 are the first known records for Orthotrichum lyellii, and the first for Cryphaea heteromalla since 1860. In VC64 Syntrichia papillosa has been refound after an interval of more than a century. During 2000 and 2001 all the known stations for Tortula cernua ( Desmatodon cernua, the flamingo moss) have been surveyed as part of Plantlife’s Back from the Brink project. This nationally rare moss is a specialist colonist of lime waste, and it has decreased from loss (and lack of renewal) of its habitat in quarries and near lime-kilns. Although it has gone from many of its old sites, there are seven surviving populations in VC63, and one just across the Derbyshire border in VC57. These are the only sites known to be extant in Britain. The list below includes new vice-county records and other records of note. An asterisk indicates a new or updated vice-county record. Recorders’ initials: TLB = T. L. Blocked; JE = J. Egan; HL = H. Lake. Aloina aloides: (61) 54/02 steep chalk bank, Hessle Chalk Pits, TLB, 10th November 2001. 144 Y.N. U. Bryological Section: Annual Report 2000-2001 Aloina ambigua: (63) 43/58 bare disturbed ground, Lindrick Common, TLB, 28th October 2000; 43/59 disturbed calcareous ground, old railtrack, west end of Conisbrough Viaduct, TLB, 3rd November 2000; (64) 44/43 stony soil, levelled ground on Magnesian Limestone, Peckfield Lane, Micklefield, TLB, 17th November 2000. Amphidium mougeotii: (63) 43/28 dripping rocks, streamside cliff, Rivelin valley, TLB, 9th December 2000. Brachythecium glareosum: (63) 43/58 old spoil heap, old Magnesian Limestone quarry, Lindrick Common, TLB, 3rd November 2000. Bryum algovicum var. rutheanum: (63) 43/58 low limestone wall, Roche Abbey, TLB, 19th May 2001. Bryum radiculosum: (61*) 54/02 mortar of brick wall, Hessle, TLB, 10th November 2001. Cephaloziella rubella: (63) 43/28 gritstone dough, Wyming Brook, HL, 11th February 2000. Conardia compacta (Amblystegium compactum): (63) 43/58 ledge on rock cutting, edge of Moses Seat Wood near Monk Bridge, Lindrick Common, TLB, 28th October 2000. Cryphaea heteromalla: (63*) 43/38 on elder by sheltered stream, Porter Brook, near Hanging Water, Sheffield TLB, 25th March 2001; 43/29 Underbank Reservoir, HL, 31st June 2001, 43/38 on willow, Shire Brook, Hackenthorpe, HL, 11th June 2001; 44/50 on Salix , bank of R. Don, near Sprotbrough Bridge, TLB, 24th November 2001; the only previous record of this pollution sensitive epiphyte in VC63 was made in 1860! (64) 44/36 on elder, long disused limestone quarry, Quarry Moor, Ripon, TLB, 12th October 2001; 44/26 on elder, stream bank, Fountains bridge, TLB, 24th July 2000. Didymodon acutus ( Barbula acuta): (64) 44/43 stony calcareous ground, quarry margin, Newthorpe Quarry, Micklefield, TLB, 17th November 2000. Distichium capillaceum: (63) 43/58 on low wall, Roche Abbey, TLB, 19th May 2001. Distichium inclinatum: (63*) 43/58 seasonally wet ground, old Magnesian Limestone quarry, Lindrick Common, TLB, 28th October 2000. Ditrichum flexicaule s. str. : (63*) 43/58 rock ledge, old shallow quarry, Lindrick Common, TLB, 28th October 2000. Fissidens adianthoides: (63) 43/58 about springs, old quarry on Magnesian Limestone, Brancliffe Lime Works, TLB, 18th November 2000. Frullania dilatata: (63) 43/38 Shire Brook, Hackenthorpe, HL, 11th June 2001; 43/38 tree bole, wooded stream banks, Limb Valley, HL & JE, 28th December 2001; 43/58 on ash, canal banks, Kiveton Park, TLB, 24th November 2001. Jungermannia hyalina: (63) 43/28 stone in wooded stream, Royd Clough, HL, 26th May 2000. Leiocolea alpestris: (63) 43/58 overgrown quarry face, woodland on Magnesian Limestone, Hawks Wood, TLB, 25th November 2000. Lophozia perssonii: (64) 44/43 stony soil, levelled ground on Magnesian Limestone, Peckfield Lane, Micklefield, TLB, 17th November 2000. Metzgeria fruticulosa: (63*) 43/38 tree bole, wooded stream banks. Limb Valley, HL & JE, 28th December 2001; (64) 44/26 on elder, stream bank, Fountains bridge, TLB, 24th July 2000. Y.N. U. Bryological Section: Annual Report 2000-2001 145 Microbryum curvicolle (Phascum curvicolle): (64) 44/43 stony soil, levelled ground on Magnesian Limestone, Peckfield Lane, Micklefield, TLB, 17th November 2000. Microbryum rectum (Pottia recta): (63) 43/59 disturbed calcareous ground, old railtrack, west end of Conisbrough Viaduct, TLB, 3rd November 2000; (64) 44/43 stony soil, levelled ground on Magnesian Limestone, Peckfield Lane, Micklefield, TLB, 17th November 2000. Orthotrichum lyellii: (63*) 43/29 tree branch 2-3 ft above water, reservoir margin, Broomhead Reservoir, HL, 31st January 2001; 43/29 on Salix, Underbank Reservoir, HL, 31st August 2001; 43/58 on ash, canal banks, Kiveton Park, TLB, 24th November 2001. Orthotrichum pulchellum: (63) 43/28 on Sambucus niger, stream valley on millstone grit, Wyming Brook, JE, 25th May 2001; (64) 44/26 on elder, stream bank. Fountains bridge, TLB, 24th July 2000. Orthotrichum stramineum: (64) 44/26 on elder, stream bank, Fountains bridge, TLB, 24th July 2000 Pohlia camptotrachela: (63) 43/29 Underbank Reservoir, HL, 31st August 2001; 43/29 Damflask Reservoir, HL, 5th September 2001; 43/29 Broomhead reservoir, HL, 11th September 2001; 43/29 Agden reservoir, HL, 17th October 2001. Pohlia drummondii: (63) 43/29 exposed mud, Damflask Reservoir, HL, 5th September 2001. Ptychomitrium polyphyllum: (63) 43/29 on wall. Midhope, HL, 9th November 2001; 43/29 on wall, Ewden Bridge, HL, 11th September 2001. Scapania aspera: (63) 43/58 rock ledge, old shallow quarry, Lindrick Common, TLB, 28th October 2000. Scapania nemorea: (63) 43/28 on grit boulders near stream, Black Brook, Rivelin valley, TLB, 9th December 2000. Schistidium platyphyllum (S. alpicola var. alpicola): (63) 43/29 reservoir overflow, Morehall Reservoir, HL, 4th October 2000. Syntrichia laevipila var. laevipila ( Tortula laevipila var. laevipila): (64) 44/36 on elder by disused magnesian limestone quarries, Burton Leonard, TLB, 12th October 2001; (64) 44/26 on elder, stream bank, Fountains bridge, TLB, 24th July 2000; var. laevipilaeformis: (63*) 43/59 on Salix, subsidence flash, R. Don, Cadeby, TLB, 22nd September 2000. Syntrichia papillosa (Tortula papillosa): (64*) 44/36 on elder by disused magnesian limestone quarries, Burton Leonard, TLB, 12th October 2001; last recorded in VC64 in 1895! Tortula cernua (Desmatodon cernuus): (63) 43/58 weathered lime waste, old quarry on Magnesian Limestone, Brancliffe Lime Works, TLB, 28th October 2000; 43/58 bare alkaline waste, old Magnesian Limestone quarry, Lindrick Common, TLB, 28th October 2000; 43/59 weathered lime waste, old quarry on Magnesian Limestone, Nearcliff, eastern end of Conisbrough Viaduct, TLB, 15th December 2001. Tortula lanceola (Pottia lanceolata): (63) 43/58 spoil heap, old quarry on Magnesian Limestone, Brancliffe Lime Works, TLB, 18th November 2000. Ulota phyllantha: (63) 43/38 Shire Brook, Hackenthorpe, HL, 11th June 2001; 43/58 dead Salix bark, South Anston, JE & HL, 25th April 2001; 43/28 on Sambucus, Black Brook, Rivelin valley, TLB, 9th December 2000. 146 Book Reviews Zygodon conoideus: (64) 44/26 on elder, stream bank. Fountains bridge, TLB, 24th July 2000. Acknowledgements T. L. Blocked gratefully acknowledges Plantlife, the wild plant conservation charity, who commissioned and supported the survey of Tortula cernua. BOOK REVIEWS The Origin of Plants by Maggie Campbell-Culver. Pp. 260, inch numerous colour plates. Headline, London. 2001. £25.00 hardback. This rather ambiguous title, clarified by the subtitle The People and Plants that have shaped Britain’s Garden History since the year 1000’, is an evocation in words and illustrations of plants introduced to this country. An anthology of garden history, it traces the provenance of many of the plants we take for granted; on a century-by-century basis, the author demonstrates, with the aid of contemporary paintings and modern photography, the enrichment of our botanical heritage. An introductory chapter touches on such subjects as early treatises on plants, the slow development of our indigenous plants for food and medicine, the rapid importation of plants through trade and exploration, and the creation of botanic gardens. Each chapter, including one entitled “Setting the Scene” which covers the period prior to 1000 AD, closes with a fascinating chronological list of plant introductions. Comparisons will undoubtedly be drawn with John Fisher’s The Origin of Garden Plants (Constable 1982), but that work lacks the attractiveness and more detailed content of Campbell-Culver’s beautifully illustrated book which is strongly recommended not only to botanists and gardeners but also to those interested in the history of the British landscape. VAH Lichens of Antarctica and South Georgia, A Guide to their Identification and Ecology by D. O. Ovstedal and R. I. Lewis Smith. Pp. xii + 411 (inch 48 b/w plates & line drawings), plus 32 pp. colour plates. Studies in Polar Research, Cambridge University Press. 2001. £70.00 hardback. This unique terrestrial environment, dominated by non-vascular plants, has been extensively researched in recent years, and although much has been published on its lichens, guides to their identification are limited. Useful monographs by Lamb on some genera and by Lindsay on the macrolichens of South Georgia require updating, and Dodge’s Lichen Flora of the Antarctic Continent and Adjacent Islands (1973) has been strongly criticized: although it covered 415 taxa, based on herbarium studies, more than half of these proved to be synonyms of species described from elsewhere. Redon’s Liquenes Antarcticos (1985), also based on herbarium studies, contains limited original material and is less widely available. The present work is therefore a most welcome identification guide covering 427 taxa (33.5% endemic to Antarctica), with keys to generic and specific level, the latter rather interestingly being easier to follow than the former. Taxonomic treatments, in alphabetical order, are sometimes accompanied by line drawings and black-and-white photographs, but the colour plates are of variable quality and do not do justice to the spectacular flora. Although large areas of Antarctica have yet to be lichenologically explored, and some locally important crustose genera (e.g. Lecanora, Lecidea s. lat., Rhizocarpon) require more detailed treatment, the authors are to be congratulated on compiling not only a useful identification guide but also a rich source of biogeographical, ecological and bibliographical material on this unique environment. It is an absolute must for the polar lichenologist, but at this price many users will only be able to consult library copies. MRDS 147 YORKSHIRE NATURALISTS’ UNION EXCURSIONS IN 2000 Compiled by A. HENDERSON and JANETTA LAMBERT BROUGH (VC61) 20 May (P. J. Cook) Introduction (G. Morrell) On the morning of the walk a little rain was forecast but the weather turned out sunny and quite warm. The group, numbering about 10, gathered at 10.30 a.m. as arranged at the Ferry Inn pub close to the River Humber. Before setting off, the different habitats to be found and the best routes were outlined, with emphasis on those preferential for various groups and individual interests. Just to the north of the village lies the southern tip of the Yorkshire Wolds with chalk grasslands and woodlands, but it was decided to take a circular 4-mile route around the southern side of the airfield towards the disused and flooded gravel workings which lie adjacent to the River Humber bank. Accordingly, members headed together towards the River Humber, at which point the majority turned east along the bank whilst the coleopterists went west to search the brackish- water creeks. Soon each specialist was finding something of interest. A clutch of Mallard eggs was found on the end of a washed-up plank in the reed-bed, and, over the next few weeks, the author of this note, a local naturalist, was able to keep an eye on them, being rewarded by being there when the eggs hatched. As members moved further eastward, all sorts of flora and fauna were discovered: Annual Wall Rocket, Fallow Deer on Reeds Island, rare snails and insects. Lunch was enjoyed sitting on the bank almost opposite Reeds Island, from where the Fallow Deer could be watched through a telescope. Afterwards members set off back along the track between the disused gravel workings before turning west along the northern side of the airfield. The route back to the Ferry Inn pub and the tea meeting was through the village and alongside the railway line, the wall colonised by a species of polypody. After the welcome chips and sandwiches laid on at the pub, the day’s foray closed with reports and discussion of the activities and finds of each section, with a number of the specimens collected during the day being passed around. Mammals (C. A. Howes) The Humber bank is an extremely popular dog-walk, the huge daily contributions of dog excrement providing a food substrate for coprophagous diptera, coleoptera and fungi, perhaps rivalling the estuary strand-line as a major ecological influence. Grey squirrels were present in the trees and gardens of urban Brough. Mole hills were frequent in lawns and agricultural land. Drain banks and hedge bottoms held Rabbit warrens and Common Rat colonies. Bold and obtrusive Common Rats were much in evidence feeding on picnic scraps in the Brough Haven Car Park. Good views were enjoyed of Brown Hare sprinting across flooded grazing-land. Roe Deer secreted by day in copses were seen at a distance but were regular quarry to local shooters. Watercress beds in a briskly running chalk stream produced Water Shrew; Water Voles were evidenced by their droppings and the grazed lawns they had left behind them; a sluggish ditch adjacent to the aircraft factory held Three-spined Sticklebacks; and Common Frogs were in wet grassland. Anglers at the local flooded clay pits were catching Bream, Chub, Eel, Flounder, Perch, Roach and Tench. Ornithology (W. F. Curtis) The ornithological members of this meeting concentrated their efforts along the Humber Estuary from the car park immediately to the west of the British Aerospace factory eastwards to Welton Water. The first stretch consisted of the raised river bank with a Naturalist 127 (2002) 148 Yorkshire Naturalists’ Union Excursions in 2000 narrow fringe of reeds where Reed Bunting, Reed Warbler and Sedge Warbler were in evidence along with three species of finch. Small numbers of Shelduck were noted offshore, whilst a lone Grey Heron fished along the edge of the reeds. To the east of the factory the short-cut grass of the airfield attracted several corvid species together with Pied Wagtail, Yellow Wagtail and Lapwing. A nearby, small flooded area revealed a single pair of Moorhen with young. Towards the eastern end of the airfield a large (c.4 hectares) flooded area caused, according to a local resident, by the collapse of the field drains, held a veritable wealth of wildfowl. Amongst those recorded were three pairs of Great Crested Grebes, two with young; some four Little Grebes; several each of Garganey; Gadwall; Ruddy Duck; Pochard; Tufted Duck; Shoveler; Mallard; Grey Lag Geese, with goslings; Canada Geese; Mute Swan, with cygnets; and at least five species of waders including Black-tailed Godwit and Ruff. Arguably the creme de la creme was a Little Egret, still not a common bird in Yorkshire, though the Black Tern seen on the return walk must be a contender. The area of bushes between the eastern boundary of the airfield and Welton Water produced a good array of passerines with five species of warbler plus Goldcrest, Long- tailed Tit and Song Thrush. In this same general area, Stock Dove and Kingfisher were observed whilst a Cuckoo was heard. Raptors reported were Sparrowhawk and Kestrel (and an Osprey recorded by a visiting bird-watcher). On returning to the car park some 25 Ringed Plovers were feeding on the mud exposed by the falling tide, together with a few Dunlin, Curlew and Redshank. A total of 72 species was recorded. CONCHOLOGY (A. NORRIS) The molluscan records produced for the Brough area by John W. Taylor, W. Denison Roebuck, J. E. Crowther and William Cash on the occasion of the 1901 YNU visit to the area number 33 species. Most of these, however, came from the chalk wolds north of Brough. On this occasion we concentrated on the Humber foreshore, in particular the brackish pools, ditches and salt-marsh. The area only produced 20 species on the day, however, 10 of which do not appear in the 1901 list. It is interesting to note that, although the salt-marsh species would have been known to the 1901 party, they either did not look for them or they did not occur in that area at that time. The Dun Sentinel, Assiminea gray ana and its associated species Ovatella ( Myosotella ) myosotis were both recorded from the River Humber at a later date. These two species were not known when Tom Petch produced his report, “The published records of the land and freshwater mollusca of the East Riding, with additions” ( Transactions of the Hull Scientific and Field Naturalists’ Club [1904]. A. Smith of York first recorded Assiminea gray ana in the salt-marsh at Hessle on June 28 1950 ( Journal of Conchology 23(5): 134), where it was subsequently recorded as fairly common (A. Norris; March 28 1965 [TA54/005255]). A further search of the Humber bank produced a small colony on the salt-flats behind King George Dock, Marfleet, Hull on April 8 1966 (TA54/153285); no further colonies could be found. The original site at Hessle was destroyed in 1998-99 to safeguard the railway line that runs close to the shore at that point. A search on 10 April 1999, of the area just east of Hessle (inaccessible in the 1960s as it was situated within the restricted area of the dock railway yards) produced a thriving colony of Assiminea and a few examples of Ovatella. Therefore when the opportunity arose to examine the salt-marsh east of Brough to see if these snails had extended their distribution further upriver, it was delightful to find that both species occurred, Assiminea in all three of the 1 km squares examined (44/9424; 9425 and 9326), and a single example of Ovatella in 9425. It would seem that these species are expanding up-river on the north bank of the River Humber. It would be interesting to know if the previously predicted changes in the tides within the Humber have made these habitats available, or if they have always been there in small numbers. It is hard to understand how the early collectors could have missed them. Ovatella ( Myosotella ) myosotis is far more common and widespread, with a marine Yorkshire Naturalists’ Union Excursions in 2000 149 variety occurring on the east coast. The main find on the landward side of the area was the first record for VC61 of the slug Arion (Mesarion) flagellus, which was found in the field being used as a landfill site. Lepidoptera (G. Boyd) Butterflies and moths were not particularly abundant either in numbers of individuals or of species observed. Just six species of butterfly were seen, with the Green-veined White (' Pieris napi ), the commonest, followed by the Orange Tip Anthocharis cardamines. Three species of macro-moths and four species of micros were reported. Perhaps the most interesting record was a newly emerged specimen of the pyralid Myelois cribrella (Thistle Ermine) found by Miss J. Lambert. This used to be considered a south of England species, barely reaching Yorkshire, but in the last 20 years it has been recorded regularly from the county and indeed from as far north as Scotland. Whether this extension in range is real or apparent is not clear. Cleaner air or generally warmer weather could have induced the moth to spread northwards, but it may have been here all the time, overlooked because most lepidopterists have concentrated their attention on the larger moths. Hymenoptera (M. E. Archer) The species of aculeate wasps and bees recorded at Brough were graded as Common, Frequent, Occasional, or Rare, for Yorkshire. These comprised five species of Bumble Bees, all Common: Bombus lapidarius, B. pascuorum, B. hortorum, B. pratorum, and Psithyrus vestalis\ and four species of Solitary Bees: Andrena scotica, A. haemorrhoa, Nomada marshamella , all Common; and Halictus tumulorum. Frequent. Diptera and Miscellaneous Orders (G. Boyd) To invite a second-string lepidopterist to collect and collate field records for flies is rather like entering a cart-horse for the Grand National. Since it may well be another century before the Union visits Brough, the meeting co-ordinator decided that any records at all for this group would be preferable to complete silence. Perhaps she thought that, having recently retired from work, the recorder should be encouraged to do a little proper entomology rather than merely matching insects to pictures as many moth and butterfly men are alleged to do ! In the end the total list for the day (lodged with Miss Lambert) ran to 36 species from within seven insect orders. These included the damsel fly Ischnura elegans, the red and black leaf hopper Cercopis vulnerata, the common earwig and fifteen assorted species of fly. The two most striking specimens, circulated as members of the party enjoyed their sandwiches on our return from the field, were the lovely picture-winged fly Euleia heraclei and the large snipe fly Rhagio scolopaceus. (The latter was erroneously introduced to members as a vicious biting Tabanid; later determination proved it to be a perfectly innocuous beast, although in a closely related family.) The diptera collected were almost all of species both widespread and common, the only exceptions being the hover-flies Platycheirus fulviventris and P. immarginatus . These are said to frequent, respectively, “lush open marsh” and “brackish marsh on the coast or along tidal reaches of rivers” (Stubbs & Falk, 1983). The habitat at Brough is clearly appropriate for both species. The species are superficially extremely similar and it was only after the two male specimens had been pinned and set so that the front legs could be examined under magnification that it was realised that they were not duplicates - which was particularly pleasing, neither having been previously taken. Plant Galls (J. & K. G. Payne) Mr A. Henderson found the gall-causing aecial stage of Puccinia phragmitis on Rumex crispus and of a Phragmidium (almost certainly mucronatum) on Rosa sp. Miss J. Lambert collected the terminal bud gall of the cecidomyid Rhabdophaga rosaria on Salix sp. 150 Yorkshire Naturalists’ Union Excursions in 2000 (Willow bushes by the Humber bank path at Brough). The galls listed below were all found by Mrs J. Payne in the area of the Humber Bridge car parks (54/0126). Most interesting was an abundance of sloes distorted by the fungus Taphrina pruni: hardly a single normal fruit was to be found. The whiteness of the distorted fruit was obvious at c.100 metres distant. Besides this there were Puccinia punctiformis on field thistle (plants spindly and pale green); P. malvacearum on a mallow (distorted leaf veins); Podosphaera clandestina on young hawthorn shoots; Phytoptus avellanae on hazel (swollen, mite-filled buds); and Aceria stenaspis on beech (narrow rolling of leaf edge). Arachnida (C. A. Howes) The windswept beds of Phragmites and Agropyron along the Humber shoreline produced only small numbers of Clubiona phragmitis, Pachygnatha clercki and Bathyphantes concolor. However, Lycosid (wolf) spiders were abundant, stalking insects settling on the Humber mud and strand-line flotsam. These included Pardosa hortensis and pullata on the higher shore, whereas the less vegetated mud subject to regular inundations produced the salt-marsh specialists Pardosa purbeckensis and Xerolycosa miniata. Tussocky vegetation in terrestrial situations in the lea of the flood embankment held good numbers of Tetragnatha extensa and Larinioides cornutus. Botany (R. Middleton) Although the meeting in 1900 appears to have ranged further afield and into the Wolds valleys, it was decided that a perambulation of Brough Airfield would be more appropriate and would still sample several habitats. The waste places of Brough supported the usual mix of urban weeds although the members attending the 1900 meeting would have been unlikely to encounter either Senecio squalidus or Lepidium draba. At Brough Haven the upper intertidal mud was dominated by a stand of Bolboschoenus maritimus. Along the upper shore to the east a good selection of halophytic species was found, including Glaux maritima, Apium graveolens, Plantago maritima and Aster tripolium. An unusual Scurvygrass was collected and later examined by Mr P. Cook who was of the opinion that it was a Cochlearia x hollandica backcross, resulting from hybridisation between C. anglica and C. officinalis. Although full access to the extensive grassland of the airfield was not available, examination of its margins did reveal some of the plants that might have been expected - Stellaria graminea, Lathyrus pratensis, Centaurea nigra, Carex hirta and Cynosurus cristatus. Limited examination of the main body of grass, via the public footpath which bisects the field, proved rather disappointing, revealing that it had been much ‘improved’ and that the margins were actually a relic of an earlier state. To the north of the airfield the light sandy soils provided a fine display of arable weed species including Spergula arvensis, Viola arvensis, Anchusa arvensis and Amsinckia micrantha. The latter taxon, although now a well established, widespread and often abundant element of the local cornfield flora, did not arrive in Britain until the early years of the 19th century or the vice county until the 1960s. The final surprise of the afternoon was a superb display of the locally infrequent fern Asplenium ruta-mutaria, which had comprehensively colonised a south facing wall adjacent to Brough railway station. Mycology (J. Payne) The following non-galling fungi were collected by Mrs J. Payne: Puccinia cnici on Cirsium vulgare; Entyloma ficariae on Ranunculus ficaria and Peronospera niessleana on Alliaria petiolata. Lichenology (A. Henderson) The riverside walk along the airfield’s southern boundary produced no lichenological Yorkshire Naturalists’ Union Excursions in 2000 151 surprises: some common Cladonia species with many squamulose patches; and a few crustose species on pebbles and stone banking, the most interesting being Thelidium minutulum with its very tiny immersed spherical pyrenocarps on harder calcareous substrates. Xanthorion, best seen here on the wooden pier, was to be a feature of the day’s flora on fencing and in hedgerows. As we turned northwards on the track to Melton Common, Rosa branches were quite yellowed by invading Xanthoria parietina, X. polycarpa and X. Candelaria on a background of grey Physcia and Amandinea punctata. Hedgerows and copse along the path east of the airfield had the Xanthorion- influenced Parmelietum now typical of such agricultural, suburban-outskirt areas, with occasional Usnea subfloridana and Ramalina farinacea. As the trackway to the north of the airfield crossed an old airfield path of concrete and asphalt, Caloplaca crenulatella was locally common. This species, only recognised as a distinct entity in this country in the last few years, appears to have one of its headquarters on disused airfield runways and tracks throughout Britain, often covering many square metres of runway. Moss-covered patches of such worn asphalt surfacing had Collema tenax var. ceranoides nestling almost hidden among spreading Hypnum cupressiforme. All in all, a day spent renewing acquaintance with lichens typical of our increasingly ‘ suburbanised ’ landscape. OLDSTEAD (VC62) 10 June (J. M. Blackburn) Introduction (J. M. Blackburn) It was on a fine warm day that 28 members assembled on the grass verge of the track leading to Cockerdale Farm, Oldstead. Forest Enterprise had given permission to visit Cockerdale and Great Cockerdale Wood, whilst the fields surrounding Cockerdale Farm were all open access. This meeting did not replicate a Union meeting in 1902, but it was a day for making comparisons. In 1902 the Union met at Coxwold station and walked from there on a day which provided one or two heavy showers. They reached Oldstead from a lane by Kilburn Thicket where the party divided into two groups, one party going up to Scotch Corner and Roulston Scar and the other up Oldstead Bank. The two groups merged again near Kilburn before returning to Coxwold. Our own party arrived at the start by car, of course, and Roulston Scar was not on the itinerary for the day. The presence of pine and larch plantations is mentioned in the 1902 report but the present coniferous cover is extensive. This has inevitably affected the character of the area, creating acidic habitats in a predominantly limestone area. Fields opposite the parking area were immediately visited. They were herb-rich and, in the marshy areas at the southern end, Lychnis flos-cuculi was in evidence, so good to see in a habitat much declining in the county and very much in need of protection. Oldstead Bank received due attention, with Daphne laureola and Helleborus foetidus noted, as they were in 1902. The central area of Great Cockerdale Wood was accessed by a permissive path leading to Mount Snever Observatory. This was visited by several members whilst others reached the observatory at the top of the scar. There was much bramble in the wood above the conifer level but progress was not unduly hampered and the initially acidic habitat, with birch, sycamore and oak, soon gave way to a calcareous zone, dominated by ash. Here the slopes were strewn with limestone boulders which had fallen from the limestone crag above. This area quickly repaid the attention it received. The marshy area and stream in the valley bottom was well covered, with mixed results. Some members did a circular walk by walking up the forest track round to Cocker Dale and crossing the fields to reach the top of Oldstead Bank and Scotch Corner. Fifteen affiliated societies were represented by 23 members at the indoor meeting. The landlord of the Black Swan in Oldstead had kindly agreed to open for us and drinks and sandwiches were soon circulating. This was followed by a very lively meeting chaired by our President, Colin Howes. Apologies were received from Heather Walker, Alan Heaton and also from Andrew Grayson who had been unable to stay for the meeting. All sections 152 Yorkshire Naturalists’ Union Excursions in 2000 had much to report and the chairman added to these by making comparisons with findings and remarks from reports on the visits to the area in 1902 and an earlier visit in 1893. Les Magee said that he had found no sign here of the disease affecting alders in some parts of the country. The trampling in the flushes and marshy areas was commented upon and duly noted. The chairman expressed his thanks to the Excursion Secretary, Divisional Secretary and the landlord of the Black Swan. Reports on our findings will be sent to Forest Enterprise and to Colin Furness of Cockerdale Farm, to whom we also extend our thanks. Mammals and Lower Vertebrates (M. J. A. Thompson) Compared to the Y.N.U. outing to Oldstead in 1902, the nine mammal species seen on the 2000 outing were similar in number, but there were species differences. Of the Insectivora on this occasion, beside numerous mole hills, a dead mole Talpa europaea was found on one of the woodland rides, whereas in 1902 a common shrew Sorex araneus was also recorded. Both rabbit Oryctolagus cuniculus and brown hare Lepus europaeus were seen, the latter being flushed near Cam Farm. Of interest, in 1902 the red squirrel Sciurus vulgaris was the resident squirrel, only to be replaced by the American grey squirrel Sciurus carolinensis a century later. The only other rodent record was the field vole Microtus agrestis, evidenced by numerous field vole runs and holes underneath some of the thick matted grass edging some of the meadows. Carnivora were represented by the badger Meles meles, with an active badger sett in one of the woods, along with numerous badger trails and footprints. A fresh fox Vulpes vulpes scat was found, but its contents did not indicate what other small rodents were in the area. No deer were recorded in 1902, but in 2000 both roe Capreolus capreolus and fallow Dama dama were present. A single roe deer was seen in one of the dense coniferous woods and there were both roe and fallow deer slots in the soft mud of the rides, with the larger fallow prints. The roe deer population explosion in this part of North Yorkshire occurred in the 1930s. A small number of fallow are known to be in the area. At least 3 different individual common or viviparous lizards Lacerta viviparus were seen sunning themselves on the small rocky limestone outcrops overlooking the study area. One of them was a pregnant female. Ornithology (J. E. Dale) The late morning was spent following the main track through Great Cockerdale Wood to the ridge near Cam Farm, and on the return following the edge of Cockerdale Wood before taking a steep descent past the farm to return to our car park. Goldcrests and Coal Tits were abundant in the trackside conifers; Willow Warblers were in song in all parts of the area; Blackcaps were less common but at least six were heard during the morning and Chiffchaffs were located twice. In the relatively open area of scattered bushes at the head of Cockerdale a few Whitethroats, and at least one Garden Warbler were heard. Kestrel was hunting here and Spotted Flycatcher (two) and Redstart were near the Cockerdale Wood track. Curlew and Skylark were heard over farmland above the woods. Part of the afternoon was spent on Modes Bank, and later the track was followed down the valley towards Oldstead. Other species added to the list were Swallow (five), House Martin, Song Thrush, Long-tailed Tit and Goldfinch. Cuckoo and Linnet were reported by others during the tea-time discussions. 38 species compared favourably with the 45 recorded during the 1902 visit when rather more time was available. Noteworthy differences in 2000 were the lack of Wood Warbler which had been ‘heard on all sides’, presumably in natural hanging woods that in this area have probably been replaced by conifers, and the inevitable absence of Corncrake. Hooded Crows were present on a keeper’s gibbet in the earlier visit, and although they had doubtless been shot before the spring months, are today quite rare in Yorkshire, and most likely to be seen near the coast. Goldcrest was very numerous during our visit whereas it was apparently treated as an interesting find in 1902. Yorkshire Naturalists’ Union Excursions in 2000 153 CONCHOLOGY (A. NORRIS) The field meeting reports for 1892 and 1902 record an interesting mollusca fauna for the area of Roulston Scar, Coxwold. In 1892 W. Denison Roebuck and F. W. Fierke recorded 38 species, and at the 1902 meeting the Rev. T. Ainsworth and J. E. Crowther listed 29 species, 8 of which had not been recorded in 1892. These two meetings produced a total list of 46 species, 8 of which are freshwater species. David Lindley and myself recorded 48 species, only 3 of which are freshwater species. The 1892 report states that a pond not far from the village of Wass produced the freshwater species but there is no specific report covering these in the 1902 report. It can be seen from these totals that we recorded several species not seen on the previous meetings. Two of these are the freshwater snails Potamopyrgus antipodarum and Lymnaea truncatula, other additions being Succinea putris, Columella edentula, C. aspera, Vertigo substriata, Arion subfuscus, Nesovitrea hammonis, Oxychilus draparnaudi, Monacha cantiana, Zenobiella subrufescens, and the segregate species Carychium tridentatum, Cochlicopa lubricella, Vitrea contracta and Euconulus alderi. 6 of these would have been unknown to the early recorders. It is also interesting to note that 8 of the above are mainly recorded from marshes. One of the highlights of the trip on both of the previous meetings was Helicigona lapicida. In the report for 1892 this species was reported as being found in a deep ravine above Wass village. The report for 1 902 records dead specimens from Roulston Scar, and the circular for that meeting (the 164th meeting), reports that two dead specimens had been recorded from a hedge bank between Kilburn Village and the Scar. David Lindley also recorded two dead specimens from the cliffs of Roulston Scar on this occasion. A full survey of the area for habitats of lapicida , a species not recorded alive from this part of Yorkshire in over 100 years would be an interesting project for the future. Other highlights recorded in the past and repeated on this occasion included very fine examples of the slug Umax cinereoniger and the very local Azeca goodalli (A. tridens in early reports), found in a marsh with Vertigo substriata and the two Columella species. Lepidoptera (J. Payne) Owing to a late start and limiting circumstances the writer mainly recorded on the lower ground. However with the observations of L. and R. Aukland, P. Tannett, G. Wadsworth and others a very good list of butterflies and moths was compiled. The only member of the Hesperidae seen was Small Skipper. The Pieridae were much in evidence - Large, Small and Green-veined were in fair numbers. 3 male Orange-tip were seen and, even more pleasing, 3 Brimstone were reported. Two members of the Lycaenidae Common Blue and Small Copper were recorded. It was noticeable that in the group Nymphalidae there was no sighting of Small Tortoiseshell. This species was scarce in 1999 and seems destined to be so again. Very pleasing was the report of 3 Comma and a single Painted Lady. Also reported was a single Peacock and a worn Red Admiral. The Satyridae showed up as Small Heath and a single Wall Brown making a magnificent total of 14 species. Most of the moths recorded were members of the Geometridae: Yellow Shell, Clouded Border, Chimney Sweeper and Streamer were noted and there was an abundance of Silver- ground Carpet (a Galium feeder) no doubt because Goosegrass is flourishing again in the hedgerows. The two noctuids recorded were the aptly named Beautiful Golden Y and the equally beautiful Angleshades in predominantly pink colouration. Entomology (W. R. Dolling) Orthoptera: It was too early in the season for any of the true grasshoppers but an adult of the Common Groundhopper Tetrix undulata was swept in a ride. Dermaptera: The Common Earwig Fortificula auricularia was everywhere. 154 Yorkshire Naturalists ’ Union Excursions in 2000 Mecoptera: Scorpionflies, Panorpa species, were numerous but none were identified though P. communis was the most likely. Neuroptera (s.l.): A single male snakefly Raphidia xanthostigma was swept in the valley bottom where we parked the vehicles; this species is said to be associated solely with willows. Diptera: The only fly identified was the Rhagionid Chrysopilus cristatus in the valley- bottom flush. Hemiptera-Homoptera: The black and red froghopper Cercopis vulneraria was noticed in several places. Leafhoppers (Cicadellidae) noted were: Eupteryx signatipennis on Meadowsweet and Oncopsis avellanae on Hazel. Planthoppers (Delphacidae): Criomorphus albomarginatus, Stiroma ajfinis and Ditropis pteridis, the last monophagous on Bracken. Cixius nervosus (Cixiidae) lcf in the flush. Hemiptera-Heteroptera: Hawthorn Shieldbug Acanthosoma haemorrhoidale, one dead under conifer bark. The lygaeids Cymus melanocephalus on rushes in the flush and Kleidocerys resedae on Alder and Birch. Anthocoris nemorum was the only anthocorid. The remainder of the true bugs were Miridae, as follows: the grass-feeding Stenodema holsatum, S. laevigatum and S. calcaratum, all as overwintered adults, and nymphs of Leptopterna dolabrata and Capsus ater, grass-feeders that overwinter in the egg; adults of the new generation of three oak-associated species were already present: Harpocera thoracica, Dryophilocoris flavoquadrimaculatus and Calocoris ( Closterotomus ) quadripustulatus; Monalocoris filicis on ferns; Macrolophus pygmaeus (= nubilus) and Dicyphus stachydis on Woundwort; and Calocoris alpestris (= major ) and Lygus wagneri on low plants in shady places. Coleoptera: Collecting methods being confined almost wholly to working plants rather than sieving, grubbing etc., the list has few specimens in common with the list from a century ago which features mainly species from cryptic habitats. The widespread growing of Oilseed Rape in the vicinity was reflected in the presence of the Pollen Beetle Meligethes aeneus (the ‘Rape Fly’) and the weevils Ceutorhynchus assimilis, C. floralis and C. pallidactylus. The flea-beetles Longitarsus parvulus and Aphthona euphorbiae suggest that Linseed is also grown in the vicinity. Undoubtedly the best aspect of the day’s work was the finding of Oedemera virescens (Oedemeridae) in some numbers, coleopterists (Frank Kenington and W.R.D.) encountering several specimens. This species has the conservation rating ‘vulnerable’ (RDB2); although previously found also in Gloucestershire and Norfolk, since 1970 it has only been taken in NE Yorkshire. Pollen beetles (in addition to the ‘Rape Fly’) were: Meligethes brunneus (on Woundwort), M. morosus (on White Dead-nettle), M. atratus (on Dog Rose), M. carinulatus (on Birdsfoot Trefoil) and Brachypterus urticae (on Nettle). Weevils were numerous with the following identified: Deporaus betulae (both adults and the characteristic leaf-rolls on Hazel, in which the leaf lamina is almost severed transversely, the dry, brown terminal portion hanging in a conical roll from the midrib of the intact, green, basal portion); Apion ulicis (on Gorse); A. apricans (on Red Clover); A. ervi (on Vetches); A. loti (on Birdsfoot Trefoil); A. violaceum (on Docks); A. curtirostre (on Sheep’s Sorrel); Anthonomus pedicularius (on Hawthorn); Curculio salicivorus (on Willow, where it is an inquiline in sawfly galls); Barypeithes pellucidus (in leaf litter); Strophosoma melanogrammun, Phyllobius glaucus, Polydrusus pterygomalis (all mostly on Hazel); Phyllobius pomaceus (= urticae) (on Nettle) and Polydrusus pilosus (on Spruce). Leaf- beetles included Oulema gallaeciana and O. ? melanopa (on grasses); Gastrophysa polygoni (on Dock); Chrysomela aenea (on Alder); Phratora (= Phyllodecta ) laticollis (by sweeping, but supposedly associated with Aspen and White Poplar although here probably on Salix)\ Galerucella lineola (on Salix ): G. tenella (on Meadowsweet); Phaedon Yorkshire Naturalists’ Union Excursions in 2000 155 cochleatriae (probably on Watercress); Phytodecta pallida (both the pale brown adults and the leaf-green larvae on the leaves of Hazel). Flea-beetles: Altica lythri (on Willowherb); Chalcoides fulvicornis (on Willow); Longitarsus suturellus (on Ragwort); Phyllotreta undulata (on crucifers); and the two linseed-associated species mentioned above. Ladybirds: Calvia 14-guttata, Propylea 14-punctata, Coccinella septempunctata. Click- beetles: Aplotarsus (or Selatosomus) incanus, Agriotes pallidus, Denticollis linearis. Rove- beetles: Tachyporus nitidulus, Tachinus laticollis, Anthophagus caraboides (the last on Rowan flowers). Longhorn (cerambycid) beetles: only Rhagium bifasciatum (also at Rowan flowers, but its larva bores in coniferous trees; it was found on one of the early excursions to the Coxwold area). Scraptiidae: Anaspis rufilabris, A. frontalis and A. maculata. Other odds and ends: Cyphon coarctatus (Elodidae), Aphodius ater (Scarabaeidae), Atomaria linearis (Cryptophagidae), Cortinicara gibbosa (Latridiidae), Rhinosimus planirostris (Salpingidae), Byturus tomentosus (Byturidae), Malachius bipustulatus (Melyridae), Cantharis nigra (Cantharidae - several other species present were uncollected) Eupuraea melanocephala (Nitidulidae, but not a pollen beetle like Meligethes ) and two additional unidentified species of Eupuraea. Plant Galls (J. & K. G. Payne) As at our recent Brough meeting, sloes distorted by the fungus Taphrina pruni were much in evidence. Fine specimens, in fact, occurred exactly at the spot identified for members to meet. Miss Janetta Lambert collected the ‘Knotting Gall’ caused by the fly Chirosia betuleti on the Broad Buckler-fern and Field Maple leaves covered by the red pustules of the gall mite Artachris cephalonea. Growing right by the door of the Black Swan, where we had tea and the meeting, was Ground Elder with the swellings caused by the fungus Protomyces macrosporus. Most members will have noticed the dense covering of pustules on Alder leaves caused by the mite Phytoptus laevis laevis\ also on Alder leaves were the swellings in the angles between midrib and veins caused by another mite, Eriophyes inangulis. In addition, J.P. noted the following: Red Campion with ‘smut’, Ustilago violacea on the anthers; Blackthorn with the leaf-edge galling of the mite Phytoptus similis\ Dog Rose with the downwardly rolled leaf-edges of the sawfly Blennocampa phyllocolpa\ Germander Speedwell with the hairy terminal pouch of the gall midge Jaapiella veronicae and Crosswort with the galling rust Puccinia galii-verni. Botany (N. Sykes) Although arable and pasture fields in the vicinity had been treated for maximum agricultural production, and ancient woodland on the surrounding hills had been largely re- planted with conifers, the area still retained pockets of near natural vegetation with interesting plant communities. Seepage from hillside springs drained into a marshy area with Juncus ejfusus, J. inflexus. Lychnis flos-cuculi, Angelica sylvestris, Valeriana dioica, Scrophularia auriculata, Lotus pedunculatus, Stellaria uliginosa, Cardamine pratensis and Menyanthes trifoliata. A nearby shallow streamside had Chrysosplenium alternifolium, C. oppositifolium , Cardamine amara, C. flexuosa, Calamagrostis epigejos, Veronica beccabunga, Callitriche sp. and several overhanging trees of Salix pentandra revealed by strong fragrance and shining leaves. On drier patches were found numerous Dactylorhiza fuchsii, Stellaria holostea , Rumex acetosella , Potentilla anserine , Briza media, Lathyrus pratensis, Alchemilla xanthochlora. Geranium robertianum, Myosotis discolor. Geranium molle, G. dissectum, Trifolium pratense T. repens, Ranunculus bulbosus, R. repens and R. acris. Medicago lupulina and Veronica chamaedrys, V. officinalis, V. serpyllifolia and V. montana were scattered around. A forest ride adjacent to Cockerdale beck was largely shaded by near conifers which doubtless served to check more vigorous plants as the trackside was carpeted with Lysimachia nemorum and Ajuga reptans) in flower. Valeriana officinalis, Scrophularia nodosa , Anthriscus sylvestris, Arctium sp., Tamus communis, Geum urbanum and 156 Yorkshire Naturalists’ Union Excursions in 2000 Vaccinium myrtillus also occurred. A steep climb through the wood revealed limited ground flora but on level less shaded ground beyond the Observatory Melampyrum pratense , Mercurialis perennis, Galium saxatile, Silene dioica and Ulex europaeus eventually gave way to more acid ground dominated by bracken amongst which numerous Trientalis europaea were seen in full flower. A grassy track climbing north-east up Modes Bank has long been known for its Helleborus foetidus which were seen in fruit. As well as the more common hedgerow plants were clumps of Daphne laureola also in fruit and large stands of Campanula latifolia not yet in flower. Bryology (J. M. Blackburn) The main thrust of the day was to spend time in the central area of Great Cockerdale Wood which seemed of great potential interest when the area was selected for this meeting. The interest was in the boulder-strewn slopes under the limestone crag. The limestone rocks here, where it is shaded and humid, are covered in mosses, with large quantities of Ctenidium molluscum, Thuidium tamariscinum and Tortella tortuosa. Some of these rocks had good-sized patches of the tiny moss Seligeria recurvata, proving to be frequent in the southern part of the National Park. Both Neckera crispa and N. complanata were present, with Anomodon viticulosa, Mnium stellare and Campylium calcareum in its only extant site in the vice county. Eucladium verticillatum was on some of the damper rocks. Campylopus fragilis was seen on a sloping bank in the wood, seen very infrequently in the vice county. The attractive thalloid liverwort Apometzgeria pubescens was present in several places. Three species of Scapania were found on the rocks, S. irrigua, S. nemorea and S. umbrosa, along with Frullania tamarisci and Porella platyphylla. In the more acidic parts of the wood Mnium hornum was present in large quantities and also Atrichum undulatum and Plagiomnium undulatum. The marshy area by the stream was inspected but was not particularly rewarding. Elders had Orthotrichum affine and O. diaphanum, with cracks in the bark supporting Zygodon viridissimus var. viridissimus. A marshy area above Cockerdale Farm had a fine stand of Climacium dendroides. The acidic banks by the forestry track were rewarding, as expected, with all three of the common species of Calypogeia, C. arguta, C. fissa and C. muelleriana, Cephalozia bicuspidata, Diplophyllum albicans and Lophozia ventricosa. This is proving to be one of the richest areas in northeast Yorkshire, due to the mix of habitats present. A total of 124 species was recorded. Lichenology (A. Henderson) Woodland to the north and east of the village had little other than a minimal Parmelietum with Xanthorion incursions on fencing and calcareous substrata. Tree bases and fallen boles among the denser woodland by the stream had occasional cover of the commoner Cladonia species. Patches of bright yellow Leproplaca chrysodeta illuminated a few darker recesses of bark and fencing. The doubtless much overlooked Gyalideopsis anastomosans with its flaky grey crust was noted on damp tree bases near the old disused observatory up the slope. The acid-loving Foraminella ambigua, Hypogymnia physodes and H. tubulosa were locally common here. Lower down, on tracks and roads leading through and from the village, hedgerows were stippled with Amandinea punctata and Cliostomum griffithii. A roadside quernstone just north of the village had an attractive golden-yellow, black and brown, grey and white mosaic cover of 14 species including Lecanora sulphurea, Catillaria chalybeia and Caloplaca decipiens. A low asbestos-cement roof above a grassy roadside bank in the village had the most finely developed Protoblastenia rupestris seen during the day, among it some tiny patches of Catillaria lenticularis . Although the lichens of the area were nowhere resplendent, particularly in diversity, they nonetheless achieved a colourful presence here and there throughout the day. Yorkshire Naturalists’ Union Excursions in 2000 157 Freshwater Biology (L. Magee) A small stream, less than one metre wide in places, flowed southwards from springs on Oldstead Moor and Cockerdale. It was fast flowing, exceptionally clear with a marly- limestone bottom. The summer water depth was between 4 cm and 20 cm. The stream was unshaded until entering Knever Wood where there was dense tree cover. The stony bottom consisted of small cascades and pools but the pH 8.8 was constant throughout the length surveyed. Alders were examined for signs of damage from Phytophthora fungus, which is fatal to them and spreading in England. No damaged trees were seen. There was evidence that the stream overflows regularly into the adjacent meadows and woodland. The aquatic freshwater shrimp Gammarus pulex was common but other species were present in small numbers. The following were identified in the open stream: Annelid Worms (frequent); Mayfly larvae Baetis rhodani (few), Ecdyonurus torrentis (few), E. venosus (few); Stonefly larvae Brachyptera risi (few); Molluscs Psidium species (frequent); Beetles Elmis species (few); Caddis larvae (Cased) Sericostoma personatum (few) and Agapetus fuscipes (frequent on stones); and Crustacea Gammarus pulex (common). No fish were found. The only invertebrates found in the woodland stream were Gammarus pulex and Chironomid larvae. The only aquatic plant found was Polygonum amphibium (Persicaria amphibia) in a drain leading from the stream. The OS map showed a small fire pond in the woodland but it was found to be completely silted, with a small stream of spring water flowing through. The only invertebrates found were small numbers of the aquatic beetle Hydrobius fuscipes. Equisetum telmateia, a plant in decline in VC63 and VC64 was well established here, as well as in other wet flushes in the valley. Several base-rich wet flushes on the slopes were difficult to investigate for larvae due to the dense vegetation but the recorded flora included: Dactylorhiza fuchsii (frequent), E. telmateia (frequent), Lychnis flos-cuculi (frequent), Carex acutiformis (frequent), Scrophularia auriculata (frequent) and Glyceria declinata (in two places on wet tracks). LINDHOLME MOOR (VC63) 1 July 2000 Introduction (C. A. Howes) Through the good offices of Mr Richard Lyon of Lindholme Grange the YNU was privileged to visit Lindholme Moor, private and isolated, a remarkable survival of ancient woodland, calcareous and acid grassland located on a glacial moraine and the adjacent, equally remarkable survival of uncut lowland raised mire, set in the central core of the extensive Hatfield Moors Site of Special Scientific Interest. A profound debt of gratitude is owed to the Lyon family, particularly the late Jack and Ethel Lyon of Lindholme Hall, for retaining these habitats in a wild and unexploited condition, and to their son Richard for returning adjacent arable to species-rich grassland and for managing habitats for wildlife conservation purposes. Acknowledgement is also due to English Nature for permission to visit adjacent areas of cutover peatland in the early stages of revegetation after being released from commercial peat extraction. Over forty naturalists from fourteen societies, including a contingent from the British Arachnological Society who had travelled up from Nottinghamshire, Derbyshire and Essex, met on a cool slightly overcast day, though conditions brightened up in the afternoon, bringing clouds of butterflies on the splendid ‘stewardship’ grassland. Members of the Doncaster Naturalists’ Society acted as guides during the day and, later, provided a lavish buffet tea for the report meeting held in the Robin Hood and Little John Pub in the adjacent village of Hatfield Woodhouse. The YNU had visited the area on two previous occasions, 12th September 1887 (. Naturalist 13: 83-89) and 12th July 1952 (. Naturalist 77: 181-182), and extracts from these reports were read over tea. Detailed reviews of natural history studies on Hatfield Moors have been compiled by Martin 158 Yorkshire Naturalists’ Union Excursions in 2000 Limbert ( Naturalist 110: 103-110 and 111: 59-60). In the evening (9.30 p.m. to 1.00 a.m.) a group of stalwart members weathered a constant downpour to undertake the evening’s moth trapping, bat and nightjar survey. The use of petrol generators, mercury vapour lamps, ultrasonic bat detectors and radio tracking devices was in telling contrast to the field equipment used by our 19th and mid-20th century forebears on their visits to the moor. Geology and Landscape (C. A. Howes) The morainic deposit forming the c.24 ha Island of Lindholme, which subtly protrudes above the surrounding peatland landscape (to c.4 m OD), is one of a series of low ridges occurring from Thorne in the north, through Bradholme, Tudworth Hall, Lindholme Hall and on to the village of Wroot in the south east. The current interpretation of its origin suggests glacial erratic debris from the west deposited along the edge of a transient ice sheet which surged south-eastwards from the lower Vale of York glacier into the late Devensian Lake Humber as far south as Wroot. Derek Allen and Colin Howes visited the quarry excavated into the morainic deposits adjacent to Lindholme Hall, locating pebbles of Permian Lower Magnesian Limestone, Triassic Sherwood Sandstone, Carboniferous (coal measures) Sandstone, as well as quartzite and flinty chert. A cryoturbation effect in the upper layers indicated the island had been part of the exposed periglacial surface and therefore subject to prolonged freeze-thaw conditions. The calcareous components within this deposit explain the ecological conundrum of calcareous grassland (managed to good effect in accordance with Countryside Stewardship practices by Richard Lyon) occurring within this otherwise lowland raised mire and sandy heath landscape. The Island is surrounded and partly covered by blown sands and dune formations now overlain by peats which commenced formation a little over four thousand years ago. These sands, visible in the deep drainage networks of the surrounding ‘peat fields’ and revealed in places by commercial peat milling activities, are extensively quarried around the western and southern perimeter of the moor. Glacial erratics in the form of large cobbles of coal measures sandstone, extracted by quarrying, were used in the ‘herringbone’ stonework walls of cottages, now demolished, at the western end of Lindholme Bank Road. Interestingly the etymological allusion to Lime trees in the name Lindholme (Linden Island) is confirmed by the pollen record from basal peat which included Tilia, along with Quercus, Corylus, Ulmus, Fraxinus, Pinus, Alnus and Betula (Whitehouse et al. (2001) Lindholme Island. In The Quaternary of East Yorkshire and Lincolnshire: Field Guide (Bateman et al ., eds): 185-193. London: Quaternary Res. Assn. The Lindholme Oaks Quercus robur, which still harbour woodland indicator invertebrates present here in the Bronze Age, have a long documented history, there being twelve oaks listed in a survey of 1607. During our visit the girths of sixty-four were measured, nine of the largest having circumferences at chest height of between ten to fourteen feet, some of the oldest dating to the building of the Lindholme Hall, erected as a shooting lodge in the 1840s. The Lindholme Pines form one of the oldest surviving populations of Pinus sylvestris in Britain. Their remains were examined in the forest zone at the base of Bronze Age peat, and throughout the peat horizons. Documentary references to their occurrence in the 17th and 18th centuries suggest a continued presence on this site for over four thousand years. A small number of mature pines and a population of seedlings were examined in the tall heather areas of the Lindholme Nature Reserve. Encouraged by Joyce Payne’s work (YNU Bulletin (1997) 28: 1-5), interest was shown in the venerable Monkey Puzzles Araucaria auracana to the south of Lindholme Hall. Among the largest specimens in the Doncaster region, it is tempting to speculate their planting being an arboricultural ‘pun’ by Sir Thomas Birkin Bt., who owned the estate for a period after 1907 and received local celebrity in having monkeys roaming freely around the hall. Yorkshire Naturalists’ Union Excursions in 2000 159 Mammals and Lower Vertebrates (C. A. Howes) Although extensive small mammal trapping, owl pellet and fox diet studies have been undertaken here ( Imprint , 1996, 24: 8-15; 1999 26: 38-42; Hatfield Moors Bird Report 1997), at this meeting species were recorded by tracks, trails, sounds and signs. A dead Wood Mouse and a dead Common Shrew were seen on the Lindholme Moor Nature Reserve where Roe-deer slots were present in damp peat along the ridings. Fox droppings, also found here, exclusively contained the remains of Rabbits. Rabbits and their workings were abundant around the edges of arable, heathland and woodland areas of the moraine. Old disused badger setts, now occupied by Rabbit and Fox were examined beneath the rooting systems of the venerable oaks. Droppings provided evidence of Hedgehog on the shorter grassland of the moraine where prey included ground beetles of the genus Pterostichus. Water Voles were well represented along the adjacent Hatfield Waste Drain. Here, according to local resident Ben Craggs, Otters still occur and have allegedly bred in recent years. Presence was confirmed by an old spraint located beneath a plank bridge. Bat detecting in the rain during the evening’s moth trapping and nightjar session indicated that Pipistrelles (45 kh) and a Mysotis species were hunting along the tree-lined Sandy Lane and were also working the insects attracted to the security lighting around the gas terminal on Lindholme Bank Road. Good fish stock, including Three-spined Stickleback, Roach, Bream and Perch, occurs in the Hatfield Waste Drain. Adders were encountered on Lindholme Moor Nature Reserve and a Grass Snake was in tall herbage on the moraine. Ornithology (J. Simmons & H. R. Kirk) In total 36 species, most of them ubiquitous, were identified by sight, song or call. It was nice to encounter with key species such as Nightjar, confirming the value of purpose- cleared glades, and Tree Pipit on the real ‘bit of bog’. Ten Acre Lake held typical waterbirds and provided a good feeding area for hirundines. The Lyon family are to be thanked and congratulated for continued preservation of the real refugia in an area of corporate carnage. Entomology (A. Grayson) Upon arrival, the weather was dull, hazy and overcast, with a temperature of 16°C. Shortly before 2.00 p.m. the sunshine began to break through the cloud, and the temperature rose to 19.5°C. Upon the rise in temperature, females of the horsefly Chrysops relictus became increasingly active around the YNU members and their vehicles. C. relictus favours areas of low-lying land with ditches and drains, and it was no surprise to discover it abundant on Hatfield Moors. The meadowland adjacent to Lindholme Hall contained an abundance of typical flower- loving and grassland insects, including the hoverflies Episyrphus balteatus, Sceava selenitica, Syrphus ribesii, Cheilosia proximo, Eristalis pertinax and Myathropa floria. Along Sandy Lane were, in the main, insects which prefer open, yet sheltered, habitat. Area 1 1 is the main area of lowland mire that has escaped the habitat-destroying peat-winning operations that run up to its border; it is, however, more reminiscent of semi-wet heathland, and is entirely afforested. The mosquito Aedes punctor was abundant here. If Area 11 was a little disappointing then Ash Dump (Area 17) was a bonus. This land has been well worked for peat but is now recovering. Although Ash Dump will not recover to its former state, it still has great potential for insects which favour open areas and ponds and ditches surrounded by tall reeds, grasses and bushes. The damselflies Lestes sponsa, Coenagrion puella and Enallagma cyathigerum were found at Ash Dump, as was the dragonfly Sympetrum danae, which was fairly abundant. Lepidoptera (H. E. Beaumont) It was a pity that rain curtailed fieldwork; it came to the stage that, with micros sticking to 160 Yorkshire Naturalists’ Union Excursions in 2000 sodden sheets, there was little value in continuing. The best moth recorded on a list for the day of 47 lepidoptera is Valerian Pug, the first VC63 record since the supplement to Porritt’s list in 1904. A number of micros were interesting, among them Phlyctaenia perlucidalis, Lozotaeniodes formosanus, Endothenia ericetana and Epinotia demarniana , all of which are still local in Yorkshire although three of them are becoming more frequent. Compilers ’ Note : During the day Gavin Boyd, who had travelled north to attend the meeting, noted the Marbled White Spot Protodeltote pygarga. Knowing the species as reasonably common in the south of England, only when back in Northampton did he realise from the literature that in Yorkshire it is rare. Concerned at first not to have obtained a voucher specimen, he later learned that several specimens had turned up during the evening trapping session he was unable to attend. Hymenoptera Aculeata (M. E. Archer) The species recorded were graded as Common, Frequent, Occasional, or Rare for Yorkshire. In total 16 species were recorded: Solitary Wasp Crabno cribarius (Frequent); Social Wasp Dolichovespula norvegica (Common); Solitary Bees: Andrena bicolor, A. haemorrhoa (both Common); Lazioglossum leucozonium, Sphecodes ephippius (both Occasional), Nomada fabriciana (Common); Bumble Bees: Bombus pascuorum, B. lapidarius, B. terrestris, B. leucorum, B. hortorum, B. pratorum, Psithyrus bohemicus, P. vestalis, and the Honey Bee Apis mellifera (all graded as Common). Svmphvta (G. Boyd) Strongylogaster luneata, Aneugmenus padi and the gall-forming Nematus viridis were recorded and a voucher specimen was retained. All the specimens were female. Coleptera (W. R. Dolling & R. Marsh) Among the 100 species noted during the day, the following are particularly noteworthy: From Compartment 1 1 * (W. R. Dolling) Magdalis barbicornis : there are only two other known Yorkshire records for this Nationally Notable A weevil. Cneorhynus plumbeus: a weevil of litter in dry grassy places; there are only a dozen or so Yorkshire records for this Notable B insect, very few of them recent. Cryptocephalus fulvus: a very local heathland chrysomelid with a dozen or so records for the county. Hippodamia variegata : plenty of county records, nearly all in the south, for this heathland Notable B ladybird. Olibrus liquidus: there are only four confirmed records for this phalacrid, a beetle associated with smutted grasses; a member of a difficult group regarding identification and easily confused with other more common species. From Compartment 15* (R. Marsh) Quedius nemoralis : easily confused with one or two other more common species; about a dozen records, all post- 1980. Ernobius mollis : this beetle develops in conifer cones and in the pith of conifer twigs; 24 records, nearly all pre-1970. Conopalpus testaceus : develops in the dead boughs of various deciduous trees, mainly oak; only two confirmed records for Yorkshire, Nationally Notable B. Omiamima mollina: a Nationally Notable A weevil; a dozen or so Yorkshire records, mostly post- 1970. Philopedon plagiatus : common and very abundant in coastal dunes and sandy habitats all around the coasts of Britain, with a very few inland records from sandy biotopes where there is some degree of openness or disturbance; a recent record from Pot Hill (P. Skidmore) and a much older (H. H. Corbett) record from Barnby Dun. Finally, Curculio venosus : 16 or so Yorkshire records for this oak-dependent weevil, nearly all post- 1970. Yorkshire Naturalists’ Union Excursions in 2000 161 * Compilers ’ Note: Compartment 11 is an area maintained as a nature reserve by the Doncaster Naturalists. Compartment 15 is a stretch of old oak - acidic heathland on glacial moraine. Hemiptera, etc. (W. R. Dolling) In the available time only two Compartments were worked: the old oaks and derelict arable in Compartment 15, and the birch wood on the dried-out peat in Compartment 1 1 . Dermaptera: Forficula auricularia in Compartment 15. Orthoptera: Tetrix undulata in rides in Compartment 11; immature Acrididae in Compartment 15. Hemiptera: these were, on the whole, unremarkable; the exception will be mentioned below. There was a good selection of the expected suite of oak species and also those of open grassland. Compartment 1 1 Birch surprisingly yielded neither of the two shieldbugs associated with it but did produce adults and nymphs of Pentatoma rufipes, the so-called Forest Bug, which was on many kinds of trees in both Compartments, mainly old nymphs but a fair number of adults. The seedbug Kleidocerys resedae , associated with Birch (and Alder) was abundant as were the mirids Lygocoris contaminatus and Psallus falleni and the psyllid Psylla hartigi, all of them common birch specialists. Nymphs of the arboreal nabid Himacerus apterus were also present; this is a scarce species in Yorkshire. Heathery rides in this area produced two heathland bugs, the cicadellid Ulopa reticulata and the mirid Nabis ericetorum. A nymph of the predaceous shieldbug Zicrona caerulea was found under the heather. Sallow on the edge of this Compartment supported the big arboreal froghopper Aphrophora costalis (= forneri, maculata ). (The presence of Myrica on the site suggested that a related species may well be present.) The delphacid Dicranotropis pteridis was numerous on Bracken. The most striking find of the day was the shieldbug Neottiglossa, of which Frank Kenington swept a single specimen in a ride in this Compartment. In a synopsis of the records in Stuart Foster’s database as at October 1996, this species did not appear; it is perhaps new to Yorkshire. Compartment 15 The old oaks had five Cicadellidae: Alebra albostriella, Typhlocyba quercus, Thamnotettix confines, Eurhadina concinna and Jaisus lanio\ the Eurhadina was female only, so there is an outside chance that it was E. ribauti, rare in Yorkshire. The delphacid Dicranotropis pteridis was frequent on Bracken beneath the trees and the cixiid Tachycixius pilosus was also frequent in this part of the area. Pentatoma rufipes was, of course, present in numbers on the oaks along with several mirids: Phylus melanocephalus, P pallipes, Psallus varians, Calocoris quadrimaculatus and Cyllecoris histrionius. Nymphs of Himacerus apterus were present on the oaks and also on the birches, as in Compartment 11. The grassy, herb-rich, derelict arable fields had a substantial hemipterous fauna. Lygaeidae: Nysius ericae, Peritrechus geniculatus, Stygnocoris fuligineus. Nabidae: Nasbicula flavomarginata. Anthocoridae (or Cimicidae): Anthocoris nemorum, Orius niger. Miridae: Hoplomachus thunbergi (on Pilosella ), Plagiognathus chrysanthemi, Heterotoma merioptera, Orthocephalus saltator, Lygus rugulipennis, Capsus ater, Pithanus maerkeli, Notostira elongata, Leptopterna ferrugata, L. dolabrata. Delphacidae: Criomorphus albomarginatus . Cicadellidae: Aphrodes albifrons, Errastunus ocellaris. Hawthorn in this area had the mirid Psallus perrisi and two shieldbugs: nymphs of Acanthossoma haemorrhoidale and an adult of the predator Troilus luridus. An oak here had the mirid Psallus diminutus (females only; so just possibly P. mollis but that would be a first for the county). 162 Yorkshire Naturalists’ Union Excursions in 2000 Arachnology (D. Carr, P. Harvey, T. Faulds, H. Williams & T. Harris) The nationally scarce Philodromus praedatus was collected off old oaks, its typical habitat. It is in fact rather common in Essex and other parts of the south-east. Identification of this group presents many arachnologists with problems, but it has been recorded as far north as Scotland so may well be under-recorded rather than genuinely scarce. Another interesting record was Tetragnatha pinicola, which is also rather widespread in Essex and easily misidentified by anyone unfamiliar with the species. As a mainly southern species it must be towards the edge of its range here. There were several species which are scarce in Essex. The possible Araneus marmoreus pyramidatus (in need of confirmation) is a very characteristic orb- web spider related to the garden spider; it is very local. Compilers’ Note: Fortunately David Carr and Peter Harvey, Recorders of the British Arachnological Society, attended the meeting. They produced a list of 67 species of spider, 29 of them on woodland heath (Lindholme Nature Reserve: peat on podsol on sand); 53 on glacial cap (moraine); with 15 species common to both habitats. An equally numerous list of spiders collected from the area by Tom Faulds, Howard Williams and Trevor Harris includes a further 32 spiders, giving an overall total of 99 species recorded. Plant Galls (T. Higginbottom) After much leaf turning the Birch scrub in square SE6906 finally provided five interesting galls. However, the swelling on the stem of Chamerion angustifolium caused by the micro- moth Mompha nodicollela was one of two highlights of the day discovered in 1 km square SE7005. There was even greater delight with the discovery of the bud gall caused by the cynipid Andricus callidoma on Quercus robur. This was the recorder’s third sighting of A. callidoma in Yorkshire. Each of these rather unusual galls were found by the disused quarry. Few galls were discovered on the mature Oak in the area which was described as ancient woodland. The dense cover of Bracken made exploration of the woodland rather difficult but a future field meeting to the site would be an interesting prospect. 39 galls (12 cecid, 11 cynipid, 11 eriophyid, 1 gall fly, 1 lepidopteran, 1 psyllid and 2 rusts) were seen on 21 hosts during the day. Botany (D. R. Grant) The area reported on is a very small piece of the original Hatfield Peat Moors. This relic area is covered with much Bracken and Silver Birch trees. Conservation workers are removing these and also have succeeded in raising the water table. This has encouraged Sphagnum mosses to become re-established. The open areas had a little Calluna vulgaris, Erica tetralix and both Cottongrasses, Eriophorum angustifolium and E. vaginatum. One part had a good stand of Ceratocapnos claviculata. Along the edge of the woodland Rubus plicatus was frequent, the plants being very tall due to the wet summer. Members were shown the very small colonies of Myrica gale and Andromeda polifolia. Adjacent set-aside fields had much Filago germanica, Conyza canadensis and Anagallis arvensis. The verges of the approach road near the gas well had Rubus nemoralis, R. polyanthemus and R. plicatus , again in quantity. Lichenology (A. Henderson) The haha-like low wall by Lindholme Hall had a mixed calcicole-acidophile crustose flora overgrown here and there by Xanthoria and Phaeophyscia orbicularis. Nearby oaks had invasive Xanthoria polycarpa and Lecanora carpinea on twigs and axils. Stones in the quarry had a mosaic of Lecania erysibe, Catillaria chalybeia, Trapelia coarctata, Rhizocarpon reductum, Lecanora albescens, L. crenulata and Caloplaca holocarpa. Sorediate Peltigera didactyla was very occasional here amongst a Cladonietum which had Yorkshire Naturalists’ Union Excursions in 2000 163 inter alia well developed Cladonia ramulosa, C. floerkeana, C. macilenta and C. pyxidata on a black background of fruiting Placynthiella icmalea. After lunch the wooded edges of the sandy ride south of Lindholme Bank Road repaid exploration with some fine patches of Peltigera rufescens, Collema tenax (v. tenax and v. ceranoides ) and C. crispum , with a trace here and there of indeterminate (probably) Vezdaea thallus, often sprinkled among numerous scattered Nostoc globules (not commune ) in the damper stretches. Oaks in the wooded heath of Compartment 1 1 had a Parmelietum developed only as far as the fruticose elements, Ramalina farinacea and Evernia prunastri. Cladonia species in the understorey included C. humilis, C. squamosa, C. subulata and a little C. parasitica. Physconia grisea was seen once only on a pathside elder in a copse, when returning to the Hall. The constitution of the lichen flora seen during the day suggests that it will benefit most from the avoidance of significant anthropogenic interference. WETHERBY (VC64) 22 July ( J. Kendrew) Introduction (J. Kendrew) On a pleasant morning 24 members and friends met at the Old Wetherby Railway Station and welcomed visitors from Suffolk Wildlife Society, London Natural History Society and a student from Liverpool University. After a briefing about possible locations of interest, members dispersed to explore their individual specialisms. People explored Woodhall, Stockeld Park area, Ox Close Wood, the old railway lines (which have now been turned into cycleways), and areas in the town of Wetherby. Some of the countryside available in 1901 has been lost to industrial and housing development and the ensuing problems of litter and waste were particularly evident on some of the railway lines. Members met at Collingham Memorial Hall for the meeting where 19 affiliated Societies were represented. The President, Colin Howes, chaired the meeting and received apologies from Bob Marsh and Terry Dolan. The reports revealed that a good day had been experienced and members enjoyed a shared buffet during the proceedings. The meeting concluded by Colin Howes thanking the Divisional Secretary for organising an interesting day, at a venue which had been visited by a similar number of Union members 99 years ago. Mammals & Fish (L. Magee) One Mink was seen foraging close to anglers. These creatures are established throughout the river and no doubt their prey includes Rabbits, which were numerous around the Woodhall Estate. Badger and Deer tracks were seen at drinking points. There was no sign of Water Voles which were very abundant until about 25 years ago. Coarse fish fry are usually abundant in backwaters during July but very few were seen during the day. A few minnows were captured among the Milfoil. Anglers confirmed the following species as present: Pike, Barbel, Chub, Perch, Brown Trout, Grayling, Dace, Bream and Stoneloach. Birds (L. Magee) Heron, Kingfisher and Goosander were seen, but very few hirundines or Swifts. Mollusca (A. Norris) The report of the YNU’s visit to the Wetherby area published in 1901 lists only 5 species of mollusc, 3 land and 2 freshwater species, and states that the “extreme heat and especially the dryness militated so much against the collection of land mollusca”. Very hot, dry conditions can make it difficult to locate some species, particularly slugs, but I have never known conditions so bad that only 2 freshwater species could be found in a river like the River Wharfe. On this occasion David Lindley and myself visited 3 different 1 km squares along the old railway line running from Wetherby to Spofforth. In the first part of this walk (44/3948) we recorded 19 land species. The next square (44/3949) produced 18 and the last 164 Yorkshire Naturalists’ Union Excursions in 2000 (44/3849) 12 species: a combined list of 25 species. It was noticeable, however, that the most common species found were mainly garden pests and introductions, such as Milax budapestensis, Arion flagellus and Boettgerilla pallens. In the afternoon we visited the River Wharfe and its banks at Boston Spa to try and locate living examples of the land winkle Pomatias elegans. We were successful in that we found a few living in loose friable soil, close to the River Wharfe within 1 km square (44/4246). The River Wharfe produced some 14 species including large numbers of the freshwater nerite Theodoxus fluviatilis and the freshwater mussel Pseudanodonta complanata. Entomology (G. Boyd) Lepidoptera On a generally overcast day few species of Lepidoptera were seen, although some were present in considerable numbers. Five common species of microlepidoptera were reported, five widespread butterflies and three ubiquitous species of the larger moths. It is perhaps of interest to highlight what was not seen; no Skipper butterflies, no Common Blues Polyommatus icarus or Small Coppers Lycaena phlaeas, no Vanessids apart from a clutch of Peacock Inachis io caterpillars spotted by Mrs Payne who also saw a Small Tortoiseshell. Even the common micromoths such as Anthophila fabriciana, usually abundant over nettle beds, were not observed. It was pleasing, however, to find Ringlet Aphantopus hyperantus abundant and reasonable numbers of Gatekeeper Pyronia tithonus - a butterfly which I never saw at all when I lived in Leeds ten years ago. Mrs Payne reported Yellow Shell Camptogramma bilineata from Ox Close and Riband Wave Idaea aversata from Wood Hall. She saw Large Yellow Underwing Noctua pronuba by the former railway and this species was also seen by Mrs Abbot at SE 395493. A leaf mine on Herb Bennett caused by Stigmella aurella was seen by John Newbould on the disused railway near Knaresborough Road at 44/4940. Hvmenoptera The Bumble Bee Bombus pascuorum and the Common Wasp Vespula vulgaris were recorded from the former Wetherby-Spofforth railway line. Diptera A mere eight species of flies were listed, all common, but including the handsome syrphid Chrysotonum bicinctum, a specimen of which was shown to those members of the Union attending the post-meeting get-together. Possibly the most interesting record was a female specimen of Eristalis abusive. This species is reported most commonly from coastal areas, but it is possibly widely overlooked inland among the hordes of “look-alikes” of the same genus. Plant Galls (J. A. Newbould) During the morning session I searched along the disused railway track which is located in squares 44/3948 and 44/3949 before returning to the Knaresborough Road which is in square 44/4048. Following the wet summer with little sunshine the NVC type W8 Fraxinus excelsior - Acer campestre - Mercurialis perennis woodland, dominated by Ash and Sycamore had few galls. The deep cutting was mostly in shade even on a hot sunny day. Even the numerous sycamore had no nail galls. However, a more open area where the track turns west towards Spofforth produced the gall Taphrina prunii on Prunus spinosa which seems to have exploded in Yorkshire and elsewhere this year. This fungal gall, which totally destroys the fruit, was on bushes also containing unaffected fruit. Towards Knaresborough Road the sawfly gall Blennocampa pusilla was observed. It rolls the leaflet upwards in a spiral from the midrib and was seen on both Rosa canina s.l. and on a rose identified by Mrs P. Abbott as R. virginiana. Eriophyes laevis inangulis on Alder from Collingham (44/3946) and Aceria erinea on Walnut in Colingham churchyard were recorded by C. A. Howes. Eriophyes galii on Yorkshire Naturalists’ Union Excursions in 2000 165 Goosegrass and E. similis on Blackthorn at Linton (44/3847) were recorded by Mrs J. Payne. E. macrorhynchus on Field Maple at Wetherby and Linton (44/4048 and 44/3847) was recorded by J. A. Newbould and Mrs J. Payne. E. goniothorax typicus on Hawthorn at Wetherby (44/4048) and Phytoptus tiliae tiliae on Common Lane at Wetherby by the Wharfe car park (44/4047) were found by J. A. Newbould. Flowering Plants (P. P. Abbott) Most people started the day walking along the disused rail track. The banks of the cutting are now very overgrown with tall scrub so that the patches of typical limestone grassland plants were very few. However, Clinopodium vulgare, Origanum vulgare, Centaureum erythraea and Scabiosa columbaria were seen on the plateau between the tracks and Knautia arvensis and Centaurea scabiosa elsewhere. A puzzling knapweed with ray florets proved to be a rayed form of the common Centaurea nigra. Introduced shrubs provided the main interest in this area. Everyone saw sprays of white flowers of Stephanandra incisa at the end of the car park. The bright pink, pleasantly scented rose, which occurred frequently along the paths above the rail track, was Rosa virginiana. Almost as frequent were the attractive, delicious cherry plums, Prunus cerasifera. Along the Boston Spa road the fruit harvest continued with damson, Prunus domestica ssp. insititia and Malus domestica. In the afternoon most of the botanists went along to Ox Close Wood local nature reserve, where they were pleased to see the Yorkshire Red Data speciality Orobanche reticulata , the nationally scarce Hordelymus europaeus and the splendid, huge, old, small-leaved lime, Tilia cordata. Mycology (J. Payne) An interesting and spectacular bracket fungus Pleurotus cornucopiae was seen by Janetta Lambert in Ox Close. This tiered agaric was growing on a fallen tree-trunk, thought to be elm, which is its usual host. Rusts were rather scarce but there was a widespread infection of Puccinia chaerophylli on Myrrhis odorata in the flood zone on the south of the footbridge. In Ox Close itself Viola hirta bore spores of the later stages of Puccinia violae. A different species of Viola , amongst which it was growing, was not affected. No rust was found on any Epilobium species and, although there was an abundance of Lapsana communis, Puccinia lapsanae was only found after a long search. P. lapsanae also occurred at Wood Hall and the Rose of Sharon planted near the chapel was affected by Melampsora hypericorum. Goat willow Salix caprea at Wood Hall and at the old station car park had a plentiful dusting of the golden spores of Melampsora caprearum. A specimen of Dogwood Cornus sanguinea, collected by J. A. Newbould, proved to be host to the powdery mildew Microsphaera tortilis. Lichenology (A. Henderson & C. J. B. Hitch) Before leaving the car park meeting place the shaded step way leading up from its southwest corner was examined. 26 species, typical of such shaded suburban stonework, were listed from the steps, their wallsides and copings. Physconia grisea was, however, something of a surprise to find here. Sarcopyrenia gibba, noted for its dumbbell-shaped spores, was as ever a delight to come upon. On the roof felting of the adjacent garage were numerous thalli of Physcia caesia. The common Physcia species are no longer considered unusual occurrences on substrata with such a tarry or bituminous admixture, whereon they frequently form swollen, raised thalli, as if shunning close contact with the substrate. The old railway track, curling away to the north-west, was then followed. Ground, stone and trees here are densely shaded, but in areas a little more insolated, trees and saplings had patches of Amandinea punctata and Lecanora chlarotera with some initial Xanthorion colonisation by Phaeophyscia orbicularis, Xanthoria Candelaria and X. polycarpa. 166 Yorkshire Naturalists ’ Union Excursions in 2000 Here and there near the foot of the cutting sides, shaded Fraxinus buttresses and boles had Porina aenea and Opegrapha vulgata with blotches of Collema crispum and Placynthiella icmalea on the damper earth. An exciting find here was the occurrence of a species of Strigula, growing on heavily canopy-shaded rock-facing, and apparently distinct from the three species in the British list. Nowhere here, however, were lichens easy of notice. They were more in evidence on wall copings towards the town centre, and on willows by the river which had a developing Physcietum. After lunch, St. James’ churchyard in mid-town was explored. Here a list of 31 species was considered a respectable total for a shady urban churchyard. Baeomyces rufus was prolific in places on the inner face and the coping of the south churchyard wall. Rinodina pityrea, recorded just inside the gateway, was the most unusual record, a species presumably overlooked in the past because unknown and unrecognised by many lichenologists. The occurrence on the inner face of this same wall of Xanthoria ucrainica, a species only recently recognised in the British flora, also deserves mention. Freshwater Biology (L. Magee) River Wharfe. Collingham Bridge to upstream of Woodhall Bridge. pH 8.4. The flora and fauna of the River Wharfe has been studied by naturalists for two centuries and there are extensive data on individual species. There is, however, much less accurate base line data on the changes in the ecology of the river over a long period. Important early ecological surveys by Union members, Percival, Whitehead and Butcher between 1928 and 1940. No species are known to have disappeared since then but there have been dramatic declines in the populations of the flora and fauna. There have also been explosions in newly established species. No species are known to have been lost but the future of some is precarious, e.g. Salmon, Water Vole, Daubenton’s Bat and Water Crowfoot. The surveys of the main river and the Collingham Beck were confined largely to a comparison of the current situation with past and ongoing surveys. The river was surveyed by wading and the area of plant cover recorded. A characteristic of the Wharfe has always been the development of shingle beds and islands. Since the droughts of the 1990s, small islands composed of silt and gravel have continued to grow in number and in some cases they have become very large (more than 100 m in length), with maturing Salix spp. Examples may be seen upstream and downstream of Pool Bridge. Some islands are protected and have an interesting flora and fauna. In this area, the former large stands of aquatic plants have decreased in recent years but several species still occur in the deeper water: Perfoliate Pondweed Potamogeton perfoliatus\ Swedish Pondweed P. x suecicus which has a very restricted distribution in the British Isles, neither parent occurring in the Wharfe. (There are claims that P. pectinatus has been found in the past.); Cooper’s Pondweed P. x cooperi, an uncommon and difficult hybrid; Stream Watercrowfoot, Ranunculus penicillatus spp. pseudofluitans, a dominant plant until 1975, since when it has steadily decreased, was seen in small patches during the day. It has been largely replaced by the Spiked Watermilfoil, Myriophyllum spicatum, first found on the Wharfe in 1968. More than 80 species of caddis fly have been identified on the Wharfe. Larvae of the following taxa were noted on the day: Brachycentrus subnubilis (the angler’s Grannom), Rhyacophila dorsalis, Limnephilus lunatus, Silo nigricornis, Halesus digitatus, Psychomia pusilla, Agapetes fuscipes, Sericostoma personatum, Mystacides nigra, Hydropsyche species, Lepidostoma hirtum and Anabolia nervosa. Mayflies seen were: (larvae) Blue winged Olive, Ephemerella ignita. Large Dark Olive, Baetis rhodani , Iron Blue, Baetis muticus, Brook Dun Ecdyonurus torrentis, and one imago of Rhythro gena semicolor ata. Odonata: males of the Banded Demoiselle Calopteryx splendens were active throughout the afternoon. Yorkshire Naturalists ’ Union Excursions in 2000 167 Collingham Beck (Collingham Bridge to junction with the Wharfe) In this stretch a pH reading was taken of 8.6. Three species of Mayfly were plentiful. There were a few larvae of the large Green Drake Mayfly Ephemera danica in deep silt. This species is uncommon on the mid- Wharfe, which has suffered from pollution in recent years. A search of a stony-bottomed part of the Beck did not produce any Crayfish. MICKLE FELL (VC65) 19 August (Deborah Millward) Introduction (D. Millward) Considering the remoteness of the location and the previous 24 hours of heavy rain, which had yet to disperse, a surprising 19 members from 12 societies attended the meeting. The clouds lifted by the start of the excursion but bank-full streams and washed away footbridges deterred a sensible proportion from reaching the goal and these members concentrated their efforts on the area of shake holes south of Connypot Beck. The more intrepid shed boots and waded the beck only to be confronted by a second stream in full spate that was crossed on a submerged strand of wire fence with members hanging on grimly to the upper strand. An unexpectedly long slow trek across the blanket bog followed which, disappointingly, considerably reduced the time available for recording on Mickle Fell. However, an intrepid few made it to the high ground and back to the meeting in Brough in time. Two members made the sound and enviable decision to continue recording, taking advantage of the now beautiful evening and maximising their investment in the long hike. The return trip for all groups was blessed with much reduced stream crossings. It is perhaps no coincidence that the Union has not visited this site since 1910. Thanks have been sent to the Strathmore Estate. Mammals and Lower Vertebrates (M. J. A. Thompson) On the 1889 excursion to Mickle Fell only two mammals were recorded, the common shrew and the mole. On the VC65 outing in 2000 four mammal species were recorded, namely, rabbit, mole, field vole and stoat. Both rabbit burrows and mole hills were abundant in the strips of limestone rock outcrops that surfaced above the heather. A single field vole was sighted, probably in an area of low population density due to moorland habitat. A dead adult stoat was found in a snapper trap set on a wooden pole across Force Beck, as part of ‘vermin’ control on a well-keepered grouse moor. Both common frogs and viviparous lizard were seen by members. Ornithology (M. J. A. Thompson) Comparison of the bird report in the 1889 Naturalist of the excursion to Mickle Fell (Sept. 1889) with that for the VC65 outing in 2000 has had its difficulties. Not only were more habitats covered in the earlier report but, also, on that occasion the excursion took three days. The ornithologists recorded 38 species in 1889 (25 residents and 13 migrants), whereas on the 2000 visit 15 species were noted (1 1 residents and 4 migrants). The birds in 1889 were listed at the end of the Union’s report under the scientific names that were in use at that time. Since then, due to more accurate taxonomy, many of the generic and specific names have changed, so sorting out which species were which proved to be an interesting exercise. Listed below are the bird lists for both excursions (1889 classification in brackets): 1889 2000 Grey Heron Teal Buzzard Kestrel Red Grouse Ardea cinerea (A, cinerea ) Anas crecca (A. crecca) Buteo buteo Falco tinnunculus Lagopus lagopus (L. scoticus ) X X X X X X 168 Yorkshire Naturalists ’ Union Excursions in 2000 1889 2000 Black Grouse Tetrao tetrix (T. tetrix) X Oystercatcher Haematopus ostralegus - X Golden Plover Pluvialis apricaria ( Charadrius pluvialis) X - Lapwing Vanellus vanellus ( V. cristatus ) X X Dunlin Calidris alpina ( Tringa alpina ) X - Snipe Gallinago gallinago - X Curlew Numenius arquata ( N . arquata) X X Redshank Tringa tetanus - X Common Sandpiper Actitis hypoleucos ( Totanus hypoleucos ) X - Wood Pigeon Columba palumbus (C. palumbus) X - Swift Apus apus ( Cypselus apus) X - Kingfisher Alcedo atthis (A. ispida ) X - Skylark Alauda arvensis (A. arvensis ) X - Sand Martin Riparia riparia ( Cotile riparia) X - House Martin Delichon urbica ( Chelidon urbica ) X - Swallow Hirundo rustica (H. rustica) X - Meadow Pipit Anthus pratensis (A. pratensis) X X Pied Wagtail Motacilla alba ( M . lugubris ) X - Grey Wagtail M. cinerea (M. melanope) X - Dunnock Prunella modularis {. Accentor modularis) X - Robin Erithacus rubecula ( E . rubecula) X - Redstart Phoenicurus phoenicurus ( Ruticilla phoenicurus ) X - Whinchat Saxicola rubetra ( Pratincola rubetra) X - Wheatear Oenanthe oenanthe ( Saxicola oenanthe) X X Blackbird Turdus merula (T. merula) X - Ring Ousel T. torquatus ( T. torquatus ) X - Song Thrush T. philomelos ( T. musicus ) X - Willow Warbler Phylloscopus trochilus ( P trochilus ) X - Blue Tit Parus caeruleus (P. caeruleus) X - Dipper Cinclus cinclus (C. aquaticus ) X X Rook Corvus frugilegus (C. frugilegus) X X Carrion Crow C. corone (C. corone) X X Raven C. corax (C. corax) X X Starling Sturnus vulgaris ( S . vulgaris) X X House Sparrow Passer domesticus ( P. domesticus) X - Chaffinch Fringilla coelebs (F. coelebs) X - Twite Carduelis flavirostris {Linota flavirostris) X - Linnet C. cannabina {Linota cannabina) X - 38 15 Thirteen of the species listed for 1889 would not (or only possibly rarely) be seen in a montane heathland habitat. Some summer migrants, such as the Swift, would have moved on by the time of the 2000 VC65 excursion in the third week of August. Black Grouse are still in the area, but not immediately local. The presence of Buzzard is indicative of the continuing recovery of this raptor in north- west Yorkshire. Oystercatchers, as an inland breeding species, were first recorded in Yorkshire in 1936 (Mather 1986) and, therefore, would not have been seen in 1889. Although 5 bird species were seen in 2000 that were not recorded in 1889, namely Kestrel, Buzzard, Redshank, Snipe and Oystercatcher, there appears to have been a reduction in the total number of species, as well as the number of birds. Declines nationally have been recorded amongst Twite, Linnet, Dunlin, Golden Plover, Whinchat and Ring Ousel (Mead Yorkshire Naturalists’ Union Excursions in 2000 169 2000), and some of these declines may account for their not being seen in 2000 and the differences in total numbers between the two excursions. References Mather, J. R. (1986) The Birds of Yorkshire. London: Croom Helm. Mead, C. (2000) The State of the Nations’ Birds. Stowmarket, Suffolk: Whittet Books. CONCHOLOGY (A. NORRIS) It is no wonder that the Upper Teesdale National Nature Reserve on Mickle Fell has never been surveyed for snails before this date. The two-hour walk to the base of the crag, from the nearest point cars could be parked, over very rough and wet moorland, was hard enough. On top of this the trio concerned (David Lindley, Tom Blocked and A.N.) had to cross several small rivers (in flood), a case of removing boots, etc., and wading through very cold water. The walk did produce at least two records, but not of mollusca. Over two of the rivers logs had been placed and pole traps set inside a wire tunnel. In both of these traps were freshly killed Weasels. Presumably the local Game Keeper had placed the traps on the logs. The visit to the limestone crags proved to be very interesting from the point of view of the mollusca. Although David Dickinson and A.N. recorded only 22 species of mollusca from the reserve, all but two, Arion ater and Pyramidula rupestris , can claim to be new altitude records for the species in England. In most cases at least 1,000 feet has been added to the highest previous record for England. The crags have an average altitude at their base of 2,400 ft and 2,450 ft at the top. Contour lines run along the base and the top of the crags, for almost the full length of the limestone outcrop. All of the records come from between these two altitudes. 22 species in total were recorded from the Upper Teesdale National Nature Reserve, 8 species of mollusc from NY35/80-24-, 21 from NY35/81-24- and 8 from NY35/82-24-. Time was the main constraint on recording in the area. We felt that further species could and will be found in the area, given time. A number of calcareous flushes occur at the base of the cliffs, which in dryer weather could produce some very interesting records. D.L. had to leave early, after only 2 hours on site. T.B. and A.N. recorded specimens from all three 1 km squares, before tackling the two-hour trek back - one hour into which the will to live was lost. The cars were finally reached at 6.30 p.m., too late for attendance at the indoor meeting. The list of species found is as follows: 80-24- 81-24- 82-24- Previous Altitude Record Lymnaea truncatula X 1,350 Carychium minimum X ? Carychium tridentatum X X 1,550 Cochlicopa lubrica X X X 1,400 Pyramidula rupestris X X X 3,054 Columella aspera X ? Vertigo substriata X 1,230 Lauria cylindracea X X 1,800 Discus rotunatus X 1,400 Arion ater agg. X X X 3,000 Arion circumscriptus X 1,250 Arion internedius X 1,325 Vitrina pellucida X X ? Vitrea subrimata X ? Vitrea contracta X X X 1,325 170 Yorkshire Naturalists ’ Union Excursions in 2000 80-24- 81-24- 82-24- Previous Altitude Record Nesovitrea hammonis X X 1,400 Aegopinella nitidula X 1,400 Oxychilus alliarius X 1,400 Deroceras leave X 1,230 Deroceras agreste * X X X ? Clausilia dubia X 2,350 Arianta arbustorumi X X 2,400 Notes: * All of the specimens of Deroceras agreste found proved to be young and therefore could not be confirmed by dissection. However, the identification is considered to be correct. f Recorded from an altitude of 2,425 ft. Crustacea (G. Fryer) Streams were in spate, making moorland crossings difficult. Small peaty pools at c.600 metres south of Mickle Fell revealed the expected paucity of micro-crustaceans, but collecting was desultory. Here the almost ubiquitous Chydorus sphaericus was accompanied by C. ovalis, a species often found in acidic habitats in the northern, but not the southern, Pennines. A rotifer Keratella sp. was plentiful, suggesting that such habitats merit attention from students of this currently neglected group. Botany (D. Millward) Lune Head Moss and Philip Reed Moss supported the typical flora of a managed blanket bog with dominant Calluna vulgaris and Eriophorum vaginatum but not to the exclusion of other more significant species. All the usual “berries” were there with noticeably frequent Rubus chamaemorus, none of which bore fruit. Vaccinium myrtillus, V. vitis-idaea, V. oxycoccus and Empetrum nigrum, were more dispersed, the first being the most frequent. Drosera rotundifolia occurred on barer peat. Closer to the summit where some calcareous influence ameliorated the very acidic conditions, Viola palustris and Chrysosplenium oppositifolium were recorded. As expected, once onto the limestone flanks of the fell, diversity increased dramatically. Search was concentrated on the southern edge of 35/8024 within the grazing enclosure where rabbits kept a close turf. Here Saxifraga hypnoides, Cochlearia pyrenaica and Galium boreale mingled with the commoner Viola lutea, Alchemilla filicaulis ssp. vestita and Carex caryophyllea. Sedges as usual were concentrated in the flushes where both Carex viridula ssp. brachyrryncha and ssp. oedocarpa flourished with pale Myosotis stolonifera, Epilobium alsinifolium and Selaginella selaginoides. Pteridophytes were uncommon throughout the day; only Blechnum spicant, Cryptogramma crispa and Dryopteris dilatata were seen on the acid ground, and Cystopteris fragilis and Asplenium viride on limestone rocks. Bryology (T. L. Blockeel) The long and arduous walk to the summit of the Fell left only limited time for bryologising. Exploration was confined to parts of the south-facing slopes of the summit ridge. At the western end there was a small area of block scree with grit boulders, supporting plentiful Andreaea rupestris and A. rothii subsp. rothii, with Barbilophozia atlantica in the hollows. Further towards the summit, grit rocks had Dicranum fuscescens and Lophozia incisa. The dry south-facing limestone was notable for the plentiful occurrence of Pseudoleskeella catenulata. Schistidium species were also plentiful, and they included S. robustrum and S. trichodon , the latter new to the vice-county but previously known from Yorkshire Naturalists’ Union Excursions in 2000 171 the Widdybank area of Teesdale. Other species on the limestone crags included Frullania tamarisci, Scapania aspera, Porella cordaeana, Didymodon ferrugineus and Pohlia cruda. Some boulders had quantities of Barbilophozia barbata associated with Ditrichum flexicaule s.str. There are a number of springs and runnels below the crags. Particularly noteworthy here was a patch of the arctic-alpine moss Cinclidium stygium. Flushes supported Dicranum bonjeanii, Hymenostylium recurvirostrum, Philonotis calcarea, Climacium dendroides and Palustriella commutata var. falcata. One springhead had Marchantia polymorpha subsp. montivagans ; this is new to the vice-county, and as subsp. polymorpha grew nearby, the two could be readily compared. 81 species were recorded along the summit ridge. Mycology (C. S. V. Yeates) Following the rather misleading information that it would only take “an hour or so” to walk to the top of the fell, the writer opted not to do the generally much more profitable slow, careful searching of a small area, but set off at a fast pace for the summit. In the event, a combination of a shortage of time at the objective, highly grazed or very wet vegetation and an apparent paucity of fungi made for a frustrating time. From a mycologist’s point of view the noteworthy plants around the top of the fell were disappointingly healthy. In the acidic rushy grassland the ivory-coloured discomycete Hymenoscyphus herbarum was frequent on dead Cirsium palustre stems; and, among a number of fungi recorded on rabbit pellets, another discomycete, the long-haired Lasiobolus cuniculi has few Yorkshire records, largely because the genus has been recently revised and earlier records cannot be redisposed without voucher material. The most interesting species by far was that found by Albert Henderson, about which the reader is directed to the lichen report. Lichenology (M. R. D. Seaward & A. Henderson) Although the two lichenologists did not make it to the top, being slow movers and deterred by a fast-flowing beck, there was much to claim attention. The juxtaposition of acid and alkaline habitats ( Callunetum , acid grassland, gritstones and limestone outcrops), in addition to walls composed of a miscellany of stone types, provided a rich source of lichen records. In all well over 100 terricolous and saxicolous species were recorded. Perhaps the most striking record from the long stretch of boundary wall was the lichenicolous fungus, Zwackhiomyces coepulonus, growing on Xanthora parietina on the northeast shoulder of mortar capping. This is the first British record for this species (det. C. J. B. Hitch, conf. D. L. Hawksworth), previously known only from Spain and the Canaries. The dark immersed or sessile perithecia have 1-septate, hyaline, finely verrucose spores, 16-20 p x 5-8 pi (see British Lichen Society Bulletin 88 [2001]: 77 and Nova Hedwigia 51: 283-360). Several lignicolous species were observed on old fence posts, marking this western boundary of our study area. Lecanora aitema (the only recent record from Yorkshire) was most notable here among the commoner lignicolous flora of Lecanora conizaeoides, L. chlarotera, L. symmicta, Lecania cyrtella, Mycoblastus sterilis and Micarea botryoides. Of particular interest in the acidic heathland areas were many Cladonia species (at least 16), especially the fine show of C. portentosa on hummocks. Much of the wet peaty earth was stippled with dark green globuloid Botrydina-i ype crusts of Omphalina , nowhere seen fruiting and therefore indeterminate to specific level; as distinct from the squamulose Coriscium- type thalli of this genus which were occasionally found in fruit and determined as O. hudsoniana. Together these representatives of this agaricoid lichen genus constituted a quite significant element of the heathland surface cover; occasional well developed patches of the thick glaucous grey squamules of Trapeliopsis glaucolepidea were another interesting feature. Limestone outcrops supported a reasonably diverse flora, although several lichens characteristic of this habitat, including the more colourful Caloplaca species, were 172 Yorkshire Naturalists’ Union Excursions in 2000 noticeably absent; the associated soil in crevices of one exposed ridge at c.500 m was dominated by Squamarina cartilaginea (much less common now on Yorkshire limestone), and a few thalli of Solorina saccata (similarly disappearing due to habitat disturbance) were found upon a smaller outcrop at c.550 m. Some of the hardest limestone bore the immersed black perithecia of Porina linearis. Acarospora glaucocarpa was seen here and there on the shoulders and tops of outcrops, growing occasionally with Staurothele caesia. Shallow soils over limestone, heavily grazed by sheep and rabbits, provided another interesting habitat with mixtures of both calcicolous and acidophilous lichens, such as Coelocaulon aculeatum/muricatum, Cladonia rangiformis, C. cervicornis and Peltigera species; and at one roadside site, P. leucophlebia, a rare lichen in England (particularly in fruit, as here) was much in evidence over an area of c. 1 m2. A short visit was also made to the spoil heaps of a former lead mine (35/834198), the stone waste supporting a lichen flora similar to but more luxuriant (particularly Stereocaulon vesuvianum ) than that observed on local walls, with the addition of Sphaerophorus globosus', the mineral soil was equally rich, with such species as Cladonia arbuscula, C. ramulosa and Coelocaulon aculeatum. EXCURSIONS IN 1999: ADDENDA See: Naturalist 126: 41-48 (2001) Leyburn Shawl Woods and Quarry (VC65) 24 July (J. A. Newbould) Entomology (J. A. Newbould) In the absence of most of the Union’s recognised entomologists, entomological recording was carried out by J. A. Newbould, C. S. V. Yeates and Helen Thornton in Leyburn Shawl, then through Warren Wood into the thicker shelter of Gillfield Wood. With temperatures at well over 20°C the south-facing slopes of Leyburn Shawl should have afforded an interesting day. Recording of lepidoptera was, however, made harder by a strong southwesterly which forecasters had said would be blowing at 25 mph, and which kept many species well hidden in sheltered places. The most common butterfly seen across the area was the Meadow Brown (. Maniola jurtina ). Not until Warren Wood was reached were other species recorded. Here Small Heath ( Coenonympha pamphilus), Ringlet (Apantopus hyperantus ) and Small Copper ( Lycaena phlaeas ) were added to the list, whilst in Gillfield Wood Green-veined White (. Artogeia napi) and Small White (A. rapae) were also present. Later at the meeting members reported Common Blue ( Polyommatus icarus) and Small Tortoiseshell ( Aglais urticae) in the area of Moor Quarry. Tim Flint reported seeing the dragonfly Sympetrum striolatum in the quarry area. Flowering Plants (I. C. Lawrence) The large quarry floor was the main focus of attention during the morning. Large areas were sparse in vegetation, the richer areas being around the pond and stream and hummocks at the western end which yielded two of the rarest plants of the day: Orobanche alba and Gentianella amarella subsp. septentrionalis. The wet areas had large quantities of Myosotis scorpioides, and Veronica anagallis-aquatica was locally common. In the pools there was Hippuris vulgaris along with Myriophyllum spicatum, Potamogeton natans and stands of Typha latifolia. Other plants of interest in the damper areas were Carex viridula subsp. brachyrrhyncha, Dactylorhiza fuchsii and the hybrid willow-herb Epilobium parviflorum x E. montanum. Other species included Galium sterneri, Sagina nodosa and the established Sedum alba and S. rupestris. Some 94 species were recorded in this area during the morning. The afternoon was spent in the nearby woodland and on the Shawl itself where another fine group of Orobanche alba was seen on the cliff edge. Here several Yorkshire Naturalists’ Union Excursions in 2000 173 calcicoles were noted including Inula conyza, Arabis hirsuta, Filipendula vulgaris, Clinopodium vulgare and Listera ovata. Other species of interest were Plantago major subsp. intermedia in the damp area of the quarry, Minuartia verna and Agrimonia procera. Finally, we were shown a stand of Sambucus ebulus on the roadside leading to the council tip. NB. At a further location for Orobanche alba in open woodland on Leyburn Shawl (44/096907), the following 2 metre square quadrat represents the position (measurements on the Domin scale): Orobanche alba , 3 spikes D1 Linum catharticum D2 Thymus praecox D3 Cirsium vulgare D2 Hedera helix D4 Hieracium pilosella D2 Teucrium scorodonia D2 Fragaria vesca D2 Centaurea nigra D2 Helianthemum nummularium D6 Sedum acre D3 Achillea millefolium D2 Sangusorbia minor D2 Bare ground D5 Festuca ovina D4 Limestone rock D4 Senecio jacobea D2 (DIO = 91-100%, D9 = 76-90%, D8 = 51-75%, D7 = 34-50%, D6 = 26-33%, D5 = 11-25%, D4 = 4-10%, D3 represents many individuals, D2 several individuals and D1 few individuals). DARNBROOK FARM, Malham Tarn Estate (VC64) 14 August Freshwater Biology (contd) (D. T. Richardson) The most outstanding feature of the calcareous flushes was the presence of very large numbers of the freshwater triclad Crenobia alpina. It inhabits spring heads and small swiftly flowing streams at high altitudes, and is incapable of enduring temperatures of above 15°C for any length of time. The water temperature of these flushes was between 8°C and 10°C, a clear indication of their deep-seated origins. Water Analyses (Streams in flood following heavy rain during the previous 24 hours): Darnbrook Beck Cowside Beck Thoragill Beck Calcium+Magnesium mgs CaCCh 1 1 56 172 130 Calcium mgs CaCCh L1 52 166 124 Alkalinity mgs CaCCL 1 1 46 162 118 pH 7.58 7.75 7.51 Water Temperature °C 14 11 12 The low mineral content of Darnbrook Beck is a direct consequence of run-off of surface water from the moor. Darnbrook Beck has its origins in the perennially wet raw peat bogs of Darnbrook and Fountains Fells. The bogs are separated from the underlying Yoredale and Lower Carboniferous Limestone rocks by a covering of Glacial Drift. Any precipitation which takes place results in immediate run-off and can cause flash floods. This also causes dilution of the mineral content of the water, which is demonstrated in the analytical figures given above. There was heavy precipitation the day before sampling the admixture of the run-off water reducing the calcium content of the water to about one third of that normally experienced. Cowside Beck, whilst taking some water from the peat bogs of Fountains Fell, draws far 174 Book Reviews more from the actual Carboniferous Limestone, and only in cases of extreme and prolonged precipitation is any degree of dilution obvious. Thoragill Beck is completely different: it has deep-seated origins within the Carboniferous Limestone, drawing water from an extensive aquifer. Precipitation does not have an immediate effect on its mineral composition and several days pass before signs of dilution are apparent. This is because the water held back in the aquifer has to be displaced before the diluted water appears. The classic example of this phenomenon is the spring at the base of Kilnsey Crag where it takes 10 to 12 days before the effects of precipitation become apparent. BOOK REVIEWS Environmental Contamination in Antarctica. A Challenge to Analytical Chemistry edited by S. Caroli, P. Cescon and D. W. H. Walton. Pp. xiv + 406 (inch numerous figures & tables). Elsevier, Amsterdam. 2001 . 147.50 hardback. This is a most valuable contribution to our knowledge of how and to what extent pollutants reach remote areas of the globe. Antarctica’s once uniquely clean environment is clearly receiving a wide range of man-made contaminants; whilst many of these are present at low levels, they are nevertheless capable of being detected by present-day analytical chemistry, such measurements providing baseline data invaluable for time-space analyses. Fifteen chapters covering many aspects of environmental chemistry and environmental monitoring of air, snow, ice, sea-water, sediments and ecosystems are provided by 36 (mainly Italian) authors. The use of plants as biomonitors receives relatively less attention, particularly, and rather surprisingly, lichens (rarely mentioned) and bryophytes (not mentioned) which have been extensively used in Antarctica for this purpose. The measurement and monitoring of radionuclides, so important for appreciating their global impact, has also been omitted. Despite these reservations, this work should be consulted by all those involved in monitoring our environment. The stimulating text is supported by a wealth of reference material, complemented by extensive bibliographies (not alphabetic), author and subject indexes. Rather pricey for individual purchase, so recommend it to your library. MRDS Food: a History by Felipe Fernandez-Armesto. Pp. xvi + 287, plus 16 pp. b/w plates. Macmillan, London. 2001. £20.00 hardback. A most readable and thoroughly enjoyable exploration of the global history of food, albeit somewhat sketchy, tracing its production, processing, preparation and ultimate consumption. This is a book on the history of science and society which touches on ecological and environmental issues of the past, current crises and future implications. MJD The Peak District Joural of Natural History and Archaeology edited by Melvyn Jones and Ian D. Rotherham. Volume 2, pp. 96, incl. colour & b/w plates, and line drawings. £5.00 plus £2.00 p.&p. from Wildtrack Publishing, P.O. Box 1142, Sheffield SI 1SZ. This annual publication contains short articles on various aspects of the wildlife, conservation, history and archaeology of the Peak District National Park and surrounding areas. This volume (2000) includes several articles of natural history interest, particularly a paper on ancient coppiced woods in Derbyshire, a study of the slug Umax cinereoniger, and a case study of deer in the urban fringe. Articles are written in non-technical language 175 and an accessible, very readable style - indeed, the one on deer is positively jocular in tone! The presentation is of a very high quality and the A4 format makes for easy reading, with large font, clear maps and diagrams, and a liberal quantity of high quality monochrome and colour plates. The glossy, full-colour cover is an attractive feature that should help to sell the journal. Whilst the content is obviously of primary interest to those working in the Peak District, several of the papers are of sufficient general interest to appeal to a wider audience. Each volume includes a featured local organisation - in this case, the Derbyshire Archaeological Society - plus a few book reviews and advertisements. I am confident that many members of the Yorkshire Naturalists’ Union would enjoy reading it. MAA CONTRIBUTORS Abbott, P. P. 165 Archer, M. E. 31-38, 149, 160 Atherden, M. A. 138, 174-175 Beaumont, H. E. 80, 159-160 Blackburn, J. M. 141-146, 151-152, 156 Blocked, T. L. 141-146, 170-171 Boyd, G. 149, 160, 164 Brand, G. M. 85-99 Carr, D. 162 Caverhill, D. A. 138 Cook, P. J. 147 Crossley, R. 17-18 Curtis, W.F. 147-148 Dale, J. E. 152 Delany, M. J. 137-138 Dolling, D. W. 153-155, 160-161 Faulds, T. 162 Frost, H. M. 38-40 Fryer, G. 40, 84, 139-141, 170 Goulder, R. 57-64 Grant, D. R. 162 Grayson, A. 159 Harris, T. 162 Harvey, R. 162 Henderson, A. 147-172 Higginbottom, T. 162 Hinton, V. A. 146 Hitch, C. J. B. 165-166 Howes, C. 41-56, 101-130, 131-132, 133- 137, 147, 150, 157-158, 159 Jones, J. 19-30 Kendrew, J. 163 Kirk, H. R. 159 Lambert, J. 147-172 Lawrence, I. C. 172-173 Magee, L. 56, 157, 163, 166-167 Marsh, R. J. 69-80, 160 Middleton, R. 150 Millward, D. 167, 170 Morrell, G. 147 Newbound, J. A. 164-165, 172 Norris, A. 81-83, 148-149, 153, 163-164, 169-170 Payne, J. 150-151, 153, 155, 165 Payne, K. 150-151, 155 Richardson, D. T. 173-174 Seaward, M. R. D. 38, 64, 68, 99-100, 132, 146, 171-172, 174 Simmons, J. 159 Suner, R. 65-68 Sykes, N. 155-156 Thompson, M. J. A. 152, 167-169 Whitaker, T. M. 3-17 Williams, H. 162 Yeates, C. S. V. 171 176 INDEX Book Reviews 38-40, 56, 64, 68, 80, 84, 99-100, 131-132, 137-138, 146, 174-175. Botany Effects of the reinstatement of coppice management on the vascular ground flora at Eaves Wood SSSI, Silverdale, Lancashire, 19-30; Aquatic plants at North Cave, a new Yorkshire Wildlife Trust Reserve comprising former gravel workings, 57-64. Bryology YNU Bryological Section: annual report for 2000-2001, 141-146. Coleoptera Coleoptera report for 1995-2001, 69-80. Diptera Notes on some Yorkshire Diptera, 2001, 17-18. Ecology Effects of the reinstatement of coppice management on the vascular ground flora at Eaves Wood SSSI, Silverdale, Lancashire, 19-30; Aquatic plants at North Cave, a new Yorkshire Wildlife Trust Reserve comprising former gravel workings, 57-64. Hymenoptera Wasps and bees of Cronelian and Cayton Bays and Osgodby Point in Watsonian Yorkshire, 31-38. Lepidoptera Status of the Small Pearl-bordered Fritillary, Boloria selene, in western Yorkshire, 3-17. Mammals Red in tooth and claw: 1. Studies on the history of the wild cat Fellis silvestris in Yorkshire (2000 Presidential Address, part 1), 41-56; Red in tooth and claw: 2. Studies on the nature history of the domestic cat Felis catus in Yorkshire (2000 Presidential Address, part 2), 101-130. Mycology Finding Fungi (2001 Presidential Address), 85-99. Obituary Dr Lewis Lloyd Evans (1916-2001), 81-83. Odonata Distribution of the Banded Demoiselle Calopteryx splendens along the River Derwent, 65-68. Ornithology Barn owls in churchwardens’ accounts of the 17th and 18th centuries: an indication of past abundance in East Yorkshire, 133-137; The dawn chorus and how its singers are conducted, 139-141. Yorkshire Naturalists’ Union 2000 Presidential Address (C. A. Howes), 41-56, 101-130; 2001 Presidential Address G. M. Brand), 85-99; YNU excursions in 2000, 147-172; Excursions in 1999: addenda, 172-174. Irish Naturalists’ Journal The Irish Naturalists’ Journal, sucessor to the Irish Naturalist, commenced publication in 1925. The quarterly issues publish papers on all aspects of Irish natural history, including botany, ecology, geography, geology and zoology. The Journal also publishes distribution records, principally for cetaceans, fish, insects and plants, together with short notes and book reviews. Current subscription rates for four issues (including postage): 25.00 (£15.00 stg); Students 8.00 (£5.00 stg). Further details from: Mr Brian Nelson, INJ, Department of Zoology, Ulster Meseum, Botanic Gardens, Belfast BT9 5AB. Titus Wilson Kent Works • Burneside Road • Kendal • Cumbria • LA9 4RL Tel. 01539 720244 Specialist printers /binders of Academic Journals , Catalogues and Private Publications Our service includes attending to worldwide distribution Binding Why not have your copies of The Naturalist bound into volumes? One year’s issues per volume, or alternatively two years in one volume at less cost than binding as two separate volumes. We are also experienced in binding theses and experts in the re-binding and repairing of all books. Spink & Thackray Broomfield Binder Back Broomfield Crescent Telephone: 0113 2780353 LEEDS LS6 3BP Fax 01 13 2304155 www.spinkandthackray.co.uk Printed in Great Britain by Titus Wilson & Son, Kendal ISSN 0028-0771 Latest publication of the Yorkshire Naturalists’ Union ALIEN PLANTS OF YORKSHIRE GEOFFREY WILMORE The first comprehensive modern publication dealing with the alien plants of any major British county, detailing the flora (almost 1300 species) of Yorkshire’s wool waste dumps, railway sidings, sewage works, shoddy fields, industrial wasteland, dockland and oil mill sidings. Pp. 316, including 15 line drawings; published January 2000 Normal price: £15.00 (plus £2.50 per copy p.&p.) Reduced price for members of the YNU: £12.00 (plus £2.50 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Mr John A. Newbold, Stonecroft, 3 Brookmead Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS THE FRESHWATER CRUSTACEA OF YORKSHIRE a faunistic & ecological survey GEOFFREY FRYER The crustacean fauna of Yorkshire reflects the great physiographic diversity of the region. Adopting an ecological approach, this book considers the Yorkshire fauna in relation to climate, topography, geology, soils and water chemistry, always keeping in mind that it is dealing with living organisms whose habits, requirements and physiological limitations determine exactly where they live. Matters covered include the ecological background; faunal assemblages and their regional attributes; an analysis of the factors that determine distribution patterns, many of which are mapped; wide geographical aspects; and conservation. Large areas, such as the Pennines, Howgill Fells, North Eastern uplands and the lowland plains are surveyed. So too are localised regions including Whernside, the Malham area, lowland heaths, and the largest lakes, as well as habitats such as upland tarns, seepages, cold springs, small lowland ponds, inland saline waters. Notes are given on every species recorded, including parasitic forms. Price £8.00 (plus £2.50 p.&p.) Special offer to members of the Yorkshire Naturalists’ Union £5.00 (plus £2.50 p.&p.) Please make cheques payable to Yorkshire Naturalists’ Union. Available from: Professor M. R. D. Seaward, Department of Environ- mental Science, University of Bradford, Bradford BD7 1DP. Heckman BINDERY, INC. Bound-lb-Please® JULY 03 N. MANCHESTER, INDIANA 46962