JANUARY - MARCH 1984 (no 968) VOLUME 109 THE NATURALIST Quarterly Journal of Natural History for the North of England Edited by M. R. D. SEAWARD, MSc, PhD, DSc, FLS, The University, Bradford PAGE CONTENTS 3 The Occurrence of Littorina spp. in Robin Hood's Bay, North Yorkshire, with a Key to British Species — I. W. Jardine 9 Marchesinia mackaii and other Nottinghamshire Bryophytes — T. L. Blockeel 1 1 Six Species of Megaselia (Diptera, Phoridae) from Northern England, New to Britain, and including Two New to Science — Ft. H. L. Disney 19 The Vascular Flora of Disused Chalk Pits and Quarries in the Yorkshire Wolds — Richard G. Jefferson 23 The Solitary Bees and Wasps (Hymenoptera: Aculeata ) of a Sand-pit at Swincarr Plantation, near York — Michael E. Archer 27 Loss of Wetlands in Sprotbrough Parish, Doncaster — P. F. Ulf-Hansen 29 Entomological Reports for 1981-82 — R. Crossley 31 The Blue-bottle Fly Calliphora vicina R.-D. as a Parasite (Primary Myiasis Agent), Particularly on Small Mammals — Y. Z. Erzinclioglu and Sharon W. Davies 35 Y.N.U. Bryological Section: Annual Report 1982 — T. L. Blockeel 37 Fungus Forays in 1981 — Spring Foray, Darlington, 7-1 1 May; Autumn Foray, Whitby, 10-14 September — T. F. Hering 40 Field Note 10, 22, 26, Book Reviews 30, 34, 40 PUBLISHED BY THE YORKSHIRE NATURALISTS’ UNION ‘The Naturalist’ is available in microform UNIVERSITY MICROFILMS INTERNATIONAL 300 North Zceb Road Dept PR Ann Arbor, Mi 48106 USA 30-32 Mortimer Street Dept PR London WIN 7RA England BINDING W'hy 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 and expert in the re-binding and repairing of all books. 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Bramley do Doncaster Museum Chequer Road Doncaster DN1 2AE THE rumiMusr U Quarterly Journal of Natural History for the North of England Edited by M. R. D. SEAWARD, MSc, PhD, DSc, FLS, The University, Bradford VOLUME 109 1984 PUBLISHED BY THE YORKSHIRE MORALISTS' WION 3 THE OCCURRENCE OF LITTORINA SPP. IN ROBIN HOOD S BAY, NORTH YORKSHIRE, WITH A KEY TO BRITISH SPECIES I.W JARDINE Department of Pure & Applied Zoology, University of Leeds, Leeds LS2 9JT Introduction The winkles (genus Littorina) are among the commonest and most familiar snails of our rocky shores, yet for the past fifteen years species indentification has been complicated by extensive taxonomic reorganisation of some species groups. There is no guarantee that further alterations will not be made, but, at the moment, a reasonable consensus exists as to the number of British species. Of the eight species now thought to occur in Britain seven are to be found in Robin Hood's Bay (Fig. 1). The purpose of this paper is to outline the changes in taxonomy which have occurred, to provide a key to the British species and to describe the varieties of these found in Robin Hood’s Bay with some notes on their natural history. Since most of the previous work was carried out in S. W. England and Wales, I felt observations from Yorkshire would be of interest, and of more direct use to local naturalists. Fig. 1. Map of Robin Hood’s Bay, N. Yorkshire: bold line giving the position of the cliff edge and fainter line illustrating the level of low water (spring tides). Locations: a) North Cheek, b) site of boulder populations of L. rudis, c) sea wall of former marine laboratory, d) South Cheek below Ravenscar, v) Robin Hood's Bay village. N represents due north. Naturalist 109 (1984) 4 The Occurrence of Littorinaspp . in Robin Hood’s Bay, North Yorkshire Taxonomic revisions 1966-1983 In the fifty years up to 1966 only four species of Littorina were recognised on our coasts; L. littorea (L.), L. neritoides (L.), L. littoralis (L.) and L. saxatilis (Olivi). In 1966 Sacchi and Rastelli revised the taxonomy of L. littoralis establishing two British species; L. obtusata (L.) and a new species L. mariae. Subsequent work has served to support this division (Goodwin and Fish, 1977) and no attempt to split these species further has gained much credence. James ( 1968) considered L. saxatilis to consist of six subspecies with twelve varieties, sparking off a decade of controversy in which it became clear that this ‘species’ was, in fact, an aggregate of several species. Heller (1975) divided in into four; L. rudis (Maton), L. neglecta Bean, L. nigrolineala Gray and L. patula Thorpe. Later, evidence was presented to show that L. patula was synonymous with L. rudis (Raffaelli 1979, Hannaford Ellis 1979), but Hannaford Ellis (1978, 1979) described a new species within this complex, L. arcana. It is probable that L. rudis should revert to the older synonym L. saxatilis unless the two can be shown to be separate species as suggested by Smith (1981). This will, however, not help to dispel lingering clouds of confusion. The existence of another species L. tenebrosa (Montagu) has not been adequately demons- trated. For a detailed review of the taxonomy and recent work on this group the reader should consult Fretter and Graham (1980) and Raffaelli (1982). Key The identification of these species is not difficult but may require dissection. Some characters used require familiarity before divisions become clear, although Fig. 2 and Fig. 3 may help initially. The key makes no attempt to reflect phylogeny and is only intended to be used for sexually mature adult snails, as indicated by the presence of a penis in the male and an oviduct below the rectum in the female. L. nigrolineala is not, as yet, recorded from the Yorkshire coast. 1. Shell with pointed spire (Fig. 2 a, d, e, f) 2 Shell without spire, apex flat or nearly so (Fig. 2 b, c) 7 2. Outer lip of aperture meets body whorl at acute angle (Fig. 2 a) 3 Outer lip of aperture rounded, meeting body whorl nearly at a right angle (Fig. 2 d, e, f) .. 4 3. Usually under 6mm in height, shell grey and unpatterned; inside of aperture glossy black or brown; characteristically a flap of periostracum, the thin proteinaceous outer layer of the shell, extends beyond the aperture lip. Tentacles not banded. Typically at the extreme upper fringe of littoral zone on cliffs or large boulders. L. neritoides Adults over 1cm in height, shell grey, black or brown frequently with faint spiral lines, columnar lip of aperture white and outer lip often striped internally. Tentacles banded. Widely distributed on shore, not on cliffs. L. littorea 4. Shell rather globose. Small, being adult at 4mm in height or less. Usually associated with barnacles in the mid and upper shore. Frequently with a black band round the base of the shell. L. neglecta Shell not globose, being more acutely spired. Never mature below 5mm in height 5 5. Shell with distinctive sculpture consisting of many flat-topped ridges around three times the width of the intervening grooves which frequently contain a black pigment. L. nigrolineala Shell often, but not always ridged. Ridges, except on base, pointed and not wider than grooves 6 The Occurrence of Littorina spp. in Robin Hood's Bay, North Yorkshire 5 6. Females possess a brood pouch and brood young. Both sexes have a large patch of pinkish cilia between the end of the gut and the columellar muscle (Fig. 3 a. b). (The variety tenebrosa will also key out here). L.rudis Females possess a large opaque white jelly gland in place of a brood pouch (Fig. 3 c). Ciliated area, if present, extending very little below gut. L. arcana 7. Penis with extended tip and one row of penial glands. In Yorkshire; aperture lip very thick and apex flat (Fig. 2 b). L. mariae Penis with short tip and two to three poorly defined rows of penial glands. In Yorkshire, aperture lip not greatly thickened and apex slightly raised (Fig. 2c). L. obtusata Occurrence in Robin Hood’s Bay 1. L. neritoides (Linnaeus, 1758); ’small periwinkle’ This species is not common due to the friable nature of the cliffs depriving it of its habitat. It does occur in small numbers on the sea walls of the Bay Hotel and former marine laboratory and on the cliffs at North Cheek and Ravenscar. 2. L. littorea (Linnaeus, 1758); ‘edible winkle’ This is a common species of rocky shores in this area where it is extensively collected for food. It occurs over most of the tidal range, sometimes in dense aggregations. Shells of juvenile specimens, below 5 mm in height, are uniform pale grey with coarse ridges but can be distinguished from L. rudis and L. arcana by the slightly concave-sided appearance of the spire, and when larger by the black and white banding of the aperture lip. The adults arc not usually ridged, and on the lower shore can reach shell heights of 3 cm, much larger than any other species. A distinctive orange-red morph of this species occurs at low' frequencies on Yorkshire coasts. 3. L. obtusata (Linnaeus. 1758) and L. mariae Sacchi and Rastclli, 1966; ‘smooth winkles’ Both species are common among fucoid weeds over most of the shore. L. obtusata, however, occurs higher up amongst Fucus vesiculosus and Ascophyllum nodosum whereas L. mariae is found lower down on F. serratus. In the former species the predominant shell morph is dark green and the aperture is not greatly thickened, whereas the latter is mostly of a tesselated brown variety and the shell is greatly thickened giving a comparatively smaller aperture. Some authors use the pigmentation of the ovipositor as an identification character (Goodwin & Fish, 1977) but this has not proved useful for Yorkshire specimens (J. Grahame, pers. comm.). 4. L. neglecta Bean, 1844 Distinguished mainly by its small adult size, this species is abundant amongst the barnacle Semibalanusbalanoides on the mid and upper shore, often inhabiting empty barnacle shells, but it is rare on sea walls and cliffs. The shells can be plain black, banded or tesselated. The females are ovoviviparous and possess brood pouches. 5. L. rudis (Maton. 1797) and L. arcana Hannaford Ellis. 1978; ‘rough winkles'. L. rudis is not a common species within the bay, being abundant only amongst boulders at Ravenscar. where it co-exists with L. arcana. Small populations also exist amongst boulders at the base of cliffs to the north of the village where L. arcana is extremely rare. Some of these boulder populations have purplish-grey shells but others have orange and brown striped shells, a variety not shown by L. arcana in this area. Populations of rough winkles on the sea walls of the Bay Hotel and former marine laboratory have been found to consist entirely of L. arcana. These animals have thinner shells with wider apertures than the boulder L. rudis but where the two species co-occur at Ravenscar the shells are indistinguishable (Fig. 2 d. e. f). L. rudis broods young while L. arcana lays egg-masses. 6 The Occurrence of Littorina spp. in Robin Hood’s Bay, North Yorkshire .Q o 2 X C/3 0J° - C 1.1 - s °'C E 3 g 2 CL fc 03 . y* o ^ “5 o: « C c o C3 C3 U OS O E E P P 5 o « E C/3 3 X ^ 6 ^ L- 5 "O — d> o x: °f .E TD O 3 ; l_ is £ 03 3 *3 • O _c 'X £ >■* G/j o o 03 — _c CQ o x X — * a X X) C U- Z c s E -5 — o o w u. ■z: w 03 > T3 — c o ^ s i u. CJ *“< Cl ~ C -a '5 3A length of metatarsus. Hairs beneath basal half of hind femur about 3 x length of hairs on anterior face. 10-14 postero-dorsals on hind tibia, with only those on middle third a little robust. wings: Mean length 1.63 mm (range 1.53-1.73 mm). Costal index 0.43-0.46. Costal ratios 3 09 — 4.39: 1 . 18—1 .67: 1 . Costal cilia 0.10-0.13. Wing membrane yellowish grey. Veins pale brownish. Sc fades out just before Rl. A minute hair at base of vein 3. Axillary ridge with two bristles which are stronger than costal cilia. Haltere with yellow knob and darker stem. Naturalist 109 (1984) 12 Six Species 0/ Megaselia ( Diptera , Phoridae) from Northern England, New to Britain female: Similar to male. Proboscis with broader labrum with convex lateral margins. Labellae almost devoid of spines below. Hairs on segments 3-6 of abdominal venter. Tergites 5-8 as in Fig. 3. Costal Index 0.46. Costal ratios 3.44:1.64:1. Costal cilia 0.12 mm. affinities: In the keys of Lundbeck (1922) M. clemonsi will run to couplets 52 in Group VI or 39 in Group VII, depending on whether the costal index is above or below 0.44. Several species described since 1922 also key to these points, with the result that considerable confusion now exists. The species which need to be distinguished from M. clemonsi are M. abdita Schmitz, M. 1 FIGURE 1 Megaselia clemonsi n. sp. hypopygium of male viewed from left side (scale line = 0.1 mm). FIGURE 2 Megaselia hilaris hypopygium of male viewed from left side (scale line = 0.1 mm). Six Species o/ Megaselia ( Diptera , Phoriciae) from Northern England, New to Britain 13 bifida Disney, M. bovista (Gimmerthal) (= M. exigua Wood), M. hilaris Schmitz, M. latior Schmitz, M. pullipalpis Colyer, and M. sylvatica (Wood). M. bifida is immediately distinguished, in both sexes, from the rest of the complex by the presence of bifid spines in the distal comb of the posterior face of the hind tibia (Fig. 8 in Disney, 1983b). M. pullipalpis has the costal index clearly shorter than 0.4 and the male’s anal tube is distinctly shorter than in M. clemonsi. M. latior and M. sylvatica have been confused with each other in the past (see below) but are readily distinguished, in both sexes, from the rest of the complex by having the middle notopleural bristle virtually reduced to a hair. In effect these two species only have 2 notopleural bristles each side. All the other species have 3. M. abdita was described on the basis of a series of females only, from Afghanistan (Schmitz. 1959). However, Dr M. Baez has sent me a series of males and a female from Tenerife. The female agreed with Schmitz’s description except in one particular. Schmitz wrote of the FIGURE 3 Megaselia clemonsi n. sp. female abdominal tergites 5-8 (scale line = 0.1 mm). FIGURE 4 Megaselia sylvatica female abdominal tergites 5 and 6, and posterior part of 4 (scale line = 0.1 mm). 14 Six Species o/Megaselia ( Diptera , Phoridae) from Northern England, New to Britain abdominal venter ‘der ganz unbehaarte Bauch’. Dr Baez’s specimen has a median band of short hairs in a dense field on segments 3-6. Through the co-operation of Dr H. Ulrich I have examined two paratypes of M. abdita and in both the same field of hairs is present. However these hairs can be readily overlooked if the specimens are not viewed from the correct angle. It is clear that Dr Baez’s female is indistinguishable from M. abdita, so that the males from Tenerife represent the undescribed male of this species. The male of M. abdita and M. bovista can be distinguished from M. clemonsi and M. hilaris by their simple labellae bearing few spines below. M. hilaris, like M. clemonsi, has densely-spinose labellae. The two species can be separated by comparing the hypopygia (Figs. 1 and 2), and in particular by the lack of hairs on the dorsal half of the epandrium in M. hilaris. The females of M. abdita are immediately distinguished from those of M. bovista and M. clemonsi by the closely crowded hairs, in several rows on each segment, on segments 3-6 of the venter. M. bovista is distinguished from M. clemonsi by having all the legs dark and by more hairs (90-110) on the frons. I have seen no authenticated female of M. hilaris (ie, a female obtained in cop., in a reared series, or strongly associated in terms of the circumstances of collection). Xlegaselia differens Schmitz, 1948b In reporting an undetermined species of Megaselia from Tow Hill Nature Reserve, North Yorkshire (Grid ref. 34/8286) as * Megaselia sp 2’ (Disney, 1979) it was noted that ‘it somewhat resembles M. differens a species only known from Austria. Through the co-operation of Dr Ulrich I have examined a specimen of M. differens from the Schmitz collection. It is indistinguishable from the Tow Hill specimen. I have a second British specimen collected by Dr A. G. Irwin at the Bridge of Brown (Grid ref. 38/1121), 15 June 1982. The Tow Hill specimen was collected 18 June 1976. These represent the first British records of this species, which is keyed and described by Schmitz and Beyer (1965, 1974). Megaselia drakei n. sp. (Fig. 5) Type locality England: Ferry House, Lake Windermere, Cumbria Type material Holotype: cf, Ferry House, Lake Windermere, Cumbria (Grid ref. 34/3995) 31 October/6 November 1981, leg. C. M. Drake. Paratypes: 2 cf, Tarn House lawn, Malham Tarn, North Yorkshire (Grid ref. 34/894672) 20/22 September 1980 and 25/26 September 1980, R. H. L. Disney; 1 cf, Flatford Mill, Suffolk (Grid ref. 62/079330) 15/17 August 1981, R. H. L. Disney. Etymology The species is named after C. M. Drake, who collected the holotype specimen. Description Only male known: head: Frons brown with 50-70 hairs. Antero-lateral, antial and upper supra-antennal bristles forming a regularly curved row, with the antials slightly nearer the antero-laterals. Lower SA’s as strongly developed as uppers, which are about as far apart as pre-ocellars. Antenna and arista dark. Palps dirty yellow with a field of shallow pits bearing more erect hairs on the external face, with 5-6 bristles, of variable length, distally and a long bristle on inner face. Labrum brownish with convex sides and tip of epipharynx darkened. Labellae a little enlarged but with only a few scattered spines below. thorax: Dark. Scutellum with a posterior pair of bristles and an anterior pair of hairs, which are as fine as (or finer than) hairs at rear of scutum. Three well developed notopleural bristles. Mesopleuron with 1-6 hairs (usually 3 or more). FIGURE 5 Megaselia drakei n. sp. hypopygium of male viewed from left side (scale line = 0.1 mm). abdomen: Tergites dark brown with shortish hairs mainly in distal halves. Venter greyish with a few hairs on segments 3-6. Hypopygium as Fig. 5, with epandrium and hypandrium brown to dark brown and anal tube dusky. The process of the left side of the hypandrium is an almost parallel-sided, rod-like structure. legs: All legs dark brown to blackish. Front tarsi w'ith hair palisades on segments 1-4. Mid-tibia with hair palisade extending 0.1 mm. The distinctive hypopygium will immediately distinguish it from other species running to these couplets. Megaselia eisfelderae Schmitz, 1948a (Fig. 6) This species can easily be confused with M. lutea (Meigen). Males are readily separated by examination of the mid-tarsi. In males of M. lutea the last segment is about 2 x length of segment 4, and usually somewhat inflated (Fig. 7). In M. eisfelderae the last segment is normal (Fig. 6). I have a series of males and a female, collected along with the common M. lutea, in mature fen carr at Malham Tarn, North Yorkshire (Grid ref. 34/888672) in August 1983. These specimens represent the first British records. M. eisfelderae has previously been recorded from Austria, Germany, Holland, Finland, and the Nearctic Region. 16 Six Species o/ Megaselia (Diptera, Phoridae) from Northern England , New to Britain FIGURE 6 Megaselia eisfelderae tarsal segments 3-5 of middle leg in male (scale line = 0.1 mm). FIGURE 7 Megaselia lutea tarsal segments 3-5 of middle leg in male (scale line = 0.1 mm). Megaselia latior Schmitz, 1936 (Fig. 8) sylvatica auctt. nee (Wood, 1910) Desig. nov. Through the co-operation of Dr H. Ulrich and B. H. Cogan I have remounted cotypes of M. latior and M. sylvatica. This revealed that the species that has been attributed to M. sylvatica in Britain by all workers since Wood is in fact M. latior. M. sylvatica has proved to be a common species which has long been known under a synonym. The details and distinction between the species are given below (under M. sylvatica). Apart from recording M. latior in August 1983 at Malham Tarn, North Yorkshire (Grid ref 34/895672) I have specimens from Berkshire, Cambridgeshire, Essex, Rotherham, Suffolk, and Norfolk, the latter having been attributed to ‘M. sylvatica’ by Disney and Evans (1979). Wood’s female specimen, erroneously attributed by M. sylvatica (see below), came from Tarrington Hereford. M. latior represents an addition to the British List. It has previously been recorded from Belgium. Megaselia melanostola Schmitz, 1942 This species will run to couplets 5, 11 or 13 in Group VI of Lundbeck’s (1922) keys. The elongated anal tube (Abb. lc in Schmitz, 1942) will distinguish it from the other species running to these couplets. A male from Rowantree Scar (Grid ref. 44/032932) 20 June 1978 and another from Tranmire (Grid ref. 45/765118) 14 September 1978 were both included in the category ‘ Megaselia spp’ by Disney etal (1981). I have also 3 males and a female collected by J. M. Nelson at Flanders Moss, Perthshire (Grid ref. 26/623976) June 1981 and 27 May/2 June 1982. This species is an addition to the British List, having been reported previously from Austria, Germany and Sweden. FIGURE 8 Megaselia latior hypopygium of male viewed from left side (scale line = 0.1 mm). FIGURE 9 Megaselia sylvatica hypopygium of male viewed from left side (scale line = 0.1 mm). Megaselia sylvatica (Wood, 1910) (Figs. 4 and 9) impolluta (Schmitz, 1920) Syn. nov. Schmitz (1939), subsequent to his original description of M. impolluta , pointed out that the female was very distinctive in that the sixth abdominal tergite was abbreviated, and a patch of long hairs was developed behind the tergite (Fig. 4). 18 Six Species o/Megaselia (Diptera, Phoridae) from Northern England, New to Britain According to a label attached to Wood’s type series of M. sylvatica the specimens were examined by Schmitz in 1926. I have also examined the same type series and have, through the co-operation of B. H. Cogan, remounted the single female and a male. I have designated the latter the lectotype. This lectotype is the same species as males of M. impolluta’ obtained in reared series (Disney and Evans, 1979, 1982), whose females are as in Fig. 4. It is clear, therefore, that M. impolluta is a synonym of M. sylvatica. The female in Wood’s series is not M. sylvatica. It is M. latior. It would seem that Schmitz failed to appreciate this incorrect association and thus concluded that he had been right to establish M. impolluta as distinct from M. sylvatica. I have M. latior females obtained in reared series, and erroneously attributed to M. sylvatica (see above), through following Schmitz. The two species are readily separated in the female sex by the form of abdominal tergite 6. In M. sylvatica it is as in Fig. 4. In M. latior it is subequal to, or longer than, tergite 5. The males are distinguished by the hypopygia. in particular M. latior has the anterior part of the penis complex developed into an elaborate expanded structure that protrudes even when the penis is withdrawn (cf. Figs. 8 and 9). M. sylvatica is widely distributed in Britain. Acknowledgements I am grateful to B. H. Cogan (British Museum, Natural History) and to Dr H. Ulrich (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn) for allowing me to re-mount critical type material. I thank the Shell International Petroleum Co. Ltd. for a further grant to aid my study of Phoridae. References Disney, R. H. L. (1979). Some scuttle flies (Diptera, Phoridae) from Tow Hill Nature Reserve, North Yorkshire. Naturalist, Hull. 104: 39-42. Disney, R. H. L. (1983a). Scuttle flies — Diptera, Phoridae (except Megaselia). Handbk. Ident. Br. Insects. 10(6): 1-81. Disney, R. H. L. (1983b). A useful new character in the giant genus Megaselia (Diptera: Phoridae), w'ith two new species from Britain. Z. ang. Zool. 70: 225-234. Disney, R. H. L., Coulson, J. C. and Butterfield, J. (1981). A survey of the scuttle flies (Diptera: Phoridae) of upland habitats in Northern England. Naturalist. 106: 53-66. Disney, R. H. L. and Evans, R. E. (1979). Further records of Phoridae (Dipt) reared from fungi. Entomologist's mon. Mag. 114(1978): 166. Disney, R. H. L. and Evans, R. L. (1982). Records of Phoridae (Diptera) reared from fungi. Entomologist's Rec. J. Var. 94: 104-105. Lundbeck, W. (1922). Diptera Danica. Part VI Pipunculidae Phoridae. Gad, Copenhagen. Schmitz, H. (1920). Drei neue europaische Phoriden (Dipt). Ent. Meddr. 13: 115-119. Schmitz. H. (1936). Vier neue europaische Phoriden. Konowia. 15: 190-195. Schmitz, H. (1939). Die ersten hundert Phoriden von Portugal (Fortsetzung). Broteria. 35: 180-193. Schmitz, H. (1942). Drei neue europaische Phoriden. Natuurh. Maandbl. 31: 10-12. Schmitz. H. (1948a). Zur Kenntnis der fungicolen Buckelfliegen (Phoridae, Diptera). Natuurh. Maandbl. 37: 37-44. Schmitz, H. (1948b). Zweiter Beitrag zur Kenntnis der Phoriden osterrichs (Diptera). Ann. natuurh. Mus. Wien. 56: 375-399. Schmitz, H. (1959). Contribution a l’etude de la faune d’Afghanistan 19 — Phoridae. Broteria. 55: 119-130. Schmitz, H. and Beyer, E. (1965), in Lindner, E. (ed). Die Fliegen der Palaearktischen Region 33. Phoridae Lief. 258, 260: 513-608. Schmitz, H. and Beyer, E. (1974). Ibid. Lief. 301: 609-637. Schmitz, H. and Dclage, A. (1974). Ibid. Lief. 301: 638-664. Wood, J. H. (1910). On the British species of Phora (contd). Entomologist’s mon. Mag. 46: 149-154, 195-202. 243-249. 19 THE VASCULAR FLORA OF DISUSED CHALK PITS AND QUARRIES IN THE YORKSHIRE WOLDS RICHARD G. JEFFERSON Department of Biology, University of York, Heslinglon, York Introduction The Yorkshire Wolds form the northernmost section of the English chalk of Upper Cretaceous age, extending from the River Humber in an arc to the sea at Flamborough Head, covering an area of approximately 775 km2. The Wolds have been extensively quarried for chalk and many workings have been abandoned, only thirteen working quarries remaining in 1983. The abandoned quarries and pits range in size from 0.05 to 10 ha. The small chalk pits were excavated primarily as a local source of building stone and aggregate whereas the larger pits and quarries were chiefly producing chalk for agricultural lime used locally (Hull and Thomas, 1974, Allison. 1976). Current working quarries produce chalk for use in paint whiting and cement manufacture. Many of the smallest disused pits have been infilled or ploughed out and some larger quarries are used as landfill sites. It is well known that some disused chalk and limestone quarries in the British Isles are important botanical sites (Davis, 1979) and the importance of such sites in the Wolds has been recognized by the Yorkshire Naturalists’ Trust, with four quarries and pits being protected as Nature reserves or management agreement sites; two of these (Wharram Quarry and Kiplingcotes chalk pit) are also Sites of Special Scientific Interest. Ratcliffe (1982) also emphasizes the botanical interest of disused Wolds quarries along the Wolds Way long distance footpath. This paper aims to show that other disused pits and quarries in the Yorkshire Wolds are of botanical conservation interest. Sample sites and methods A total of 227 quarries originally located from aerial photographs and 1:10000 Ordnance Survey Maps were visited between November 1981 and February 1982; thirty of these w'ere subsequently selected for botanical survey. During June and July 1982, a total list of vascular plants was compiled for each of these sites as part of a more detailed ecological survey, the results of which are reported elsewhere (Jefferson, in press). Additional plant records were added, where necessary, from observations made in May and June 1983. Table 1 gives details of the sites surveyed. Results and discussion The survey recorded 256 species of vascular plant. Table 2 lists some of the more interesting species recorded and the number of sites at which they occurred. This list includes species restricted both locally and nationally, some typical calcicoles and naturalized and introduced species. A number of species of the genus Hieracium (excluding H. pilosella) were recorded from seven sites; unfortunately identification to species level was not possible. Jefferson (in press) showed that a range of plant community types occurred in Wolds quarries including early successional communities with discontinuous vegetation cover on quarry floors, spoil heaps and chalk scree through to later successional tall grassland communities with some scrub invasion, occurring on quarry floors and side slopes with deeper soil. Clearly there is an age component influencing the plant communities and the overall species composition. On a smaller scale. Usher (1979) demonstrated the effects of age on plant community structure by looking at parts of the quarry floor at Wharram which had been successively abandoned over a thirty-year period. Botanical diversity decreased through the succession from the open dicotyledon stage to the grassland stage. Additionally. Jefferson (in press) showed that site area and distance of a quarry from semi-natural chalk grassland can have an important influence on species composition and species richness. Other factors such as substrate characteristics and extent of grazing can also be important in quarries generally (Humphries. 1980). The quarry flora includes pioneer colonists of bare chalk (for example, Desmazeria rigida, Hieracium pilosella). many species typical of chalk grassland (for example. Carlina vulgaris, Naturalist 109 (1984) 20 The Vascular Flora of Disused Chalk Pits and Quarries in the Yorkshire Wolds TABLE 1 Site Details Chalk Quarry/Pit Grid reference Cartographic Area (ha) Total number of species Rifle Butts SE 898427 0.25 91 Etton SE 970434 1.12 89 Millington SE 831535 0.09 55 Bishop Wilton SE 805559 1.15 73 Staveley (A) SE 913501 0.05 41 Staveley (B) SE 918501 0.06 39 Pilmoor TA 091788 0.53 68 Whitegate SE 958756 0.63 86 Poor Wood SE 829509 0.22 58 Thirkleby SE 915698 0.38 68 Burrow House SE 983668 0.29 54 Wharram SE 859653 6.22 111 Eppleworth TA 021324 1.30 93 Nunburnholme SE 857478 0.25 64 Burdale SE 872626 2.83 108 Middleton SE 955499 0.29 40 Kipiingcotes SE 915435 1.80 90 North Grimston SE 857670 0.21 84 Aldro SE 813626 0.28 68 Helperthorpe SE 960705 0.42 57 Low Mowthorpe SE 885659 0.25 67 Flixton TA 039793 0.96 93 North Dalton SE 942503 0.40 80 Linghall SE 917727 0.08 62 Muston TA 087798 0.09 55 Humberfield TA 012264 3.20 81 Pefham SE 858578 0.10 22 Staxton TA 011783 0.66 91 High Mowthorpe SE 891699 0.08 45 Sledmere SE 921634 0.33 76 Thymus praecox), arable and pasture weed species (for example, Stellaria media , Cirsium arvense, Senecio jacobea) and naturalized plants such as Centranthus ruber and Buddleja davidii. Robinson (1902) mentions the presence of a number of naturalized and casual species occurring in quarries at Hessle and Willerby on Humberside including Diplotaxis muralis and Antirrhinum majus. The species list (Table 2) contains no national rarities but a number of species such as Astragalus danicus and Cirsium eriophorum are restricted nationally (Perring and Walters, 1976) and C. acaule, Ophrys apifera and Blackstonia perfoliata are rare in the North of England being near the northern limit of their range in Britain. Crackles (1974) states that the latter two species show a preference for open habitats in the North of England where competition pressure is less. This is partly confirmed by Julian (1983) who showed that at Wharram, O. apifera is restricted to the open quarry floor where the soil is approximately 5 cm deep and of low nutrient status. As soon as the soil has developed sufficiently to support a closed community of tall grasses (Arrhenatherum elatius , Dactylis glomerata) and herbs ( Centaurea nigra, Lathyrus pratensis) O. apifera is outcompeted and excluded. The Wolds is predominantly an area of intensive arable farming where extensive areas of semi-natural chalk and neutral grassland are now restricted to the dry valley systems. In the last twenty years an estimated 43 per cent of this grassland has been changed to arable, improved 21 The Vascular Flora of Disused Chalk Pits and Quarries in the Yorkshire Wolds TABLE 2 A partial list of vascular plant species recorded from chalk pits and quarries in the Yorkshire Wolds during 1982/83. Acinos arvensis 1 XErucastrum gallicum 1 *Amsinkia intermedia 1 Euphorbia exigua 1 Anacamptis pyramidalis 8 Festuca gigantea 1 Anthyllis vulneraria 6 Filipendula vulgaris 1 Astragalus danicus 1 Galium album 2 Arabis hirsuta 6 Gentianella amarella 7 Avenula pratensis 4 XGeranium sanguineum 1 Blackstonia perfoliata 1 Gymnadenia conopsea 1 Brachypodium pinnatum 9 Helianthemum nummularium 6 B. sylvaticum 1 Hypericum hirsutum 3 Bromus e rectus 8 Koeleria macrantha 7 B. ramosus 1 XLathyrus tuberosus 1 * Buddleja davidii 1 XLinaria repens 1 Campanula glomerata 10 Listera ovata 2 tC. latifolia 1 Malva moschata 1 Carex sylvatica 1 *Melilotus officinalis 2 Carlina vulgaris 5 Mycelis muralis 4 Centaurium erythraea 3 Onobrychis viciifolia 1 *Centranthus ruber 1 Ophioglossum vulgatum 1 Cerastium arvense 2 Ophrys apifera 2 Chaenorhinum minus 2 Origanum vulgare 7 Cirsium acaule 1 Picris hieracioides 4 C. eriophorum 4 Poly gala vulgaris 5 Cornus sanguinea 1 Prunus avium 1 iCoronilla varia 1 Reseda lutea 5 Coronopus squamatus 1 Sagina nodosa 1 Dactylorhiza fuchsii 9 Sanguisorba minor 15 Daucus carota 4 Scabiosa columbaria 8 Desmazeria rigida 7 Sedum acre 1 *Diplotaxis muralis 1 Senecio erucifolius 1 Echium vulgare 1 S. viscosus 1 Erigeron acer 1 Stachys arvensis 1 Eriophila verna * Naturalized species 1 Thymus praecox 10 t Introduced species $ Casual species pasture and forestry, and in some cases cessation of sheep grazing has resulted in extensive scrub development (Rafe and Jefferson. 1983). Disused quarries therefore provide refuges for chalk grassland species declining due to habitat change as well as for early successional species characteristic of open ground on calcareous soil. It is therefore important that some of the botanically interesting disused quarries and pits on the Wolds are retained for conservation and research. Many sites will, however, require some form of management to prevent the succession proceeding to a less diverse community dominated by tall grasses such as A. elatius. Management has already been implemented at Wharram and the rationale of this is discussed in the reserve management plan (Anon. 1973). Acknowledgements I am grateful to the Yorkshire Naturalists' Trust. Humberside County Council. Birdsall Estates Ltd. and numerous farmers and landowners who gave me permission to work on their land. I 22 The Vascular Flora of Disused Chalk Pits and Quarries in the Yorkshire Wolds thank Dr M. B. Usher for commenting on an earlier draft of this paper and Dr R. W. Rafe for useful discussion. Finally, I acknowledge the financial support of the Natural Environment Research Council. References Allison, K. J. (1976). The making of the English landscape. The East Riding of Yorkshire landscape. Hodder & Stoughton, London. Anon (1973). Wharram Quarry Nature reserve management plan from 1 January 1979 to 31 December 1983. Duplicated report. Yorkshire Naturalists’ Trust Ltd. Crackles, E. (1974). Seeking to understand the flora of the East Riding of Yorkshire. Naturalist. 99: 1-17. Davis, B. N. K. (1979). Chalk and limestone quarries as wildlife habitats. Minerals and the Environment. 1: 48-56. Hull, J. H. and Thomas, I. A. (1974). Limestones and dolomites. In The geology and mineral resources of Yorkshire. (D. H. Rayner and J. E. Hemingway, ed.): 345-359. Yorkshire Geological Society, Leeds. Humphries, R. N. (1980). The development of wildlife interest in limestone quarries. Rec. Rev. 3: 197-207. Jefferson, R. G. (in press). Quarries and wildlife conservation in the Yorkshire Wolds, England. Biol. Conserv. Julian, S. J. (1983). The autecology of Ophrys apifera L. through the study of its distribution in Wharram Quarry. BSc Honours project, University of York. Perring, F. H. and Walters, S. M. (1976). Atlas of the British flora. EP publishing, Wakefield. Rafe, R. W. and Jefferson, R. G. (1983). The status of Melanargia galathea (Lepidoptera: Satyridae) on the Yorkshire Wolds. Naturalist. 108: 3-7. Ratcliffe, R. (1982). The Wolds Way. Long distance footpath guide number 12. HMSO, London. Robinson, J. F. (1902). The flora of the East Riding of Yorkshire. Brown, London. Usher, M. B. (1979). Natural communities of plants and animals in disused quarries. J. Envir. Manage., 8: 223-236. BOOK REVIEW Distribution Atlas of Woodlice in Ireland by Declan Doogue and Paul T. Harding. An Foras Forbartha, St Martin’s House, Waterloo Road, Dublin 4, Ireland. 1982. £2.50, including postage. The main part of this work is a series of maps, one for each of the species recorded in Ireland, with text on a facing page. There is a Preface and an Introduction, followed by a Check List of the Irish species with a further list of species absent from Ireland but occurring in Britain. A section on the history of recording and the growth of the Irish species list through time gives a valuable insight into the origins of the present work. This is followed by sections on the operation of the Woodlouse Recording Scheme, the storage of records, identification and factors affecting distribution. The text accompanying the maps gives a very brief description of identifying features, habitat preferences, and overall distribution. Each account ends with the number of vice-counties from which the species has been recorded and a key to those references in the Bibliography mentioning the species. All in all this is far more than a simple atlas. It contains a wealth of information about the Irish Woodlice (and indirectly about the British ones too) and with 146 references, leaves no stone unturned in the exposure of their past investigation. The remarkable thing is that this has been achieved by a tiny band of dedicated people who between them have covered just about every 10 km square in Ireland. They deserve our congratulations and thanks. The Atlas is a worthy monument to their achievement. SLS 23 THE SOLITARY BEES AND WASPS ( HYMENOPTERA : ACULEATA) OF A SAND-PIT AT SWINCARR PLANTATION, NEAR YORK MICHAEL E. ARCHER 17 Elmfield Terrace , Malton Road , York Y03 OEH The sand-pit (SE 6856) was found on 21 August 1967 and its solitary bees and wasps were studied from April to July 1968 and from March to September 1969. Access to the site was then prevented and the pit has now been filled. The sand-pit (Fig. 1), which was about 22 metres x 22 metres, had standing water at its centre surrounded by an area of damp sand and then of dry sand. The extent of the water and damp sand varied during the year. No vegetation was present in the pit, while the path and an area immediately to the left of the pit was largely bare of vegetation. There were thirty-five species of solitary bees and wasps recorded, some of which were found nesting in very large numbers. The three most numerous species w'ere Andrena clarkella (Kirby) which in 1968 was represented by over 900 freshly-dug burrows, Mellinus arvensis (Linn.) in 1969 by about 140 burrows and A. barbilabris (Kirby) where a burrow count was not possible but probably approached that of A. clarkella. Males of A. clarkella emerged in early April closely followed by the females. After a period of mating the males disappeared by the middle of April or a little later. The females dug the burrows and provisioned the cells but had disappeared by the middle of May. There was one generation each year. On emerging, the males walked and flew over these sunlit banks and flat areas waiting for the females to emerge. Males approached and pounced on any small dark object, including inaminate objects and other males, but left immediately after making contact if the object was not a female. The male grasped the female with fore and middle pairs of legs around the waist and rear thorax and extruded his genitalia towards the female opening. Females usually struggled so the two bees separated and no successful copulations were seen. Males were seen to chase females in flight even if carrying pollen. The females dug their burrows in the flat area mainly to the left of the sand-pit (Fig. 1). The nesting area had a stable sand surface and was moss-covered but with intervening bare patches. In 1969 the left area contained 354 freshly-dug burrows but in 1968 a sample area of 33.5 square metres contained 308 burrows: the estimate for the left area would be therefore about 900 burrows. The female dug the burrow with her forelegs, excavated sand being pushed backwards by the middle and hind legs. As the burrow became deeper, the sand was first brought to the surface and then moved horizontally away from the entrance so that a mound of about 7 cm diameter was produced. On flat ground, the entrance of the burrow was in the centre of the circular mound, while on sloping ground the mound became oval in shape, with the entrance at the upper edge. The presence of the mound enabled one to separate freshly-dug burrows from emergence holes. At first the entrance might be closed or open, but as soon as the female started to collect pollen the entrance was kept closed. Thus, on arriving with a pollen load on the hind legs the female dug through the entrance plug with her forelegs, sometimes assisted by the middle legs. As the female disappeared into the burrow, the sand collapsed behind her, filling the entrance. Similarly, on leaving, the sand collapsed behind the female, but she also moved sand from the edges of the mound to the entrance with her hind legs, which were held at right-angles to her body. First departures from the freshly-dug burrows were accompanied by walking around the mound followed by flying around the nesting area in order to learn the locality of the burrow. The female then proceeded to stock her cells with pollen and nectar. Some burrows were excavated to determine their structure: the main burrow on the flat area went straight down for about 11.5 cm. having a diameter of about 1 cm, with an entrance plug about 0.6 cm in depth. About six cells were present per burrow, either directly attached to the main burrow or with a short side-burrow. The cells were separate from each other, with external dimensions of about 1.6 cm length and 0.8 cm diameter. The inside walls of each cell were smooth and shiny, except for the plug of soil with which it had been sealed. The larvae, which usually laid in a curved position on the pollen-nectar ball, were capable of very little movement, mainly an up-and-down movement of the head. Naturalist 109 (1984) 24 The Solitary Bees and Wasps of a Sand-pit near York 15 metres Andrena clarkella Andrena barbilabris >*u*+*»** Mellinus arvensis FIGURE 1 Sketch map of sand-pit and path with nesting areas of Andrena clarkella , A. barbilabris and Mellinus arvensis. (P = Path, W = Standing water, boundaries of sand-pit and path.) Fully-grown larvae were found in early May and early June. Nixon (1954) reported the larvae pupate and emerge as adults in the autumn, with the adults over-wintering in the cells. The cleptoparasite of A. clarkella is Nomada leucophthalma (Kirby), which appeared in the spring with its host but disappeared after its host at the end of May. During the phase when the burrows were open female cleptoparasites were seen to enter the burrows. The female Nomada would follow a female Andrena in flight and moved onto the surface of the mound after the female Andrena had dug in. Soon after the middle of April the female Nomada were seen to dig into the burrow mainly with her forelegs but with the middle legs assisting. Bernard (1951) reported that underground the female Nomada laid an egg in the cell of its host. On hatching the Nomada larva would kill the egg or young larva of its host before eating the pollen-nectar ball. The adults of A. barbilabris emerged soon after the middle of May, the males a few days before the females. The males had disappeared by the middle of June and the females by early July. The nesting areas were the bare dry mobile sand areas of the path and the right side of the sand-pit (Fig. 1). At first males were numerous, 20-30 males being seen together either flying near, walking over or stationary on the sand surface, often at the entrances of the emergence holes. The males entered burrows looking for the females and followed the females in flight. The mating posture was similar to that of A. clarkella and again, although many attempted copulations were seen, a successful copulation was not observed. The female would raise her legs dorsally above her body and the pair would separate. Several males were also seen clustered around a female. The female dug a burrow, creating a mound of about 2.5 cm diameter and about 0.6 cm at its highest point. During the provisioning phase the burrows were kept closed. The cleptoparasite of A. barbilabris is Specodes pellucidus (Smith, F.). Only females of Sphecodes emerged a few days before that of their hosts. The female Sphecodes spent much time walking over the sand surface tapping the sand with her antennae, paying particular attention to the mounds. From the middle of June the female Sphecodes started to dig into the mound with The Solitary Bees and Wasps of a Sand-pit near York 25 her forelegs, the middle and hind legs pushing the excavated sand to the side. The female cleptoparasites had disappeared by early July but unlike Nomada both sexes emerged in mid-August to mate. Afterwards the males died and the females over-wintered underground. The adults of Mellinus arvensis emerged in late July to early August, the males up to a week before the females. Males disappeared towards the end of August while some females were still active towards the end of September. M. arvensis is a fly-hunter, mainly stocking its cells with Muscidae (Hamm & Richards, 1930). The nesting area was associated with the banks of the sand-pit (Fig. 1). A count of freshly-dug burrows in 1969 produced an estimate of 144 burrows. Fig. 1 shows that the high-density nesting areas of the above three species do not overlap so that interference between burrow systems was unlikely. Each species was associated with a nesting area of different characteristics, each habitat being presumably selected by the nesting female. The times of appearances of the three species also showed no overlap. Three other species were found nesting. A few burrows of Halictus rubicundus (Christ) were found in the A. clarkella area. Six females were found from early May to the middle of June, one seen entering an open burrow with a pollen load. The cleptoparasite of H. rubicundus is Sphecodes gibbus (Linn.) and it was seen from late July to early August. A few burrows of the second species, Colletes succinctus (Linn.), were found in the vertical bank faces during August. The burrows were left open and its cleptoparasite Epeolus cruciger (Panzer) was seen to follow female Colletes and enter burrows when females were absent. Three small patches of burrows of the third species Psen equestris (Fabricius) were found on sloping ground in the A. clarkella area. In 1969 counts of freshly-dug burrows in the three patches gave 46 burrows (10+27+9). P. equestris was active during July. The small number of individuals and/or small area of nesting site required by these three species was unlikely to have produced much burrow interference with the three main species. The other species recorded are as follows. Females of the wingless scolioid wasp, Myrmosa atra (Panzer), were found from July to early September. Little is knowm of the habits of this cleptoparasite but its hosts are probably ground-nesting sphecoid wasps. Two species of pompiloid wasps, the spider-hunting Priocnemis parvula Dahlbom and the cleptoparasite Evagetes crassicornis (Shuckard) and one solitary vespoid wasp, Ancistrocerus scoticus (Curtis) were also found. The additional sphecoid wasps were the acridid nymph-hunter Tachyspex pompiliformis (Panzer), the fly-hunters Crabro cribrarius (Linn.), C. peltarius (Schreber). Crossocerus quadrimaculatus (Fabricius), Oxybelus uniglumis (Linn.), and the aphid-hunter Diodontus tristis (Vander Linden). The Andrena mining bees included A. denticulata (Kirby) and A. fuscipes (Kirby) with their cleptoparasite Nomada rufipes Fabricius, A. haemorrhoa (Fabricius), A. tarsata Nylander, A. wilkella (Kirby) with its cleptoparasite N. striata Fabricius. The cleptoparasite N. marshamella (Kirby) was also found but its very common host A. jacobi Perkins, R. C. L. was not recorded. Among the halictine bees were Lasioglossum calceatum (Scopoli), L. fratellum (Perez), L. punctatissimum (Schrenck), and L. rufitarse (Zetterstedt). The cleptoparasite Sphecodes fasciatus von Hagens was found in both the female and male sex so its identity is certain. The megachilid bee. Megachile willughbiella (Kirby) was also taken but since this bee is an aerial nester (usually in decayed trees), it must have been just passing over the sand-pit. With the exception of L. fratellum and L. rufitarse all the above species are widely distributed throughout England, although many such as M. atra. T. pompiliformis. C. succinctus , and A. tarsata are local in distribution, being restricted to sandy places. L. fratellum is probably widespread in the north and west of England and L. rufitarse in the north and west midlands areas. References Bernard. F. (1951). Super-famille des Apoidea ou abeille. In Traite de Zoologie (P.-P. Grasse, ed.). 10: 1198-1257. Masson et Cie., Paris. Hamm, A. H. and Richards. O. W. (1930). The biology of the British Fossorial Wasps of the families Mellinidae. Gorytidae, Philanthidae. Oxybelidae. and Trvpoxyidae. Trans, ent. Soc. Lond. 78: 95-131. Nixon, G. (1954). The World of Bees. Hutchinson. London. 26 BOOK REVIEWS Resource Competition and Community Structure by David Tilman. Pp. xi + 296, including many diagrams. Princeton University Press, 1982. £19.40 hardback, £7.05 paperback. Since their first monograph in population biology Princeton has regularly produced excellent innovative texts presenting to ecologists new ideas and new hypotheses to test, and Number 17 by David Tilman is no exception. Against a background of resource utilization and partitioning, he explains, via consumption constraint curves, why there is such a diversity of plants and animals. The ideas are carefully developed in a well written, easily understood text which proceeds through the use of graphical rather than mathematical argument and which examines well-known data such as that from the Rothamsted Park Grass Plots. The ideas presented may not be immediately accepted by all ecologists, but they do provide testable hypotheses for the sceptics to reject by experiment. One may not find some of David Tilman’s statistics entirely satisfactory, but it is this type of text which enables ecology to make progress through the presentation of innovative ideas for others to test. This book should be on the bookshelf of every serious student of ecology who has wondered why rare species exist and why an ultraefficient ‘superspecies’ has never evolved. JEPC The Ecology of Pests: Some Australian Case Histories, edited by R. L. Kitching and R. E. Jones. Pp. viii + 254, with many line drawings, figures and b/w plates. CSIRO, 1981. This is a collection of papers about both plant and animal pests. Many of the examples covered are introduced organisms which have become pests as a result of having no natural predators. Examples of this type are skeleton weed ( Chondrilla juncea) and the Sinex woodwasp ( Sinex noctilio). The authors of these papers discuss how these organisms have become pests and what measures have been taken to control them. The measures used include chemical and biological type controls as well as an integrated approach. The other examples are naturally occurring pests, such as the arid zone kangaroos ( Macropus robustus and Megaleia rufa) and mosquitoes ( Aedes vigilax and Culex annulirostris) . As with most collections of papers written by authors with differing communication skills, the accounts vary tremendously in the clarity of presentation and amount of previous knowledge required to understand clearly the message of the case history. The standard in this case is generally high, but there are one or two of the twelve that let the others down. The coverage of this volume is good and the examples, each with their own lessons for pest control in a wider context, are to be recommended to all who have an interest in the ecology of pests. JEPC Grasses and Grasslands: Systematics and Ecology, edited by James R. Estes, Ronald J. Tyrl and Jere N. Brunken. Pp. 312. University of Oklahoma Press, 1982. $12.50. A collection of eleven papers presented at a symposium at Oklahoma State University in August 1979. The symposium was seen as necessary to make up for a shortage of papers about grass taxonomy and grassland ecology at other symposia. All the papers have an underlying evolutionary perspective. The volume is divided into two parts. Part 1 covers the taxonomic topics with four papers on the Triticeae, one on the Poaceae and one of a rather more general nature. As may be inferred from the families considered in this section, the papers all have an applied aspect to them and most are clearly illustrated with diagrams and tables. Part II contains five papers of a profoundly ecological character. Again, these papers are essentially of an applied character, the interaction between grasslands and grazing animals being a major theme. This volume draws attention to the economic importance of grasses and grasslands and points to a paucity of published material, on a worldwide scale, that views these subjects in their evolutionary context. JEPC LOSS OF WETLANDS IN SPROTBROUGH PARISH, DONCASTER 27 P. F. ULF-HANSEN University of Western Ontario, London, Canada N6A 5B7 Introduction Wildlife habitats are being lost and damaged at an increasing rate, and if this continues, the area of semi-natural vegetation in Britain will be halved within a generation (Goode, 1981). Freshwater systems are particularly vulnerable, with the principal threats being intensive land use, drainage and pollution. There is a need for quantitative data on such destruction and improverishment. Presented here are data for one small area (about 1100 hectares), the parish of Sprotbrough, near Doncaster, South Yorkshire, which was collected as part of an investigation into mobility as a causal factor of rarity in wetland flora (Ulf-Hansen, 1981). Methods A survey of Sprotbrough parish sought to identify the approximate date of formation of wetlands. Ordnance Survey maps from the First Series (1851-52) through to 1971-72 and aerial photographs from 1966-68 to 1978-80 provided a continuous monitor of possible wetland formation. However, it was found that no wetlands were created post- 1890 and the causes and dates of their disappearance were therefore noted; where possible this information forms the basis of results given here. All still-water bodies were counted, ranging from ponds in old parkland to cut-off canals and field ponds. Results and Discussion The loss of wetlands over a period of 117 years from 1850 to 1967 is given in Table 1, which shows extant wetlands and numbers lost over five periods of time. Since the time intervals vary, the percentage rate of loss per annum is given. The table clearly indicates an accelerating rate of loss, reaching 1.23 per cent of sites in the pre- and post-war period to 1967. This represents a total loss of twenty-two wetlands or 65 per cent over the complete period. TABLE 1 Loss of wetlands 1850-1967, variable intervals, Sprotbrough parish Year No. of wetlands extant No. of wetlands lost % rate of loss per annum 1850 34 — — 1890 28 6 0.44 1910 25 3 0.54 1930 22 3 0.60 1967 12 10 1.23 The nature of the loss is illustrated by Fig. 1, which shows the situation in 1850, 1910 and 1967. The fine scatter of sites apparent in 1850 has been markedly reduced, the most recent plot showing only four clusters and a singleton. The trend has been for smaller single wetlands (mostly field ponds) to be eliminated, with the remaining sites being represented by larger water-bodies, often sited in old parkland. It has been possible to establish probable causes of destruction for fourteen of the twenty-two sites: eight were eliminated due to agricultural land use, either by filling in or drainage, four to urban/residential development, and two to road and rail construction. The 65 per cent loss of wetlands is broadly comparable to other studies, although the area sampled here is smaller. Ratcliffe (1977) reports a study of the Leicestershire area, where 37 per cent of ponds have been filled in since 1930. For comparison, over the same time period, this investigation yields a figure of 55 per cent. The higher loss for Sprotbrough may reflect the Naturalist 109 (1984) 28 Loss of Wetlands in Sprotbrough Parish, Doncaster 1850 1910 1967 FIGURE 1 Loss of wetlands, 1850-1967, Sprotbrough parish. Doncaster Loss of Wetlands in Sprotbrough Parish, Doncaster 29 urbanized nature of parts of the parish. Also in the Leicestershire area, the number of field ponds was reported to have decreased by 50 per cent (Beresford, 1981), although no time period was noted. These declines show the heavy pressure on such wetland habitats and evidence suggests that other wildlife habitats are equally under threat. Approximate estimates of habitat losses over the same time span as given above were made from figures presented by Goode (1981). These indicate losses of about 30 per cent for Dorset heathlands. Dorset chalk grasslands, and northern mosslands. Goode (1981) also cites evidence of a 30-50 per cent decline of all ancient, semi-natural deciduous woodland over the last thirty years. Small wetlands are thus equally in danger, and Haslam (1975) in fact considers them to be one of the most threatened British habitats. It is to be hoped that the rate of destruction of these and other valuable habitats slows in the near future. Acknowledgement I would like to thank Ms C. A. Lupson for help and encouragement. References Beresford, J. E. (1980). The aquatic macrophyte flora of field ponds: their conservation value and distribution. Paper presented at British Ecological Society Winter Meeting. Goode, D. (1981). The threat to wildlife habitats. New Scientist. 89: 219-223. Haslam, S. M. (1975). The management of British wetlands. II. Conservation. J. Environ. Mgmt, 1: 345-361. Ratcliffe, D. A., ed. (1977). A Nature Conservation Review. Vol 1. Cambridge University Press. Cambridge. Ulf-Hansen, P. F. (1981). Species mobility as a causal factor of rarity: a study of the colonization of wetlands by macrophytes. BTech dissertation. University of Bradford. ENTOMOLOGICAL REPORTS FOR 1981-82 R CROSSLEY Hemiptera In addition to recording notable discoveries made in the two years under review, this report includes a number relating to earlier periods which were overlooked previously or have only recently come to hand. Of particular interest is Mr Foster's discovery of the mirid bug Brachyarthrum limitatum Fieb. , which is another addition to the southern element of the insect fauna of the Doncaster area, not having been found previously further north than Suffolk. The recognition of the variety lateralis (Hahn) of Polymerus unifasciatus (F.) in 1979 was of national significance and full details have appeared elsewhere (Crossley. R. (1981) Polymerus unifasciatus (F.) var. lateralis (Hahn) (Hem. Miridae) in Britain. Entomologist's mon. Mag. 116(1980): 155). It is probable that all northern specimens of P. unifasciatus are of the variety lateralis , the normal form being confined to southern Britain. Records have been received from Messrs W. A. Ely, J. H. Flint. S. Foster, D. Horsfield. B. S. Nau, T. Potter, to all of whom I express my thanks. In the list which follows new county records are indicated thus t and new vice-county records thus *. Heteroptera * Macrodema micropterum (Curt.) (61) Allerthorpe Common. 7/7/82; S.F. *Plinthisus brevipennis (Lat.) (63) Sandall Beat, Doncaster. 12/6/82; S.F. Barnby Dunn, 23/7/82; S.F. Only previous Yorks, record. Cloughton (VC 62), 1924. t Brachyarthrum limitatum Fieb. (63) Wadworth Wood. Doncaster. 26/6/76: S.F. *Chlamydatus pullus (Reut.) (63) Balbv. Doncaster. 20/8/76: S.F. Only previous Yorks, record. Richmond (VC 65), 1927. 30 Book Reviews *C. saltitans (Fall.) (63) Balby, Doncaster, -/7/77; S.F. IGlobiceps woodroffei Wag. (63) Thorne Moors, 25/6/67; R.C. Hatfield Moor, 29/6/68; R.C. (61)* Hotham Carrs, 11/7/71; R.C. (All specimens det. late G. E. Woodroffe). Allerthorpe Common, 7/7/82; S.F. *Orthotylus adenocarpi (Perr.) (63) Sandall Beat, Doncaster, 26/6/82; S.F. *0. prasinus (Fall.) (63) Sandall Beat, Doncaster, 30/6/82; S.F. *0. tenellus (Fall.) (61) Jeffrey Bog, Kirkham, 11/7/81; B.S.N. (63)* Sandall Beat, Doncaster, 12/6/82; S.F. The only previous record for this species in Yorks, is for Sandsend (VC 62), 1925. t Polymerus unifasciatus (F.) var. lateralis (Hahn) (62) Ashberry, 9/7/78; R.C. (first specimens of this'var. to be recognized in Britain). (64)* Timble Ings, Otley, 25/7/78; R.C. Dallowgill, 7/7/79; J.H.F. Records for the nominate form are: (61) Allerthorpe Common, 1932. (62) Hovingham, 1935. (63) Ecclesall, 1921. (64) Askham Bog, 1942. In the absence of specimens these records should now be considered doubtful. Acetropis gimmerthali (Flor) (61) Allerthorpe Common, 7/7/82; S.F. Only known otherwise in Yorks, at Houghton Wood (VC 61). The following species are recorded for the first time in the vice-counties indicated but do not call for special comment: VC 61: Megacoelum infusum (H-S) VC 62: Gerris lateralis Schumm.; Corixa panzeri (Fieb.); Sigara falleni (Fieb.) VC 63: Picromerus bidens (L.); Gastrodes grossipes (Deg.); Xylocoris galactinus (Fieb.); Phyllus pallipes Fieb. VC 65: Stalia major (Cost.); Heterotoma planicornis (Pall.); Stenotus binotatus (F.); Capsus ater (L.). BOOK REVIEWS Darwin’s Finches by David Lack, with Introduction and Notes by Laurene M. Ratcliffe and Peter T. Boag. Pp. 208, including 8 plates, 27 figures and 32 tables. Cambridge University Press, 1983. £7.25 paperback, £19.50 hardback. A reissue of Darwin’s Finches must be welcomed; it is a volume which should never fade into obscurity, marking as it does the state of knowledge in 1944 concerning the evolution of species, and representative of the erudition of a great evolutionary ornithologist. The Prefaces to the two previous editions (1947 and 1961), written by Lack himself, illustrate changing views on matters such as the likelihood that small differences between sub-species are adaptive. The Introduction constitutes a valuable resume of Lack’s contribution, based on careful observation and analysis of pattern, to evolutionary thought in the 1940s. The Notes provide explanations of Lack’s work and ideas with modern hindsight. There is a comprehensive list of modern references. DJH Genes from the Wild by Robert and Christine Prescott- Allen. Pp. 101, with diagrams. Earthscan: International Institute for Environment and Development, London, 1983. £3 paperback. This Earthscan paperback is a mine of information. It seems to be intended as a source of evidence for those who would argue a case for conservation of wilderness on pragmatic grounds. There are five chapters; the first contains the genuflexion to Mendel mandatory in every elementary genetics text since the first decades of the century. In the remaining chapters many examples of the transfer of genetic material from wild species to crops and domestic animals are given. The costs and benefits of such transfers are discussed; likely recipients of ‘wild genes’, threats to the existence of such genes, and possible conservation measures are examined. An ‘Executive summary’ consisting of 41 aphoristic quotations from the text, and a 193-item list of references complete the work. The book contains a few terminological errors (eg, ‘a given set of chromosomes is called a genome’) and a few meaningless colloquialisms (eg, ‘zapping chromosomes with radiation’); it is readable and is probably comprehensible to non-biologists. DJH 31 THE BLUE BOTTLE FLY CALL1PHORA VICINA R.-D. AS A PARASITE (PRIMARY MYIASIS AGENT), PARTICULARLY ON SMALL MAMMALS Y. Z. ERZINCLIOGLU and SHARON W. DAVIES Department of Zoology, University of Durham, South Road, Durham DH1 3LE Introduction The purpose of this paper is to put on record some cases where the common and ubiquitous blue-bottle species Calliphora vicina R.-D. (= erythrocephala Mg) laid eggs on living woodmice, Apodemus sylvaticus L. These cases, together with certain other records of the same blowfly attacking the hedgehog, Erinaceus europaeus L. , form an ensemble of evidence that suggests the possibility that this blue-bottle species may be a relatively frequent parasite of small mammals, a role that has hitherto virtually escaped detection The attention of students of small mammals is drawn to this subject in the hope that additional records may be revealed, since it seems probable that this blue-bottle parasitism may occur during a limited time of year, but has escaped attention mainly because the results would be mistaken for the fly’s other common activity as colonizer of dead small mammals. The roles of C. vicina in attacking living large mammals (including man) and its well-known occurrence as a secondary myiasis agent of sheep, are reviewed in order to put the above possibility of its importance on small mammals into context. C. VICINA ATTACKING THE WOODMOUSE AND HEDGEHOG During September 1982, a group of five or six woodmice were observed to behave in a rather unusual manner at the edge of a piece of waste ground in Boston Spa. West Yorkshire. They seemed to be lethargic and unusually tame, and did not resist being handled. All the mice were immature. One was easily caught by a cat, but another was found huddled in a corner and covered with blowflies of the species Calliphora vicina which were ovipositing on the lower back and around the tail. Regrettably the mouse was not captured for further examination as it crept away and, presumably, died as it was not seen again. Lethargic behaviour of this sort has previously been reported in woodmice (Morris, 1968) but, as far as we know, this is the first report of this species being attacked by C. vicina. The European hedgehog has long been known to be the host of a variety of blowfly species. Nielsen et al (1978) published a paper on blowfly myiasis in hedgehogs in Denmark. In only five cases did they find that one blowfly species acted alone, three of these being due to C. vicina. the other two were due to Lucilia ampullacea. C. vicina larvae, in the three above mentioned cases, were found in the nostrils, mouth and anus and not in wounds: clearly these were cases of primary myiasis. As in the woodmouse case cited above, all the affected hedgehogs were described as lethargic and immature. Another relevant observation was the finding of a blowfly parasitized hedgehog near York during October 1980. The animal was not captured however and. therefore, it was not possible to determine the species of blowfly maggot concerned. Collecting records at the time show a total absence of Lucilia species in the vicinity; thus it is probable that the parasitizing fly was a species of Calliphora in this case. The sites of infestation were the left ear and eye. C. VICINA ATTACKING SHEEP Some species of blowflies (Calliphoridae) and flesh-flies (Sarcophagidae) have long been known to act as parasites of larger mammals, causing myiasis (ie the invasion of living tissue by maggots). In warmer parts of the world some species are obligate parasites (eg Cordylobia sp., Auchmeromyia sp.) that include man as host, while others are facultative parasites that normally confine their egg or larval deposition to wounds on host animals. We are concerned here with a different phenomenon of myiasis caused by aviposition on unwounded domestic animals, usually sheep. This ‘sheep-strike problem' occurs in cooler parts of the world, such as Britain. Davies (1934). working in North Wales, found that the green-bottle fly, Lucilia sericata Mg., occurred in all 182 cases of sheep-strike that he studied. C. vicina accompanied L. sericata in Naturalist 109 (1984) 32 The Blue-bottle Fly as a Parasite on Small Mammals only two of these cases; in all the other cases L. sericata acted alone. Later, Ratcliffe (1935), working in Scotland, confirmed these results by showing that L. sericata was the only species to emerge from his sheep-strike samples (except in one doubtful record where C. vicina was present as well). Experiments by Ratcliffe showed that the larvae of C. vicina cannot survive a temperature of 32°C, and he concluded from this that these maggots cannot act as primary strike agents in sheep because the skin and body temperature are much higher than 32°C. Later work by Haddow and Thomson (1937) and MacLeod (1937) showed that other Calliphorid species could act as sheep-strike agents in Britain, namely Lucilia caesar (L.), Lucilia illustris Mg., Phormia terraenovae R.-D. and (very rarely) Calliphora vomitoria (L.). Of the 121 cases from England and Scotland discussed by MacLeod, C. vicina occurred in 8 (7 in Scotland and 1 in England); in 2 of the Scottish cases C. vicina acted alone. Attempts by MacLeod to produce C. vicina strike experimentally in sheep failed, except in one case where the larvae developed and produced a wound, but were confined to the cotton-wool cover of the lesion. He concluded that C. vicina was incapable of acting as a primary agent either (a) because the high body temperature of sheep is fatal to the larvae, or (b) that the larvae are unable by themselves to liquefy solid tissue and must depend on the previous activity of Lucilia larvae. Of these two alternatives, it would seem that the first is more probable, for the following reasons: (1) C. vicina is known to cause primary myiasis in mammalian species with a suitable temperature; (2) the first instar larvae of both C. vicina and L. sericata are structurally very similar indeed and there seems to be no morphological feature present in L. sericata that would facilitate its entry into living tissues. It is possible, of course, that L. sericata may release more effective enzymes during extra-oral digestion than C. vicina', there is no evidence for this, however. It is unfortunate that MacLeod’s statement that C. vicina 'is physiologically unable to act as a primary fly, and it is probable that this species is a true obligatory secondary striker’ has been repeated out of context in the literature many times. As it stands it is a statement of universal applicability, whereas MacLeod, like Ratcliffe and Davies before him, was referring specifically to sheep, and moreover to sheep under British conditions. C. VICINA ATTACKING OTHER MAMMALS, INCLUDING MAN Zumpt (1965) collected many records of C. vicina as a myiasis agent. Of these, several were cases of primary myiasis in Man from Libya, one was a cause of urinary myiasis in an old man from Belgium and one was a severe traumatic myiasis in several specimens of the common noctule ( Nyctalus noctula (Schreber)) from Germany. Another record concerned a captive vervet monkey (Cercopithecus aethiops L.); in this case C. vicina was the only Calliphorid present, but larvae of the Muscid species, Muscina stabulans and Fannia canicularis , were also involved. Harvey (1934) reared 114 specimens of C. vicina from a farm labourer in England. The patient had undergone several operations following obstinate urethral strictures, and a permanent opening into the bladder was made for extravasation of urine; it is probable that this was the original site of infection. The man died about one month after the first maggots were observed. Cases of intestinal myiasis in babies are also on record (Nuorteva and Auvinen, 1968). Several other cases of human myiasis by C. vicina in Britain are known to us from personal experience; permission to publish the details of these cases has not been forthcoming, however. All these cases were from elderly invalids and it is worth pointing out here that, although of minor importance, human myiasis in Britain would seem to be more widespread than is generally supposed. Discussion As was noted above, we believe that the low incidence of Calliphora vicina 'strike' in sheep is due to the normally high body temperature of this animal. This view is supported by the fact that C. vicina does attack small mammals with a lower body temperature. It now remains to review briefly the available information on the body temperature of the mammalian species concerned. According to Blaxter et al (1959, a and b) the rectal temperature of sheep range from 37.5°-41°C at ambient temperatures of 8°-20°C. Skin temperatures, however, show a great range according to whether the animal is shorn or not; thus unshorn sheep have a skin temperature The Blue-bottle Fly as a Parasite on Small Mammals 33 range of 36.8°-38.7°C at ambient temperatures of 13.4°-36.2°C. while shorn sheep show a range of 26°-40°C at ambient temperatures of 8°-40°C. It should, however, be pointed out that these figures were obtained from experiments conducted under controlled conditions in the laboratory. Under field conditions the effects of wind and rain may well reduce the skin temperatures below the lower limits given above, while direct radiant sunshine may elevate these temperatures. Regarding Apodemus sylvaticus , this species has been suspected to undergo a spontaneous drop in temperature (torpor) during a few hours each day (Walton and Andrews. 1981a). Walton and Andrews (1981b) found that, at an ambient temperature of 23°C. the body temperatures of woodmice, that had been starved for twenty-four hours, dropped from 34°C to as low as 25.5°C (in single mice) and to 26.5°C (in groups of four huddled mice). This phenomenon is quite well-known in a wide range of mammalian species including Rodents, Insectivores and Marsupials. Although daily torpor occurs typically in winter, it also occurs during the autumn months. Furthermore, cases of summer dormancy are known; this latter phenomenon is not fully understood, however (Hudson, 1978). Morris (1968) measured the body temperatures of two lethargic woodmice by placing a thermometer against different parts of the abdomen. He recorded temperatures of 19.5°C and 16.5°C respectively. The ambient temperature was 2°C. Normally active woodmice whose temperature was measured in this way registered temperatures that were never less than 30°C. The two lethargic mice later became active. The temperatures of the woodmice in the Yorkshire case mentioned above were, unfortunately, not measured. Herter (1965) states that the temperature of hedgehogs will drop to ambient when the latter reaches the critical level of 17°-15°C. This, however, applies only if the hedgehog is physiologically prepared for hibernation. This drop in body temperature reaches 6° or 5° at which point the body temperature of the hedgehog does not drop any further. In the Danish cases mentioned above the temperatures of the hedgehogs were not measured, but the animals were described as being cold and lethargic. These cases were all recorded during August and September 1977. With regards to the cases of human myiasis caused by C. vicina in Britain, patients w'ere all both elderly and unwell (previous to infection). This suggests that body temperature of these individuals was below the normal level, and might explain the attractiveness of these patients to the fly. It is obvious from the above that C. vicina is capable of acting as a primary myiasis agent in mammals with suitably low body temperatures. With regards to sheep, the low skin temperatures recorded by Blaxter et al in shorn sheep may account for the occasional case of strike in sheep w-hen conditions are suitable. It is noteworthy that the tw-o cases recorded by MacLeod in which C. vicina acted alone were from Scotland where the ambient temperature may have been low. It is also worth repeating Ratcliffe's comment that Scottish shepherds firmly believe that the early season strikes (ie in June) are due to the blue-bottle (C. vicina): the green Lucilia only appearing later in the season. Further observations on this aspect of blowfly biology are needed, and any information on myiasis, particularly in small mammals, in Britain w'ould be of great interest. Acknowledgements We would like to thank Dr Lewis Davies for much useful discussion and for reading and commenting on the manuscript. Our thanks also to Mr Barrie D. S. Smith for sending us his observations and specimens from the Apodemus case cited above, and to Dr Nigel Dunstone for his comments. References Blaxter. K. L., Graham. N. McC.. Wainman. F. W., and Armstrong. D. G. (1959a). Environmental temperature, energy metabolism and heat regulation in sheep. II The partition of heat losses in closely clipped sheep. J. agric. Sci. 52: 25-40. Blaxter. K. L., Graham, N. McC. and Wainman. F. W. (1959b). Environmental temperature. energy metabolism and heat regulation in sheep. Ill The metabolism and thermal exchanges of sheep with fleeces. J. agric. Sci. 52: 41-49. 34 The Blue-bottle Fly as a Parasite on Small Mammals Davies, W. M. (1934). The sheep blowfly problem in North Wales. Ann. appl. Biol. 21: 267-282. Haddow, A. J. and Thomson, R. C. M. (1937). Sheep myiasis in south-west Scotland, with special reference to the species concerned. Parasitology. 29: 96-116. Harvey, W. H. (1934). A case of myiasis, due to Calliphora erythrocephala , occurring in man. Parasitology. 26: 306-307. Herter, K. (1965). Hedgehogs , 1-69, Phoenix House. Hudson, J. W. (1978). Shallow daily torpor: a thermoregulatory adaptation, 67-108. In Strategies in Cold: Natural Torpidity and Thermogenesis. (L. C. H. Wang and J. W. Hudson, ed.). Academic Press, New York. MacLeod, J. (1937). The species of Diptera concerned in cutaneous myiasis of sheep in Britain. Proc. R. ent. soc. Lond. (A). 12: 127-133. Morris, P. A (1968). Apparent hypothermia in the woodmouse. Apodemus sylvaticus . J. Zool. 155: 235-236. Nielsen, S. A., Nielsen, B. O. and Walhovd, H. (1978). Blowfly myiasis (Diptera: Calliphoridae Sarcophagidae) in the hedgehog ( Erinaceus europaeus L.). Ent. Medd. 46: 92-94. Nuorteva, P. and Auvinen, E. (1968). A case of intestinal myiasis caused by Calliphora vicina (R.-D.) (Dipt., Calliphoridae) in a baby. Ann. ent. Fenn. 34: 244. Ratcliffe, F. N. (1935). Observations on the sheep blowfly (Lucilia sericata) in Scotland. Ann. appl. Biol. 22: 742-753. Walton, J. B. and Andrews, J. F. (1981a). Torpor induced by food deprivation in the Wood Mouse Apodemus sylvaticus. J. Zool., Lond. 194: 260-263. Walton, J. B. and Andrews, J. F. (1981b). Torpor in single and huddled wood mice (Apodemus sylvaticus (L.)). Acta Universitatis Carolinae — Ser. Biol. 9: 181-184. Zumpt, F. (1965). Myiasis in Man and animals in the Old world. Butterworths, London. BOOK REVIEWS The Natural History of the Mediterranean by Tegwyn Harris. Pp. 224, including numerous coloured illustrations and plates. Pelham Books, 1982. £7.95. A very attractive little book which will undoubtedly be of interest to all naturalists who visit the Mediterranean region. There are short essays on the natural history of different types of habitat in the region, followed by many pages of illustrations of organisms accompanied by short descriptions of each. The range of organisms covered is comprehensive, from seaweeds through higher plants, jellyfish, worms, molluscs, crustaceans, insects, fish, birds, and mammals. The illustrations are clear and descriptions reasonably precise but in a volume this size they are necessarily only a ‘taster’, to stimulate those who have a strictly limited knowledge of natural history. Most readers will quickly outgrow the sections covering the groups of most interest to them and turn rapidly to the further reading section to discover where they can get a more comprehensive account. This guide, which covers approximately 1000 species of plant and animal, is therefore recommended as an introduction to the wildlife of this fascinating area. JEPC Insect Herbivory by I. D. Hodkinson and M. K. Hughes. Pp. 77. Outline Studies in Ecology, Chapman and Hall, 1982. £2.75 paperback. The editors of this series are assembling an impressive collection of small volumes of particular value to students of ecology. The volumes feature clearly-written text and easily understood diagrams. Insect Herbivory introduces the reader to those adaptations of both plants and insects which determine the outcome of interactions between them. Woody and non-woody plants are treated separately and the authors then examine the wider contexts of plant community and ecosystem. Having been given a taste for the subject by this excellent introduction, there are no fewer than 222 references for the reader to explore. JEPC 35 Y.N.U. BRYOLOGICAL SECTION: ANNUAL REPORT 1982 T. L. BLOCKEEL 20 Heathfield Close, Bingley, West Yorkshire BD16 4EQ Excursions The spring meeting for 1982 was held on 1 and 2 May to compile record cards for the British Bryological Society mapping scheme in some under-recorded squares near York. Bolton Percy (VC 64) was chosen as the starting point because of the records of Myrinia pulvinata and Scleropodium cespitans made during the Y.N.U. meeting of 1943 (Cheetham, 1943). Riparian mosses were found to be frequent in the village, even at some distance from the stream, indicating extensive flooding periodically. Tortula latifolia and Leskea polycarpa were plentiful, Homalia trichomanoides was in small amount, and Myrinia pulvinata was found among material of Leskea taken home for examination. Another interesting find, at the base of an old stump, was a small quantity of Radula complanata, a sensitive epiphyte absent from much of the county and quite unexpected in this part of the Vale of York. Barbula trifaria was on brickwork by the stream in an immature form resembling Gyroweisia and with the protonemal gemmae recently described by Whitehouse (1980). Other records in the village were Physcomitrium pyriforme, and in the churchyard Tortula intermedia. Orthotrichum anomalum and Thuidium tamarisci- num. A brief visit was made to the stone bridge south of the village towards the Ings, and here Scleropodium cespitans was present in good quantity, along with Barbula nicholsonii and Orthotrichum cupulatum var. riparium. None of the remaining sites visited proved as interesting as Bolton Percy. At Hagg Wood, near Copmanthorpe, some common woodland species were added to the card. Later in the year Miss J. Robertson was able to add some ephemerals from this area, including Riccia glauca, R. sorocarpa, Dicranella staphylina, Bryum violaceum, and B. klinggraeffii. Such species were not much in evidence at the time of the meeting, although Fissidens exilis and Dicranella schreberana were found in a field by a lane at Scagglethorpe. Redhouse Wood was coniferized and the few bryophytes were almost entirely confined to the rides and dykes. A visit was also made to Moorlands on the eastern side of the R. Ouse (VC 62), where again only common woodland species were present, including Campylopus introflexus. However Pottia truncata growing as an epiphyte on elder made for the unusual. The autumn meeting was held near Richmond (VC 65) on 4 September. Much of the time was spent on the wooded banks of the Swale at Hudswell. Rocks by the river were largely base-poor, and had among other species Blepharostoma trichophvllum, Distichium capillaceum and Heterocladium heteropterum. Close to the water or submerged were Fissidens crassipes, Cinclidotus fontinaloides, Schistidium alpicola, and Hygrohypnum luridum. The higher parts of the woods had extensive limestone outcrops with some luxuriant bryophyte communities, including Neckera crispa, Anomodon viticulosus, Rhynchostegiella tenella, Metzgeria pubescens, M. conjugata, Leiocolea badensis, Porella platyphylla. Radula complanata c. per., and Lejeunea cavifolia. The tiny hepatic Cololejeunea rosettiana was notably plentiful on the shaded cliffs. The woodland ground flora was also locally luxuriant and included Hookeria lucens and Rhytidiadelphus triquetrus. Some time was also spent further to the w'est, near Downholme. The only additional records of note were Frullania dilata and Tortula laevipila on an ancient elm. the only species to enliven a surprisingly poor epiphytic flora. Literature In her presidential address to the Y.N.U. Mrs J. Appleyard gave a broad survey of Yorkshire bryology (Appleyard, 1982). My own notes have included a report of Andreaea in sites near Sheffield and a review of George Stabler's records from Dentdale (Blocked, 1981a. 1982). There have been the usual sectional and excursion reports. Naturalist 109 (1984) 36 Y.N. U. Bryological Section: Annual Report 1982 Records The most important event during the past year has been the description as a new species of Barbula tomaculosa, a moss found in arable fields on the coal measures near Wakefield, Leeds and Rotherham (Blocked, 1981b). It was discovered by chance in 1978 when its rhizoidal tubers were observed under the microscope in material from a field at Stocksmoor Common. In addition to the published stations (VC 63, 64) it has been found recently in a pasture on millstone grit near Cullingworth (44/03. VC 63), a habitat rather different from the earlier ones. Nonetheless there is a clear preference for moist clayey soil. The records which follow are believed to be new stations or the rediscovery of old ones. Riccia cavernosa: (65*) 44/28 Floor of grave! pit, Thorp Perrow nr. Bedale, JR, July 1982. Riccia huebenerana: (64) 44/17 Moist sandy soil, Gouthwaite, TLB, September 1982. Sphenolobus minutus: (64) 34/77 Twistleton Glen. Ingleton, TLB, January 1982. Plagiochila britannica: (64) 34/77 Thorns Gill, Ribblehead, on limestone, TLB, August 1982. Plagiochila spinulosa: (64) 34/77 Thorns Gill, Ribblehead, on limestone cliff face, TLB, August 1982. More usually found on non-calcareous rock. Nowellia curvifolia: (63) 44/04 Old logs. North Beck Woods, Keighley, TLB. January 1982. Scapania scandica: (63*) 34/93 Bank of stream, Crimsworth Dean above Lumb Falls, TLB, January 1982. Lejeunea ulicina: (64) 34/64 Plodder Banks near Whitewell, TLB & EO, March 1982. Lejeunea lamacerina: (63) 44/03 Grit rock by stream, Goitstock Wood near Harden. TLB, February 1982. Sphagnum quinquefarium: (63) 44/04 Among grit rocks, Newsholme Dean, TLB, December 1980; 43/29 Wooded clough, Strines Reservoir. TLB. May 1981. Andreaea rothii: (64) 34/65 Grit boulder. Slate Delph Clough, Bowland Forest, EO, March 1982; 44/16 Grit rock, Ravensgill, TLB. September 1982. Archidium alternifolium: (63*) 34/94 Bare ground in pasture near Glusburn. TLB, January 1982; (64) 44/17 Moist sandy soil, Gouthwaite, TLB, September 1982. Brachydontium trichodes: (64) 44/06 Crumbling grit rock. Ravens Crag, Thorpe, TLB, August 1982. Dicranella staphylina: (65*) 44/37 Arable field west of Skipton-on-Swale, TLB, December 1981. Fissidens exilis: (64) 44/43 Woodland floor, near Micklefield, TLB. March 1982. Aloina brevirostris: (64) 44/43 Disturbed ground on magnesian limestone, Newthorpe Quarry near Micklefield, TLB. March 1982. Pottia lanceolata: (64) 34/87 Soil in crevice on high Yoredale cliffs, Pen-y-ghent. TLB & EO, April 1982. An unusually high altitude for this species. Pottia recta: (64) 44/43 Disturbed ground on magnesian limestone, Newthorpe Quarry near Micklefield, TLB, March 1982. Phascum curvicolle: (64) 44/43 Disturbed ground on magnesian limestone, Newthorpe Quarry near Micklefield, TLB, March 1982. Barbula acuta: (63) 44/42 Roadside bank on magnesian limestone, near Darrington, TLB, February 1981 . Barbula nicholsonii: (63*) 34/95 Stonework by river, Carleton Bridge near Skipton, TLB, Julv 1977. Funaria obtusa: (64) 34/77 Peaty ground. Thorns Gill, Ribblehead, TLB. August 1982. Ephemerum serratum var. serratum: (64) 44/17 Moist sandy soil, Gouthwaite, TLB, September 1982. Pohlia bulbifera: (64) 44/17 Moist sandy soil, Gouthwaite. TLB, September 1982. Bryum gemmiferum: (62* & 65*) 44/37 Soil on banks of Swale, Skipton-on-Swale, TLB, December 1981. Bryum dunense: (64*) 44/43 Disturbed ground on magnesian limestone, Newthorpe Quarry near Micklefield, TLB, March 1982. Bryum sauteri: (63) 34/94 Soil in pasture near Glusburn, TLB, January 1982. Bartramia hallerana: (64) 34/77 Wet rock crevice in ravine, Twistleton Glen, Ingleton, one patch only, TLB. January 1982. Orthotrichum sprucei: (61*) 44/64 Tree base by R. Ouse. Fulford, York, TLB. May 1982: (64) Fungus Forays in 1981 37 44/14 By R. Wharfe, Ilkley, TLB, May 1982; 44/35 By R. Nidd, Little Ribston, TLB. June 1982; 34/65 By R. Hodder near Dunsop Bridge, TLB, July 1982. Orthotrichum pulchellum: (64) 44/35 On elder, Guy’s Crag, Nidd banks south of Knaresborough, TLB, June 1982. Homalia trichomanoides: (63*) 44/13 Tree base by R. Aire, Bingley, TLB, October 1982. Myrinia pulvinata: (65*) 44/28 By R. Ure, Masham, TLB. July 1982. lsothecium striatulum: (64) 34/67 Shaded limestone, Swilla Glen. Ingleton. TLB, August 1982. Entodon concinnus: (64) 44/36 Among grass. Burton Leonard lime quarries, TLB. May 1982. An asterisk indicates a new VC record or amendment to the Census Catalogue. Recorders’ initials: TLB = T. L. Blocked, EO = E. Ormand, JR = Miss J. Robertson. References Appleyard, J. (1982). Bryology in Yorkshire Past and Present. Naturalist. 107: 41-45. Blocked, T. L. (1981a). Andreaea in the Sheffield district. Naturalist. 106: 159. Blocked, T. L. (1981b). Barhula tomaculosa , a new species from arable fields in Yorkshire. J. Bryol. 11: 583-589. Blocked. T. L. (1982). George Stabler and the bryophytes of Dentdale. Naturalist. 107: 71-72. Cheetham, C. A. (1943). Yorkshire naturalists at Bolton Percy. Mosses and lichens. Naturalist (1943): 89. Whitehouse, H. L. K. (1980). The production of protonemal gemmae by mosses growing in deep shade. J. Bryol. 11: 133-138. FUNGUS FORAYS IN 1981 Spring Foray, Darlington, 7-11 May Autumn Foray, Whitby, 10-14 September T. F. HERING In the spring we used a workroom at Polam Hall School. About twenty of us explored the northern edge of the old county (north-west Yorkshire), and the adjoining part of Co. Durham. Teesdale Nature Reserve was by far the richest site; wet weather and a late spring curtailed the lists from other sites. The Autumn Foray, attended by twenty-five people, was a return visit to Whitby, where our 1976 foray had been marred by bad weather. Our headquarters and workroom were at the Queensland Hotel. Mulgrave Woods, though much visited in the past, still managed to produce an agaric new to Britain, and two others new to Yorkshire, and there were also new records from Littlebeck Nature Reserve. I am indebted to Mr W. G. Bramley. Mr P. D. Orton and Dr R. Watling for many of the records below. List of sites Spring Autumn H = Hamsterlev. NZ/093312 R = Rokebv. NZ/083142 B = Brignail Banks. NZ/077122 HF = High Force. NY/885285 and Sun Wood NY/875282 D = Deepdale, NZ/038168 M = Mulgrave Woods. NZ/861 125 L = Little Beck. NZ/879049 A = Arnecliffe. NZ/785050 BH = Beck Hole. NZ/821025 G = Grinkle Park. NZ/742149 * = new record for Yorkshire Naturalist 109 ( 1984) Fungus Forays in 1981 Ascomycetes Ceratocystis ulmi on Elm, L Chaetosphaerella phaeostroma , D Cheilymenia stercorea, H Dasyscyphus pulverulentus , D Humaria hemisphaerica , M Kriegeriella minuta* on Juniper, HF Leptotrochila ranunculi , A Linospora capreae , H Lophodermium juniperinum on Juniper, HF Melastiza chateri, D Morchella esculenta , B Neobulgaria pura, L Phaeohelotium flexuosum , HF Rosellinia mammiformis , D Rutstroemia conformata , D Scutellinia subhirtella* , F (new to Britain) Bastdiomycetes Uredinales and Ustilaginales Gymnosporangium cornutum on Juniper, HF Milesina kriegeriana* on Dryopteris dilatata, HF Puccinia glechomatidis on Glechoma, G Pucciniastrum vaccinii on V. myrtillus, L A Urocystis eranthidis on Eranthis hiemalis , R Heterobasidiomycetes Calocera gloss oides, M Dacrymyces punctiformis on Juniper, HF Aphyllophorales Acia membranacea* , M Amylostereum laevigatum* , HF Basidioradulum radula*, HF Ceriporia reticulata* , HF Cristella confinis* , HF Cristina mucida* , A Cylindrobasidium evolvens* , HF D Fomes fomentarius, M Hydnum rufescens, F Hypochnicium lundellii*, HF Phlebia hydnoides* , HF P. ru/a*,’D Typhula setipes* on Aspen leaves, HF Agaricales Agrocybe brunneola sensu Watling non Fange*, M A. praecox , H Amanita punctata* (Cleland) Reid, F Armillaria ostoyae* , A Bolbitius titubans* , A Boletus appendiculatus* , M B. porosporus , M B spadiceus, A Chamaemyces fracidus , M Clitopilus hobsonii , M C. pinsitus , M Conocybe brunnea, A 39 Fungus Forays in 1981 Coprinus cordisporus on rabbit dung, H Cortinarius armillatus, L C. pseudosalor, L C. rigidus, L Entoloma sordidulum , A Flammulaster aff. carpophiloides* , D (possible new species) Galerina ampullaecystis , D Hebeloma fragilipes* , A Hygrophorus chrysaspis, M H. strangulatus , A Inocybe leptocystis* , M /. obscura, M /. petiginosa , G Lactarius chrysorrheus , L P. cyathula with Alder, A L. glaucescens , L P. hepatic us , L L. obscuratus , L A L. pterosporus , BH G L. volemus , A Leccinum holopus, L Lentinellus cochleatus, A Leptonia babingtonii , A Marasmius hudsonii , A A/, recubans, A M. wynnei, A Mycena rorida, L M. tortuosa , L Naucoria luteolofibrillosa , M Nolanea tenuipes, L N. versatilis* , A Oudemansiella nigra Dorfelt*, M (new to Britain) Pleurotus pulmonarius* , L Psathyrella albidula, B P. obtusata, HF P. pannucioides* , HF (new to Britain) Russula curtipes , L /?. densifolia , L G P. illota*. L P. laurocerasi, M L A P. pue llaris, A P. subfoetens . M G Tubaria autochthona , M P. conspersa. HF P. pellucida , D Tricholoma saponaceum, A P. sciodes , M Fungi imperfecti Bispora antennata, M Cytospora rhododendri on Rhododendron, R C. taxi on Taxus. HF Monilia aurea, M Ovularia obliqua, M Ptychogaster albus, M Fungus Forays in 1981 40 Other interesting records in 1981: Albugo tragopogonis (Pers.) S. F. Gray on Senecio vulgaris. This parasitic fungus resembles the widespread "white blister' of cruciferous plants, with white spores in pimple-like pustules. Long known on Tragopogon , it was unrecorded on groundsel in Britain until 1981, when Mr W. G. Bramley found it near Pickering. Later it was found near Leeds, and in Notts, and Leics. Experiments reported from Sheffield in 1978 by Whipps and Cooke ( Transactions of the British Mycological Society. 70: 389-392) showed no infection of groundsel by races from Tragopogon. Dr S. M. Francis (Commonwealth Mycological Institute, Ferry Lane, Kew, Richmond TW9 3AF) is interested in further reports of this newcomer. Peronospora lepigoni* Albugo caryophyllacearum* . These two parasitic fungi were recognized by Dr Francis on material of Spergularia maritima from Barmston (VC 61). FIELD NOTE Dewberry — Raspberry hybrids My colleagues Dr S. C. Clark and Dr D. D. Bartley recently observed near Pickering, VC 62, a colony of Rubus caesius x idaeus. The plants were intermediate between the two parents, with arching stems, white undersides to the leaves and red fruits with few drupelets. In Stace’s "Hybridisation and the Flora of the British Isles’ (1975: 201) this hybrid is cited by A. Newton for only six English vice-counties, VC 62 being one of those listed. In answer to my enquiry Mr Newton has kindly informed me that his entry for VC 62 was based on a personal field record made near Helmsley in 1975 and that he has since seen the hybrid both in VC 61 near Malton and in VC 63 near Maltby. So far as I am aware this hybrid has not previously been reported in our annual botanical records from any Yorkshire locality. W. A. Sledge BOOK REVIEW Fungi: Folklore, Fiction & Fact by W. P. K. Findlay. Pp. 1 12, including 13 monochrome plates. Richmond Publishing, 1982. £4.75 soft covers. This book is essentially a pot-pourri of anecdotes and quotations relating to man’s interactions with these fascinating organisms throughout history — both the problems they have posed and the pleasures he has derived from them. The text, which is divided into eleven short chapters commences by sampling the writings of the Greeks and Romans, progresses to those of the herbalists and then gives accounts of fairy rings, witches butter and luminous fungi, of ergotism and of dry rot. There follows a chapter dealing with the sexual connotations of some fungi (chiefly Phallus and its allies) and then a compilation of quotes from poetry and fiction of more recent times. The book then turns its attention to our consumption of fungi for nourishment, and the occasional poisonous consequences. It concludes with an introduction to the history of hallucinogenic fungi in general, and devotes its final chapter to the particular case of the fly agaric. Although offering some light-hearted background observations on fungi which may delight the natural historian, the book as a whole is a disappointment. The exposition of most topics are too cryptic to be satisfying, teasing rather than informing; yet this short text abounds with blank pages, spaces and rather arbitrary whole page illustrations. A six-page index has been appended and yet the reader who wishes to delve further into the subject is fobbed off with a mere eight-entry bibliography. The latter does at least include reference to J. Ramsbottom’s excellent Mushrooms and Toadstools in the New Naturalist series, where most of this subject matter is treated in a more scholarly and thorough fashion, and to which the reader would more profitably direct his attention. AB Printed by the Leeds University Priming Service ISSN 0028-077 1 FOCUS ON OUR PRICES HERON 8x40 BWCF BINOCULARS The Birdwatchers Glasses. As used by the forestry commission. Ideal for the young and beginners. 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SEAWARD, MSc, PhD, DSc, FLS, The University, Bradford PAGE 41 53 55 61 67 78 59-60, 69-66, 79 CONTENTS Walter Garstang (1868-1949): Zoological Pioneer and Poet — ft. A. Baker and ft. A. Bayliss Y.N.U. Bryological Section: Annual Report 1983 — T. L. Blockeel The Status of the Mountain Hare, Lepus timidus, in the Peak District — D. W. Yalden Lichen Flora of the West Yorkshire Conurbation — Supplement III (1981-83) — M. ft. D. Seaward a nd A. Henderson Changes in the Status of the Stonechat in the South-West Pennines during the 1970s — J. B. Sykes Field Notes Book Reviews PUBLISHED BY THE YORKSHIRE MORALISTS’ UNION ‘The Naturalist’ is available in microform UNIVERSITY MICROFILMS INTERNATIONAL 300 North Zeeb Road Dept PR Ann Arbor, Mi 48106 USA 30-32 Mortimer Street Dept PR London WIN 7RA England 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. 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Institutions and Subscribers £9.50. All subscriptions should be forwarded to: Mr D. Bramley c/o Doncaster Museum Chequer Road Doncaster DN1 2AE MSP Wf WALTER GARSTANG( 1868-1949): L ZOOLOGICAL PIONEER AND POET R A BAKER University of Leeds and R. A. BAYLISS Robert Gordon’s Institute of Technology, Aberdeen ‘He certainly looked more like a real scientist. He was untidy and always wore rough tweeds and heavy boots.’ This was Leo Walmsley’s description of Garstang as he saw him at Robin Hood’s Bay around 1914 and in marked contrast to the neat and precise Alfred Denny, professor of Zoology at Sheffield (Walmsley, 1969) Garstang spent twenty years as a marine biologist before he came to Leeds University as its first Professor of Zoology and never lost his love for the sea or his association with it. Walter Garstang, born in Blackburn, Lancashire on 9 February 1868 was the eldest son of a medical practitioner of wide intellectual interests who had seven children. The descendants of Walter (1832-1899) and Matilda Mary Garstang, and the bequest for a lecture trust in Blackburn set up by members of the family, are described in a book by one of their daughters (Gurney, 1970). Like his brothers, Walter junior was educated at Queen Elizabeth’s Grammar School, Blackburn and later spoke of its candle-lit interior (Garstang Family, 1956). The school’s Garstang room and portrait commemorate a notable Blackburn family. Walter Garstang went up to Jesus College at 16 xfi years of age in 1884, followed later to Oxford by his two younger brothers. John, the youngest brother, became Professor of Archaeology at the University of Liverpool. Henry Nottidge Moseley who had sailed on the Challenger expedition (Deacon. 1971) and was Linacre Professor of Zoology and Comparative Anatomy in the 1880s, and E. B. Poulton, the entomologist, were significant influences and friends at Oxford and later (Carpenter, 1945). He graduated in 1888 in Natural Science, having abandoned an earlier intention to read medicine and after a nervous illness probably caused by excessively hard work and going up to university at a very young age (Hardy, pers.comm.). Plymouth With the support of a number of leading scientists, the Marine Biological Association was founded in 1884. Garstang began his career at Plymouth a week before the opening day of the new marine laboratory in 1888 at the invitation of Gilbert Bourne. His entry into marine science was described in an unpublished and most unfortunately lost, autobiographical manuscript (Hardy, 1951 A): ‘Fifty years ago, on June 21, 1888. having taken up my degree at Oxford in the morning, I journeyed to Plymouth, picking up the old broad gauge express at Swindon, to begin duty as Secretary and Assistant to the Director of the newly founded laboratory of the Marine Biological Association.’ Garstang went on to describe the last minute preparations for the opening ceremony, including his first meeting with Ray Lankester, ‘the man who was largely to rule my destinies and those of most other young zoologists, for the next twenty years’. A diary kept by Garstang gives an early view of his many pastimes: he was involved in the YMCA and church activities, played the piano and violin, was interested in literature, concerned about his health, had doubts about his faith and was hoping for a demonstrator's post Naturalist 109 (1984) 42 Walter Garstang (1868-1949): Zoological Pioneer and Poet at Oxford. He refused an offer to work at the British Museum. One comment reveals something of the excitement of discovery. ‘17 January 1889. What abundance of life on every bit of rock near low water — enough to last a man a good ten years hard work and then he would say only “Omnia exeunt in mysterium!”’ Bourne left Plymouth after two years and Garstang unsuccessfully applied for the directorship. In the customary printed application (Garstang, 1890) he described his work as ‘secretary, sub-librarian and scientific assistant’, together with ideas on economizing on the small staff and enclosing the usual supporting testimonials. Publications at this time included his first paper on nudibranchiate mollusca (Garstang, 1889). In ‘Notes on the marine invertebrate fauna of Plymouth’ (1892) Garstang made reference to the practical problems never far from under-funded scientists. He wrote that the ‘frequent breakdowns of the small and antiquated launch belonging to the Association seriously interfere with the continuity of our work’. It has been suggested in the memoir of a Director at Plymouth (Kemp, 1943) that there were substantial financial difficulties. A vivid account of the laboratory including Garstang’s own work was written in 1899 (Crossing, 1900). In 1891 Garstang was appointed to a Bishop Berkeley Fellowship at Owens College, Manchester (Thompson, 1886). The fellowships were inspired by the research work of Johns Hopkins University, Baltimore. Garstang was fortunate to work with Arthur Milnes Marshall, Professor of Zoology, whose scientific achievements were complemented by enthusiastic extra-mural teaching (Kelly, 1950). Marshall (1893) referred to Garstang as ‘one of the brightest and most enthusiastic naturalists I have ever come across; a man devoted to science for its own sake’. Oxford After a brief return to Plymouth, Garstang was elected to a fellowship at Lincoln College, Oxford, in 1893 and a lectureship the following year, but he retained his links with Plymouth by visiting the marine station in the summer months. He was supported, amongst others, by W. A. Herdman, who declared that Garstang had been ‘one of the most promising young zoologists’ who had ‘that fertility of resource ... so necessary for the successful investigation of scientific problems’. Poulton, who had already published an account of Garstang’s researches (Poulton, 1890), wrote to Garstang advising him to go to Lincoln College and added that ‘I feel sure that Miss Ackroyd would agree with my advice’ (Poulton, 1893). Lucy Ackroyd of Newnham College, whose family home was in Bradford, had met Garstang at Plymouth and later married him in 1895. Garstang lectured at Lincoln and Jesus Colleges, worked with Lankester and lectured in extension classes. Garstang’s letter to Michael Sadler, then Secretary of the Oxford Extension Delegacy, explained his view that certain branches of zoology were especially suitable for extension classes, ‘chiefly those which deal with animals themselves as living creatures rather than their minute organization’. Lincoln College was small and a historian has recently described it as being in the ‘complacent afternoon of the Indian summer of the late Victorian age’ (Green, 1979). Although Garstang was the only scientific fellow, he enjoyed the friendship of a notable senior colleague, William Warde Fowler, classical scholar and ornithologist, who was an important influence in developing in the 1880s the practice of systematic observation of wildlife with ‘notebook and a first class telescope or field-glass’ (Allen, 1978). Fowler was co-editor with Louis Miall, Garstang’s predecessor at Leeds University, of an edition of Gilbert White’s Natural History and Antiquities of Selborne (Miall & Fowler, 1901). It is clear that Garstang had a great affection for Fowler. In Songs of the Birds, Garstang (1922 A) included a dedication, in the form of a poem, ‘as a memorial of my debt to him’. The lines included: ‘Three things old friend in youth revealed With you are interwoven, A common room, rich walks afield. Rare evenings with Beethoven.’ 43 Walter Gars tang (1868-1949): Zoological Pioneer and Poet Garstang’s association with Plymouth continued. The first vacation classes were started by him in 1895 when he took a group of Oxford students there. It was during his period as Fellow and Lecturer at Lincoln that he first propounded his ideas on the ancestry of vertebrates. ‘My theory postulates a common ancestor for the Echinodermata, the Enteropneusta and the Chordata (Garstang, 1894) and emphasizes the value of functional as well as structural considerations in matters of phylogeny’ (Garstang, 1897 A). Garstang visited Toronto in 1897 to attend the British Association meeting. A letter written on the voyage (Garstang, 1897 B) included a characteristic comment on a distant view of the Isle of Man, the sighting of Herdman’s laboratory at Port Erin, with 'high ground inland suggesting capital walks and invigorating breezes’. He arranged to collect plankton with the help of the captain and wrote that ‘I shall be able to collect a complete set of samples with no difficulty at all.’ Garstang returned to Plymouth in 1897 as Naturalist in charge of Fishery investigations. In his final years at Plymouth important moves were underway as the fisheries industry of Europe was ‘jostled from its thousand year slumber’ (Schlee, 1973). Technological change, notably in the application of steam power and later motor power, had led to international co-operation with a series of important conferences in Scandinavia. Garstang attended the second International Conference for the Exploration of the Sea at Christiania, Norway in 1901 with another notable scientist, D’Arcy Wentworth Thompson (Mill, 1902). F. Nansen and J. Hjort were the Norwegian delegates. The International Council for the Exploration of the Sea, founded in 1902 as a result of the second International Conference, brought together a distinguished group of scientists including Johan Hjort, Fridtjof Nansen and C. G. J. Petersen. Garstang later wrote to Hardy about his friendship with Hjort and his ‘admittance to the Viking fraternity' (Hardy, 1951 B). He became convenor of what was initially called the Over-Fishing Committee from 1902 to 1907 as part of a complicated programme of both hydrographical and biological work of eight European nations (Schlee, 1973). Britain supported this work and the Scottish Fishery Board and Marine Biological Association organized the scientific work. When W. C. M’lntosh, a pioneer in marine biology, occupant of the University chair in Natural History at St Andrew's and an authority on Scottish fisheries, published his book on the ‘Resources of the Sea’ in 1899, he claimed that the sea fisheries were inexhaustible. This view received much criticism and ‘the most trenchant criticism came from Dr Walter Garstang’ (Gunther, 1977). Garstang took the opposite view, suggested in the title of his paper, ‘The Impoverishment of the Sea', published a year later, in which he was led to the conslusion ‘that the bottom fisheries were not only not inexhaustible, but in rapid and continuous process of exhaustion; that the rate at which sea fishes multiply and grow, even in favourable seasons, is exceeded by the rate of capture’ (Garstang, 1900). M’lntosh used Huxley’s earlier claim in supporting evidence, but Garstang was quick to point out that Huxley had included the phrase ‘in relation to our present modes of fishing’. Clearly M’lntosh took no account of the immense advances in the powerful sea trawlers built in the 1890s and much of the basis of his book was subsequently questioned. Lowestoft In 1902 a new laboratory was established at Lowestoft, and Garstang began there a productive period, described as ‘those classical investigations into the natural history of the North Sea plaice which have laid the foundations of English fishery research’ (Hardy, 1951 A). A steam trawler was bought by another important marine scientist, G. P. Bidder, and leased to the Association, one of several acts of generosity (Gray, 1972). The appropriately named SS Huxley was equipped with cabins in the fish hold and a laboratory on deck and in an account entitled ‘Down in the deep — the romance of fish life’, Garstang described the work of catching, measuring and ‘tagging’ fish with the discs devised originally by C. G. J. Petersen (Garstang Family, 1902). He conceded, no doubt ironically, that the fish would be ‘lopsided’ but otherwise unharmed. Fishermen were to be paid for returned specimens. The Lowestoft laboratory was not lavishly housed, occupying for years a little white fronted 44 Walter Garstang (1868-1949): Zoological Pioneer and Poet shop in Waveney Road. ‘From here the North Sea investigations work was directed between 1902 and 1906 when a tiny staff moved to a house in the Marina. In this building . . . was set up the first internal semblance of a laboratory’ (Atkinson, 1955). Garstang’s team caught and freed large numbers of plaice, studying natural growth rates and migrations. Experiments in transplantation pointed clearly to the benefits to be gained in increased growth by moving plaice to richer feeding grounds. The establishment of the International Council at which Britain stressed above all the importance of studying over-fishing made this revival of fishery research essential. The over-fishing subject involves studies on the reproduction, migration and growth of fish and until these investigations were carried out at Lowestoft there was ‘no exact knowledge applicable to the fishing grounds’ around our coasts (Garstang, 1906). Accounts of the work were published by the International Council and in Government publications (Garstang, 1905, 1909 B, 1909 D). His own revealing comment in his application for the Oxford chair in 1921 makes clear his feelings: ‘The sequel (to the International Conference of 1901) is well known . . . although few zoologists appear to have realized what it meant to carry on one’s regular work of investigation and at the same time maintain almost single-handed, a steady pressure towards the goal in sight with fluctuating counsels behind, active and relentless opposition in front and cross currents of intrigue and misrepresentations constantly at work ... I would add simply that the programme drawn up by Hjort, Heincke and myself in 1901, the policy I advocated and the results achieved under my direction have long been accepted as the basis of organized fishery investigations in this country and have placed the utility of Marine Zoology, especially on the Bionomical side, outside the pale of controversy.’ From 1902 to 1908 Garstang was scientific adviser to the British delegates to the Council and convenor of the International Committee on trawling investigations. Hardy’s point (1951 A) that Garstang ‘felt he would never be happy if he was not entirely free to shape his own policy’ echoes Garstang’s own views. One might speculate that Garstang was not happy in handling the particular mix of politicians, civil servants, industrialists, and scientists in fisheries research. As Hardy (1950) put it in an obituary, ‘Garstang was an individualist, a lover of freedom and independence, who resented what appeared to him to be government interference in his scientific programme when official policy did not coincide with his own plans.’ Leeds The Board of Agriculture and Fisheries was about to extend its control over fishery investigation when Leeds University decided to create two new departments and professorial chairs of botany and zoology to replace the biology department headed by L. C. Miall, who had held the foundation chair since 1876 (Baker & Bayliss, 1983). In 1910 the Fisheries Department eventually took over and the laboratory at Lowestoft was dismantled while the research vessel Huxley was sold by the Marine Biological Association. In a letter to the Vice-Chancellor at Leeds, Garstang (1907 A) wrote ‘I earnestly wish to resume academic work’ and two days later wrote to Miall that ‘several of those upon whom I should have depended for support are now no more — Professor Moseley, Weldon, Howes, and Milnes Marshall, but several of my friends, though embarrassed by the late announcement of my desire, have promised to write on my account’ (Garstang, 1907 B). Garstang wrote to Bodington, the Vice-Chancellor, again, after his interview amplifying his proposals for research: ‘The important fisheries carried out from Hull and Scarborough seem to me to show that an expansion of the present field of biological research carried out at the college in the direction of the sea fisheries would be legitimate.’ He was however conscious of the needs of the whole area, not just the coastal fisheries and went on: ‘I should therefore propose, if elected, to devote my first year to making myself acquainted with all the branches of biological activity at present underway, or contemplated, while winding up my own reports on past investigations’ (Garstang, 1907 C). Garstang, with an Oxford DSc awarded in 1906, was a candidate in a very strong field (University of Leeds, 1907 A): Ernest MacBride, already a professor at McGill and an FRS (Caiman, 1941) was to play a significant part later in Garstang’s life. Edwin Goodrich, also an FRS by 1905, later occupied the Linacre chair at Oxford for which Garstang competed (Dc Beer, Walter Garstang (1868-1949): Zoological Pioneer and Poet 45 1972). F. W. Gamble FRS had been a Berkeley fellow at Owens (Hickson, 1927) and J. S. Gardiner, later appointed Professor of Zoology at Cambridge, had worked at Naples and was an authority on corals (Forster-Cooper, 1948). He also was later to become an FRS. The committee on the biological chairs resolved to report to the Council that ‘the Chair of Zoology should be offered to Dr Garstang and the Chair of Botany to Mr Blackman and that to each chair should be attached a fixed salary of £550 a year' (University of Leeds, 1907 B). Garstang’s success may have owed something to the Vice-Chancellor's former links with Lincoln College. His background in applied research was thought to be one of the reasons for his appointment ( Yonge, pers.comm.). He had by 1907 some fifty publications and was well known at Oxford and Manchester, and on parliamentary committees and international forums. It may be noted that the successful Professor of Botany, Vernon Herbert Blackman (1872-1967) became a distinguished plant physiologist and left Leeds for Imperial College in 1911. Garstang (1935) wrote of ‘being brought to anchor in an inland town, away from the sea which had hitherto absorbed my interest’. The immediate task was to create a new department in a small university only a few years old. with limited advanced work and no students engaged in advanced zoology. With a staff of three, transferred from the previous biology department, including a ‘keeper of the insect collection’, he began to build up his new activities. Garstang’s predecessor, Louis Compton Miall, was well known as an entomologist (Baker & Bayliss, 1983). In the University annual reports, the work is outlined. For 1908-1909 there was a note of research ‘for a large part of their vacations’ and publications listed for Garstang and a new member of staff, Marie Lebour, who later worked for many years at Plymouth (University of Leeds, 1909). Marie Lebour went to Plymouth in 1915 on temporary ‘loan’ and remained there nearly fifty years (Russell, 1972). She was a friend of the Garstang family and later worked with Robert Gurney, Garstang’s brother-in-law. In 1910 Garstang’s investigations into insect colours and markings were reported, with Lebour’s vacation work at the Millport and Cullercoats marine stations and a significant reference to a student engaged on an honours course (University of Leeds, 1910). An important part of the University’s work was the education of teachers. One Saturday morning course was held in the session 1912-13. Students were asked to provide ‘a few simple instruments’ and the more squeamish were advised that there would be ‘dissections rare and optional’. There were also summer vacation courses at Bingley Training College, and Garstang later described his work to ‘overcome the ill-considered antagonism of the Board of Education to the encouragement of Zoology in schools’, confirmed in Jenkins’ (1979) detailed analysis of science in education. In University affairs, Garstang was involved in a number of activities. He continued his early interest in rowing by supporting the University Boat Club and was prominent in the Officers’ Training Corps. He was Dean of the Faculty of Science (1917-1921), Pro-Vice-Chancellor (1929-1931) and claimed to have undertaken a ‘more than average share of administrative duties’. This was an important stage in the development of the University: Yorkshire College had received its charter on 25 April 1904 and by the 1920s had developed the early technological bias by a broader range of subjects. Most of the students were reading for degrees, including postgraduate work, and the First World War not only brought a later influx of more mature students but also substantial encouragement of research (Gosden & Taylor, 1975). In the First World War, Garstang did not go on active service but was a machine-gun instructor and was later commissioned in the volunteer battalion of the West Yorkshire Regiment. He also visited France in 1918 to lecture on zoological and educational topics to the troops, arranged through the YMCA and Army GHQ. France. Garstang noticed what he believed to be a mistake in a memorandum which stated ‘The primary object of the lectures will be the raising of the morals of the troops’ and found that ‘morale' was intended. He later wrote in his diary ‘Imagine my disappointment when I was told that I was marked down for the convalescent camps in the Trouville area, not up the line. My subject was supposed to be interesting enough to entertain the wounded but not pointed enough for the front line men — for that you must be talking on “War Aims”.’ His lecture titles included camouflage and mimicry, North Sea Fisheries, birds and their songs and the social life of animals (Garstang. 1918). ’ 46 Walter Garstang (1868-1949): Zoological Pioneer and Poet Garstang (1919 A) wrote of Oxford contemporaries who died in the Great War, including Edward Minchin, a protozoologist of note and Arthur Darbishire, Edinburgh University’s first genetics lecturer. An important part of Garstang’s work was the support he gave to amateurs and their societies. A former student later commented, ‘He never isolated himself from the outside world ... he stimulated the interests of the amateur biologists of Yorkshire’ (Eastham, 1949). In 1921 he wrote ‘Throughout my residence in Leeds I have endeavoured to revive the waning interest of the public in Zoology by stimulating the work of local Natural History and other Societies, in the course of which I have given a great number of semi-popular lectures up and down the Ridings on various aspects of animal life. I have also on several occasions conducted long vacation courses in Natural History upon the invitation of the West Riding Education Authority at the Bingley Training College. Some may deny the value of such work as involving a diffusion of energy, but for my part I am convinced that in my present position it has been a duty.’ At Plymouth he had been one of the founders and first President of a local Field Naturalists’ Club and was a past President of the Norfolk and Norwich Naturalists’ Society. His presidential address read in March 1905 was on ‘The natural history of the North Sea’ (Garstang, 1909 A). He joined the Yorkshire Naturalists’ Union and wrote for The Naturalist (Garstang, 1909 C, 1919 C, 1920). He was President in 1918 and his address on ‘Nature and Man’ (Garstang, 1919 B) was an interesting and wide ranging one. He thought that the ‘mantle of the prophet is passing — indeed has already passed — from the poet to the man of science’. He concluded: ‘In my opinion the future welfare of humanity depends more upon the training of the sentiments and emotions in the light of a knowledge of nature than upon anything else.’ In the same journal thirty years later, Henson (1949) in an obituary wrote ‘This was the first of his writings betokening a rapidly widening interest in terrestrial natural history and a rapidly developing artistic appreciation of nature. With Professor Priestley, Garstang also helped to revive the fortunes of the Leeds Naturalists’ Club by providing a home and bringing its members into closer contact with the University (Donnan, 1951). The Leeds Philosophical and Literary Society also had scientific interests. Garstang’s contributions included the presidential address on 20 October 1925 on ‘Wordsworth’s Interpretation of Nature’ which appeared as a Supplement to Nature (Garstang, 1926 B). What was initially called the Yorkshire Universities’ Marine Laboratory, and more recently the Wellcome Marine Laboratory (now closed and sold) had originally been a joint venture of the Universities of Leeds and Sheffield. Garstang in 1912 persuaded the University to rent part of a row of buildings adjacent to the slipway as a combined laboratory-living cottage for student field studies at Robin Hood’s Bay. The buildings were purchased in 1922, ‘a good investment if nothing else said the University Surveyor’ (undated typescript, Wellcome Marine Laboratory, Box 1-A2 History & Development, University of Leeds Archives). His main connection with living marine animals during his period at Leeds was through this laboratory. He first visited the village at Easter 1909 with T. H. Taylor, then one of his lecturers, and LI. Lloyd, a student. Lloyd (1949) later recalled the occasion — ‘We stayed at the Bay Hotel and its verandah, the sea washing its walls at high tide, was our laboratory.’ Apart from the work at Robin Hood's Bay and the links with Plymouth, Garstang became a member in 1919 of the Scientific Advisory Committee on Fishery Research of the Development Commission established in 1909 where he initially sat with G. C. Bourne and E. W. MacBride. He wrote articles for The Times (Garstang, 1926 A) on the effects of the Great War on fishing stocks and gave the Buckland lectures in 1930 at Grimsby and Hull. Garstang referred to the potentialities of a North Sea ‘great fish farm’, which only awaited more knowledge before it could be cultivated rather than exploited. ‘So far it must be admitted it is a farm in which Nature has done all the sowing and man only the reaping.’ He envisaged the future prediction of fishing stocks using ‘knowledge of physical and planktonic conditions’. ‘The subsequent history of fishing in Northern Europe has yet to point to an easy solution of what are now old problems’ (Garstang, 1930). Garstang’s musical and literary interests were expressed in various ways and especially as part of an interest in bird song. A reviewer of his book ‘Songs of the Birds’ in Nature noted the ‘welcome reaction from the too mechanical conceptions that are common but there is at the 47 Walter Garstang (1868-1949): Zoological Pioneer and Poet same time some danger of their leading towards too anthropomorphic ways of thinking’. The reviewer decided not to comment on the verse. Another reviewer in Ibis commented that the book ‘makes an appeal to all bird lovers, but it is a difficult book to analyse’. Garstang lectured extensively on the subject. A report in The Daily News (7/1/1922) describes a lecture on bird song to the Parents’ National Educational Union in London. ‘Professor Garstang relied upon his own vocal powers when his supply of gramophone records ran out.’ He was able to articulate the sounds of almost all the British song birds. Garstang’s verse was also written in light hearted parodies like ‘The Students’ Opera’ (Garstang, 1922 B). He also wrote verse for more serious scientific argument and to illustrate his ideas on evolution ‘for in truth he often made his points with greater force in his light hearted verse than in his more technical scientific prose’ (Hardy, 1956). He found some of these were not acceptable to the editor of Nature. Gregory (1922) wrote ‘I do not think they are at all comparable with your poems on Bird Songs’ and in another letter he quoted the comments of a referee: ‘unless this sort of thing is exquisitely well done, with delicate rhymes and real wit in the allusions, it oughtn’t to be done at all.’ Walter Garstang on the shore at Robin Hood's Bay. (Photograph by courtesy of Dr H. Henson) Retirement Garstang retired in 1933. The Gryphon referred to his ‘sailor's breeziness’ and ‘his happiest classes ... on the shore at Robin Hood's Bay’ (Lloyd. 1933). A colleague. H. Henson (1974), commented on his autocratic style as head of department and on an attitude towards letters that will strike a sympathetic chord with many, ‘If you leave them long enough they answer themselves.’ The Zoology Department had grown, and by 1933 Garstang had a staff of five including a Reader, Llewellyn Lloyd, who had been the first research student in the department. Garstang's enthusiasm for scientific investigation did not diminish with age. He published major works long after his retirement and he retained a close working friendship with Robert 48 Walter Gars tang (1868-1949): Zoological Pioneer and Poet Gurney. Gurney never held an official position but worked continuously in his private laboratory, first in Norfolk, then in Oxford, his speciality being decapod and copepod Crustacea. Garstang went to Bermuda twice, in 1935 and 1938, with Robert Gurney and wrote with excitement and enthusiasm to his wife of the ‘discovery of a most amazing globular larva’ (Garstang, 1938). He later found the larva had already been described as the genus Planctosphaera . Correspondence with discovery staff at the British Museum in 1948 (Totton, 1948) and the draft of a paper (Hardy Collection) show that Garstang was still working on the description of a new species when he was eighty years old. Two contributions to the Zoology reports of the ‘Terra Nova’ discoveries appeared after his retirement, one with Elizabeth Georgeson, a former PhD student (Garstang, 1934; Garstang and Georgeson, 1935). Garstang attended a Linnean Society discussion on bird song on 10 February 1949, and died on 23 February in Oxford. The obituaries included one by a former student, Leonard Eastham (1949), who became Professor of Zoology at Sheffield, who paid a tribute to work done at Leeds: ‘Great as were his contributions to marine zoology, both academic and economic, zoology was the gainer by this change. For his virile mind was thereby released to pursue the philosophical and speculative tasks for which he was so patently fitted . He inspired them (the students) with a love for the subject and . . . made them think and see for themselves. Few who go from us will leave behind so much affection and such a sense of gratitude.’ Other obituaries described him as a ‘kindly, sympathetic, highly cultured English gentleman’ (Henson, 1949) ; ‘a great teacher’ (Hardy, 1951 A); as a man who when obliged to fight ‘thoroughly enjoyed it’ (Hardy, 1950). These and other memories echo the earlier words of Michael Sadler (1921): ‘He is young in mind and is in sympathy with young men. From him they and their elders catch the infection of a love for nature and an exact and appreciative observation of the habits and beauty of living things.’ The University of Leeds commemorates his memory in the name of the Garstang student flats and in the Walter Garstang Fund administered by the Zoology Department to give financial help to ‘meritorious students working in the area of marine or freshwater biology and in particular to enable undergraduate students to attend vacation courses at the Plymouth laboratory’. Assessment of Garstang’s Work Garstang’s scientific work may be divided into several broad areas. The first twenty years were very productive in a branch of science under rapid development. He worked with distinguished scientists and made a notable contribution. His studies in invertebrate marine biology at Plymouth and Oxford from 1888 to 1897 led to twenty-nine papers mainly on the morphology, ecology and distribution of marine animals and in this, the first phase of his working life: ‘He made delightful studies on the habits and life of nudibranchs, he did early experiments with fish, on the theory of warning colourations . . . and studies on the burrowing crab Corystes' (Hardy, 1973). For the next decade he carried out pioneer work on fishery research at Plymouth and Lowestoft, which provided a scientific basis for the analysis of fishery problems. Working particularly in the field of bionomics on the races and migrations of the mackerel and later on the plaice, his studies included transplantation experiments on fish, ocean currents, the artificial rearing of sea fish, investigations of the plankton, the physical conditions in the English Channel and the impoverish- ment of the sea. Recent work using enzyme electrophoresis tentatively suggests that Garstang’s work on races of mackerel was ‘ahead of its time and probably correct’ (Southward, pers. comm.). His own accounts of what was achieved are necessarily brief and do not give a full story of years of difficult work by scientists of many nations in an atmosphere of political complication and in an industry of sturdy independence. An obituary ( The Times , 1/3/1949) said of this period: ‘He was the moving spirit in the development of English fisheries research ... a pioneer in bringing about the International Council for the Exploration of the Sea 49 Walter Garstang (1868-1949): Zoological Pioneer and Poet . . . (and carried out) classical investigations in the Natural History of North Sea Plaice.’ Before he was forty he had achieved more than most might do in a lifetime of scientific work but these fundamental investigations were in some ways to be overshadowed by what was to come. It is in the field of evolutionary theory that he is perhaps best remembered for his speculative and controversial ideas. Two words predominate in any assessment of his contribution to zoology — Recapitulation and Paedomorphosis. Many zoologists believe that Garstang’s (1922 C) paper on ‘The Theory of Recapitulation’ was his most important contribution. Henson (1974) described it as ‘one of the most intellectual accomplishments from the Department of Zoology at Leeds’ . The ‘Biogenetic law' first put forward by Haeckel in 1866 had been revived by MacBride (1914) in his Textbook of Embryology. Garstang attempted to re-define the foundations of the law, the basis of which he believed to be unsound. In playing a major part in the downfall of Haeckel’s theory he came into direct conflict with MacBride (1926). Corresondence in Nature later, highlighted their differences when a series of letters on Natural Selection contained these sentences: ‘the confusions and obscurities in Professor Mac- Bride’s recent letter’ (Garstang, 1929 B) and ‘Professor Garstang’s letter appears to me to be trivial’ (MacBride, 1929). De Beer, however, was one of the first to recognize the contribution that Garstang had made; referring to the theory of Haeckel, he stated ‘we are rid of a mental straight-jacket which has had a lamentable effect on biological progress' (De Beer. 1940). A more modern synthesis of Garstang’s views and their full significance for evolutionary theory are reviewed by Hardy (Huxley, Hardy & Ford, 1958). In the same 1922 paper, Garstang first introduced his own term Paedomorphosis (like a child) to describe the prolongation of the early phase of development into sexually adult life. He used this as the basis for his theory of evolution. It was almost the direct opposite of the biogenetic law. Gould (1977) describes this contradictory process: ‘Recapitulation requires that adult features of ancestors appear in the juvenile stages of descendants. Nothing therefore can be more contrary to its operation than the incorporation of previously juvenile features into the adult stages of descendants. ' He also believes that Garstang by writing in the context of evolutionary theory ‘revised by Mendelian genetics . . . regarded paedomorphosis as orthodox . . . (and) assured the downfall of recapitulation’. Garstang was the first to look for traces of the ancestors of the vertebrates in early instead of adult stagesof invertebrates; and he focussed his attention on the larvae of Echinoderms ( De Beer. 1 940). J. Z. Young (1981), referring to Garstang’s theory of vertebrate ancestry, believed it to be ‘necessarily speculative but it has certain strong marks of inherent probability.’ Three other important papers by Garstang appeared later. In his presidential address to the British Association Zoology Section in 1928 on ‘The origin and evolution of larval forms’ (1929 A) Garstang again put forward his speculative views on the course of evolution, ending with an old German saying, ‘Behaupten ist nicht beweisen,’ meaning that to assert something is not to prove it and asked his audience to throw the windows of the mind open. In the same year he gave that address, he published his paper on 'The morphology of the Tunicata and its bearings on the phylogeny of the Chordata’ (Garstang. 1928) in which he developed in greater detail the theory he first proposed in 1894 ‘that the Chordates were derived by paedomorphosis from the pelagic larval form of sedentary invertebrates’ (Hardy, 1950). Finally, three years before he died, he published ‘The morphology and the relations of the Siphonophora' (Garstang, 1946), a long paper which prompted a worker at the British Museum to comment ‘a masterly digest of Siphonophore literature ... am filled with admiration' (Totton, 1948). Garstang maintained his interest in larval forms and evolution throughout his life. We can gain some idea of how Garstang himself viewed his theories: in a typed draft of his proposed work on ‘Larval Forms’, Garstang intended to quote on the front page from Victor Franz’s Geschichte der Organismen (1924) the following: ‘One cannot deny that Goethe often had a very shrewd judgment on morphological principles, just because he did not depend on dominant theories, but sought himself independently to find in Nature how she presents herself.’ There seems little doubt that Henson (1974) was right when he said that Garstang ‘was a much greater zoologist than he was normally given credit for'. Gould (1977) describes him as ‘an excellent functional biologist'. However. Garstang's attack on Haeckel’s law brought him into 50 direct conflict with some of the very influential zoologists of his time. The absence of the highest academic honours and also the honours bestowed by Governments may be partly explained by this. Garstang’s scientific stature is still a matter of controversy. He was not elected to a fellowship of The Royal Society and several people close to him have suggested that personal animosity may have been partly the reason . Some people did not take his phylogenetic speculation seriously. Evolution- ary studies of this kind were being replaced by more experimental methods using modern techniques in zoology and speculation as to the origin of different groups of animals was becoming old fashioned. Lloyd (1933) referred to the subject as ‘that zoological country of lost endeavours where the backboned emerge from the backboneless’. There is also the absence of substantial written scientific work for many years after his arrival at Leeds. The poetry, sometimes written with serious intent, but often as doggerel as an aid to student learning, might have been too slender a vehicle to draw substantial attention and may have counted against him . The attempt to capture bird song on paper could be interpreted as a brave attempt, but unsatisfactory in its results and unacceptable to the specialists in animal behaviour. With regard to honours and The Royal Society Fellowship , it has been suggested that Garstang, because he had worked on a wide range of research topics, was not identified with a special subject, ‘But there was also a bias against “fisheries” which was regarded as “trade”, not pure science’ (Southward, pers.comm). History is concerned with the recoverable past. In an analogy particularly relevant to the study of a marine scientist, Lytton Strachey wrote of the biographer who could ‘row over that great ocean of material and lower down into it, here and there, a little bucket, which will bring up to the light of day some characteristic specimen’ (Briggs, 1965). Many interesting questions remain unanswered because the material is not there to be dredged. Garstang’s own account would have been of great value to this brief assessment of his life and work and a larger scale evaluation is clearly necessary. Garstang was ‘a poet and a lover of nature’ (Hardy, 1950) and a ‘zoological entertainer’ (Garstang, 1951), and there can be no doubt that he was also what the Challenger seamen would have called him — a philosopher. Acknowledgements We wish to acknowledge the help of Sir Alister and Lady Hardy , Michael Hardy , H . Henson , P . F. Johnston, M. R. D. Seaward, A. J. Southward, M. Stone, and Sir Maurice Yonge, and thestaff of the libraries in Aberdeen, Blackburn, Leeds, Lowestoft, and Plymouth. Note A complete bibliography has not been attempted but Alister Hardy’s list of works in Larval Forms (1951) contains over 100 titles and Garstang’s published papers, extensive use of which have been made here, are in three bound volumes held in the Edward Boyle Library at the University of Leeds. References Allen, D. E. (1978). The Naturalist in Britain. Penguin, Harmondsworth. Atkinson, G. T. (1955). Current research in the Lowestoft Laboratory. Eastern Daily Press, 11 August 1955. Baker, R. A. and Bayliss, R. A. (1983). Louis Compton Miall FRS: Scientist and Educator 1842-1921. Notes and Records R. Soc. Lond. 37: 201-234. Briggs, A. (1965). Victorian People. Penguin, Harmondsworth. Caiman, W. T. (1941). E. W. MacBride. Obit. Not. Fell. R. Soc. Lond. 4: 747-759. Carpenter, G. D. H. (1945). E. B. Poulton. Obit. Not. Fell. R. Soc. Lond. 4: 655-680. Crossing, W. (1900). A Visit to the Marine Laboratory on Plymouth Hoe. Doidge's Western Counties Illustrated Annual. Plymouth. De Beer, G. R. (1940). Embryos and Ancestors. Clarendon Press, Oxford. De Beer, G. R. (1972). E. S. Goodrich. Dictionary of Scientific Biography. 5: 466-467. Deacon, M. (1971). Scientists and the Sea, 1650-1900: A Study of Marine Science. Academic Press, London. Donnan, J. V. (1951). The Leeds Naturalists’ Club, 1870-1950. Naturalist. 109-114. 51 Walter Garstang (1868-1949): Zoological Pioneer and Poet Eastham, L. (1949). W. Garstang. Nature, Lond. 163: 518—519. Forster-Cooper, C. (1948). J. S. Gardiner. Obit. Not. Fell. R. Soc. Lond. 5: 541-553. Franz, V. (1924). Geschichte der Organismen. G. Fisher, Jena. Garstang Family (1902). Down in the Deep. Blackburn District Libraries. B921. Garstang Family (1956). The Garstang Family. Newspaper cutting from the Blackburn Times, 21 September 1956. Blackburn District Libraries, B921. Garstang, W. (1889). Report on the nudibranchiate mollusca of Plymouth Sound. J. mar. biol. Ass. UK. 1: 173-198. Garstang, W. (1890). Application for Directorship at Plymouth Laboratory. 3 November 1890 (Alister Hardy Papers). Garstang, W. (1892). Notes on the marine Invertebrate Fauna of Plymouth for 1892. J. mar. biol. Ass. UK. 2: 333-339. Garstang, W. (1894). Preliminary note on a new theory of the phylogeny of the Chordata. Zoologischen Anzeiger. 17: 122-125. Garstang, W. (1897 A). Physiology and the ancestry of vertebrates. Trans. Jun. Sci. Club, Oxford. (NS). 1: 1-6. Garstang, W. (1897 B). Letter dated 29 July 1897 written from Steamship Laurentian on voyage to Toronto (Alister Hardy papers). Garstang, W. (1900). The impoverishment of the sea. J. mar. biol. Ass. UK 6: 1-69. Garstang, W. (1905). General report on fishery investigations: report on experiments with marked fish during 1902-03, etc. 1-198; Fishery and Hydrographical Investigations in the North Sea and Adjacent Waters (Southern Area). 2nd report Cd. 2670 HMSO, London. Pari, papers XIV 663. Garstang, W. (1906). The fisheries of the North Sea and the bearing of recent investigations upon the problems of supply. Jl. Soc. Arts. 59: 401-409. Garstang, W. (1907 A). Letter to Vice-Chancellor, Leeds University. 30 May 1907. Archives, Bursar’s Office, University of Leeds. Garstang, W. ( 1907 B). Letter to L. C. Miall. 1 June 1907. Archives. Bursar's Office, University of Leeds. Garstang, W. (1907 C). Letter to Vice-Chancellor, Leeds University. 19 June 1907. Archives, Bursar’s Office, University of Leeds. Garstang, W. (1909 A). The natural history of the North Sea. Trans. Norfolk & Norwich Nat. Soc. 8: 5-14. Garstang, W. (1909 B). Report on Experiments with marked plaice during 1904 and 1905; Fishery and Hydrographical Investigations in the North Sea and Adjacent Waters (Southern Area), 153-224. 4th Report Cd. 6125 HMSO. London. Pari, papers 1913 xxviii, 153. Garstang, W. (1909 C). The disappearance of the plaice. Naturalist. 403-407. Garstang, W. (1909 D). The distribution of the plaice in the North Sea, Skagerrak and Kattegat according to size, age and frequency. Rapp. P-v. Reun. Cons. perm. int. Explor. mer. 11: 65-135. Garstang, W. (1918). Diary of France (Alister Hardy papers). Garstang. W. (1919 A). The Return to Oxford. A Memorial Lay. Blackwell, Oxford. Garstang. W. (1919 B). Nature and Man. Naturalist. 89-96. 123-124. Garstang. W. (1919 C). Songs of the Birds. Naturalist. 195-198, 231-233. Garstang. W. (1920). The Ring Ousel's Rondo. Naturalist. 158. Garstang. W. (1922 A). Songs of the Birds. Bodley Head. London. Garstang, W. (1922 B). The Students' Opera. Jackson. Leeds. Garstang. W. (1922 C). The theory of recapitulation: a critical re-statement of the Biogenetic Law. J. Linn. Soc. (Zool.). 35: 81-101. Garstang, W. (1926 A). Plaice in the North Sea. The Times. 21 and 26 April 1926. Garstang, W. (1926 B). Wordsworth's interpretation of nature. Nature. Lond. 117: Supp. 1-8. Garstang, W. (1928). The morphology of the Tunicata. and its bearings on the phylogeny of the Chordata. Q. Jl. Microsc. Sci. 72: 51-187. Garstang, W. (1929 A). The origin and evolution of larval forms. Rep. Br. Assn. Advmt. Sci. Glasgow: 77-98. 52 Walter Garstang (1868-1949): Zoological Pioneer and Poet Garstang, W. (1929 B). Natural selection. Nature, Lond. 124: 410. Garstang, W. (1930). The Buckland Lectures, First Series for 1929. Fishing News, Aberdeen. Garstang, W. (1934). Report on the Tunicata. Pt. 1. Doliolida, 195-251. British Antarctic (Terra Nova) Expedition 1910. Natural History Reports Zoology IV. British Museum, London. Garstang, W. (1935). Songs of the Birds, revised edition. Bodley Head, London. Garstang, W. (1938). Letter to Lucy Garstang from Marine Biological Station, Bermuda, 15 July 1938 (Alister Hardy Papers). Garstang, W. (1946). The morphology and relations of the Siphonophora. Q. Jl. Microsc. Sci. 87: 103-197. Garstang, W. (1951). Larval Forms and other Zoological Verses. Blackwell, Oxford. Garstang, W. and Georgeson, E. (1935). Report on the Tunicata. Pt. II. Copelata, 263-282. British Antarctica (Terra Nova) Expedition 1910. Natural History Reports Zoology IV, British Museum, London. Gosden, P. H. J. H. and Taylor, A. J., eds. (1975). Studies in the History of a University. Arnold, Leeds. Gould, S. J. (1977). Ontogeny and Phytogeny. Harvard University Press. Gray. J. (1972). G. P. Bidder. Dictionary of National Biography 1951-1960: 111-112. Green, V. H. H. (1979). The Commonwealth of Lincoln College 1427-1977. University Press, Oxford. Gregory, R. A. (1922). Letters to Garstang, 11 and 22 May 1922. Gunther, A. E. (1977). The Life of William Carmichael M'lntosh. Scottish Academic Press. Gurney, S. G. (1970). From Generation to Generation. The Story of the Garstangs of Lancashire. The Research Publishing Company, London. Hardy, A. C. (1950). W. Garstang. Proc. Linn. Soc. Lond. 162: 99-105. Hardy, A. C. (1951 A). W. Garstang. J. mar. biol. Ass. UK. 29: 560-566. Hardy, A. C. (1951 B). W. Garstang. Journal du Conseil perm. int. explor. mer. 17: 7-12. Hardy, A. C. (1956). The Open Sea, Its Natural History: Pt. 1. The World of Plankton. New Naturalist Series, Collins, London. Hardy, A. C. (1973). Lecture on 1 March 1973 at Leeds University (Alister Hardy papers). Henson, H. (1949). W. Garstang. Naturalist. 49-50. Henson, H. (1974). Tape recorded conversations with staff of the Zoology Department, Leeds University. Hickson, S. J. (1927). F. W. Gamble. Proc. R. Soc. Lond. 101: xxv-xxviii. Huxley, J., Hardy, A. C. and Ford, E. B. (1958). Evolution as a Process, 2nd edition. Allen & Unwin, London. Jenkins, E. W. (1979). From Armstrong to Nuffield. Murray, London. Kelly, T. (1950). Outside the Walls. 60 years of University Extension at Manchester. 1886-1946. University Press, Manchester. Kemp, S. (1943). Edgar Johnson Allen 1866-1942. Obit. Not. Fell. R. Soc. Lond. 4: 357-367. Lloyd, L. (1933). Professor Garstang. The Gryphon , New Series. 14: 217. Lloyd, L. (1949). Walter Garstang. University of Leeds Review. 1: 233-240. MacBride, E. W. (1914). Textbook of Embryology. Invertebrata. London. MacBride, E. W. (1926). The Recapitulation Theory. Science Progress. 20: 461-474. MacBride, E. W. (1929). Natural selection. Nature, Lond. 124: 689. Marshall, A. M. (1893). Testimonial Letter to Garstang. McGill University, Montreal, Application for Chair. 26 July 1893 (Alister Hardy papers). Miall, L. C. and Fowler, W. W., eds. (1901). Gilbert White’s Natural History and Antiquities of Selborne. Methuen, London. Mill, H. R. (1902). First meeting of the International Council for the Exploration of the Sea. Nature, Lond. 66: 346-347. M’lntosh, W. C. (1899). The Resources of the Sea, as shown in the Scientific experiments to test the effects of Trawling and of the closure of certain areas off the Scottish shores. Cambridge University Press. Poulton, E. B. (1890). The Colours of Animals. Kegan Paul, London. Poulton, E. B. (1893). Letter to Garstang, 27 September 1893 (Alister Hardy papers). 53 Y.N. U. Bryological Section: Annual Report 1983 Russell, F. S. (1972). Marie Lebour. J. mar. biol. Ass. UK. 52: 777-788. Sadler, M. (1921). Testimonial letter in Garstang application for Oxford Chair, 2 July 1921 (Alister Hardy papers). Schlee, S. (1973). A History of Oceanography. Hale, London. Thompson, J. (1886). The Owens College. Cornish, Manchester. Totton, A. K. (1948). Letter to Garstang, 10 March 1948 from British Museum. University of Leeds (1907 A). Minutes of Council Committees 9. Committee on Biological Chairs, 22 May. Archives, University of Leeds. University of Leeds (1907 B). Minutes of Council Committees 9. Committee on Biological Chairs, 20 June. Archives, University of Leeds. University of Leeds (1909). Sixth Report 1908-09, Faculty of Science. University of Leeds (1910). Seventh Report 1909-10, Faculty of Science. Walmsley, L. (1969). So Many Loves. Collins, London. Young, J. Z. (1981). The Life of Vertebrates , 3rd edition. Clarendon Press, Oxford. Y.N.U. BRYOLOGICAL SECTION: ANNUAL REPORT 1983 T. L. BLOCKEEL 20 Heathfield Close, Bingley, BD16 4EQ In April the British Bryological Society held its spring meeting for 1983 in Ilkley and excursions were made to characteristic sites within the Dales. Many of the notable bryophytes of the region were seen, and although most of the localities were bryologically well-known, three species were added to the flora of VC 64. The autumn meeting of the Section was held at Skipwith on 3 September, principally for a visit to the Common. Fuller reports on both these meetings are being published elsewhere. Records Although a number of scarce epiphytes appear in the records which follow, there is no evidence that such species are recovering lost ground in Yorkshire. Most of them occur in small quantity and have a tendency to occur only on ancient tree boles, suggesting that at least some species have difficulty in colonizing younger trees. Marchantia alpestris: (63*) 44/13 Pavement outside the Parish Church. Bingley. July 1983; (64) 44/25 On path. Harlow Carr Gardens. Harrogate, July 1983. This taxon, formerly considered a variety of M. polymorpha. may prove to be common as a weed. Metzgeria fruticulosa: (64*) 34/77 On elder near the Viaduct. Batty Moss, Ribblehead. The segregate M. temperata appears to be commoner in western Yorkshire, though both species are rare. Calypogeia trichomanis: (64) 34/77 On peat, below Meregill Head, Ingieborough. July 1983. C. muellerana : (61*) 44/63 On wet peat. Skipwith Common. Sept 1983. Anthelia julacea: (64) 34/77 Still present on wet millstone grit rocks, with Campylopus atrovirens, at Meregill Head, Ingieborough. July 1983. Lophozia perssonii : (64) 44/42 On soft magnesian limestone in small ancient quarry between Ledston and Ledsham, near Castleford. Dec 1982. Barbilophozia hatcheri: (64*) 34/87 On limestone ledge on the high cliffs. Pen-y-ghent, M. E. Newton. Apr 1983 (B.B.S. Excursion). Jungermannia caespiticia: (64*) 44/16 On moist gritty soil on bank of stream. Upper Skell Gill, Naturalist 109 (1984) 54 Y.N. U. Bryological Section: Annual Report 1983 Sept 1983. This is a nationally rare species which was found to be not uncommon in parts of Calderdale by H. Walsh in the post-war years, but otherwise appears to be a great rarity in Britain. Marsupella sprucei: (64*) 34/76 On Silurian rock east of Wharfe village, Dec 1982; 34/78 In small quantity on millstone grit at c. 1750 ft alt., Force Gill, Whernside, Oct 1983. Cladopodiella francisci: (61*) 44/63 On peaty sand, Skipwith Common, Sept 1983. Cephaloziella elachista : (61*) 44/63 Skipwith, W. Ingham, 1899 (MANCH) (Bull. Br. bryol. Soc. 42 (1983): 48). Frullania fragilifolia: (64) 34/77 On sheltered rock face, Crummackdale, Feb 1983. Brachydontium trichodes : (64) 34/76 On Silurian rock east of Wharfe village, Dec 1982. Campylopus subulatus: (64) 34/67 Among Ingletonian rocks on grassy bank near Thornton Force, Ingleton, Feb 1983. Leucobryum juniperoideunv. (64*) 44/05 On millstone grit boulder near the Strid, Bolton Woods, T.L.B. and M. O. Hill, Apr 1983 (B.B.S. Excursion). Fissidens crassipes: (61*) 44/73 On stonework at water level, Derwent Bridge, Bubwith, Sept 1983. Fissidens celticus: (64*) 34/77 Bank of stream in ravine, Twistleton Glen, Ingleton, July 1983. New to Yorkshire. Tortula virescens : (64*) 34/96 Sycamore by the Village Green, Linton, Wharfedale, A. Newton, Apr 1983 (B.B.S. Excursion). Hyophila stanfordensis: (63) 44/41 On earth at base of limestone cliff, woodland north of Wentbridge church, H. L. K. Whitehouse, Apr 1983. Pottia recta: (64) 44/42 Magnesian limestone pasture between Ledston and Ledsham, near Castleford, Dec 1982. Barbula ferruginascens: (64) 34/77 Soil among rocks, Studrigg, Crummackdale, Feb 1983. Coscinodon cribrosus: (64*) 34/77 On dry slate rocks in old quarry, Twistleton Glen, Ingleton, T.L.B. and E. Ormand, Jan 1983, and with fruit, T.L.B. , July 1983. A rare species, previously known in Yorkshire only from the famous Mielichhoferia site in Cleveland. Discelium nudum: (64) 44/14 On clay-with-shale bank near top of Shipley Glen, Eldwick, Sept 1983. Tetraplodon mnioides: (64) 34/77 Plentiful and in fine fruit on old sheep dung, more rarely on bones, among grit boulders, Combe Scar, Whernside, June 1983. Bryum riparium: (65*) 34/69 Amongst dripping shaley scree on bank of stream running into Cautley Spout, 580 m alt., west of Cautley Crag, A. R. Perry, 1969 (Bull. Br. bryol. Soc. 42 (1983): 54). New to Yorkshire. Zygodon conoideus: (64) 34/77 On elder, Clapdale, Dec 1982. Orthotrichum stramineum : (64*) 34/97 On sycamore by R. Wharfe, Hubberholme, Apr 1983; 34/96 on elder by roadside near Netherside Hall, Threshfield, May 1983. Orthotrichum rivulare: (64) 34/95 Tree root by R. Aire, Coniston Cold, Feb 1983. Ulota phyllantha: (64*) 34/74 Old willow by R. Ribble, West Bradford, Jan 1983; 34/96 On elder, riverside woodland, Netherside Hall, Threshfield, May 1983. Leucodon sciuroides: (64) 34/97 On old ash tree by R. Wharfe near Hubberholme, Apr 1983. Neckera pumila: (64) 34/97 Base of alder overhanging R. Wharfe, near Hubberholme, Apr 1983. Brachythecium salebrosum: (64*) 44/14 Wet Molinia dominated grassland, Baildon Moor, E. Newton and L. Warburton, Nov 1983 (Cliffe Castle, Keighley). Except where stated otherwise, the records are my own. An asterisk indicates a new vice-county record or an amendment to the Census Catalogue. Corrigendum The moss reported as Ditrichum pusillum in the report of the Y.N.U. Meeting at Bedale and Thorp Perrow (Naturalist 108 (1983): 153) has since been examined by several bryologists and the consensus of opinion is that it is a form of Dicranella varia. In particular it lacks the characteristic rhizoidal tubers of the Ditrichum. 55 THE STATUS OF THE MOUNTAIN HARE, LEPUS TIMIDUS , IN THE PEAK DISTRICT D. W. YALDEN Department of Zoology, University of Manchester M13 9PL Introduction There are three principal herbivores on the Peak District moorlands, domestic sheep, Red Grouse Lagopus lagopus and Mountain Hares Lepus timidus. Sheep are enumerated by the agricultural census taken in June each year; there were about 143,000 in the moorland parishes in 1976 (Yalden, 1981), though not all of these would be actually on the moorland, as some would be on lower pastures. The Red Grouse population has been estimated at about 10,000 pairs (Yalden, 1979). An earlier paper on the Mountain Hare, Lepus timidus, based on 163 sightings, described the distribution of the species, but made no attempt to estimate the population size (Yalden, 1971). As a result of continued fieldwork, it is now possible to attempt this, as well as re-examine the distribution. Methods Field survey records are assigned to 1 km squares of the national grid, with altitude and vegetation type being noted. The most important fieldwork has been carried out in March or April, when the hares are still white but the moorlands are often bereft of snow. To obtain one estimate of the population, the highest count obtained in each one kilometre square, at any time in the past sixteen years (1967-1982), has been taken as the population size in that square. In addition to my own fieldwork, Sheffield-based naturalists have been studying the species (Clinging, 1982; Clinging & Whiteley, 1980; Whiteley & Whiteley, 1976). A regular 'Mountain Hare Watch’ undertaken one day in March each year to survey the population along the eastern edges has produced higher counts than my own for some squares, and I have used the highest available figure. Another estimate has been obtained by calculating the average density per 1 km square, and multiplying that by the total number of squares in which the species has been recorded. It was noted that Mountain Hares tend to gather on steep rocky slopes (Yalden, 1971) and that this made it easier to count them. The converse of this is that hares on flatter moorland might be overlooked, and the population thus underestimated. In an effort to investigate this point further, on some of the field trips in the last three years I have been accompanied by a Labrador dog which certainly has a ‘good nose’ for hares and rabbits. Because of the regulations covering ‘Access Land', it has not been possible for the dog to roam freely. Results The present paper is based on 1090 sightings of my own. accumulated during 222 field days; Sorby Natural History Society (Sheffield) hare watches recorded between 58 and 161 hares each year. The distribution map resulting from this fieldwork (Fig. 1) is similar to the earlier one, but rather more extensive; in 1971, the species was recorded from 106 squares, whereas it is now recorded from 246 squares. The Mountain Hare has spread slightly further south down the eastern moors on Burbage Moor (cf. Whiteley & Whiteley, 1976), but the major change is the larger population in the north-west. As well as the two main areas of population mentioned before, covering the Hayfield-Kinder Scout area and the eastern edges (Yalden. 1971), there is now a third area around Arnfield and the Chew moors. Mountain Hares are now regularly seen, though they are sparse, on the Holme Moss-Black Hill plateau (cf. Yalden. 1971). Numerically, the eastern edges, between Derwent Edge and Outer Edge, continue to be much the best area for the species, with 60 per cent of the population located in the 10 km squares SK18. SK19 and SK29 (Table 1). There is still no evidence that Mountain Hares occur on Comb’s Moss. Goyts Moss. Danebower. or Eyam Moor at present, though all these areas have been revisited by myself and other naturalists. Since they were reported in these areas by earlier naturalists (Coward & Naturalist 109 (1984) 56 The Status of the Mountain Hare, Lepus timidus, in the Peak District O 1 FIGURE 1 Distribution and abundance of mountain hares, Lepus timidus, in the northern part of the Peak District, plotted by 1 km squares of the national grid. The heavy outline represents the boundary of the Peak District National Park, and the graticule the 10 km National Grid squares. 57 The Status of the Mountain Hare, Lepus timidus, in the Peak District Oldham, 1910; Stubbs, 1929; Hcwson, 1956), they must, as suggested earlier (Yalden, 1971), have become extinct locally — all these are rather isolated, southern, patches of moorland in the Peak District, and it would be difficult for Mountain Hares to colonize or recolonize them from further north. The population estimate derived by summating the highest count in each 1 km square is 735 animals. The species is, of course, highly clumped; in 112 squares, nearly half the total in which it was recorded, the highest count was only one hare, but conversely, 24 hares were seen in the most populated square, and there were 23, 22 and 19 hares recorded in three further squares (Table 1). The average number per square (for squares in which at least one hare was recorded) was 2.1, with the figure in different years ranging between 1.4/km: (in 1979 and 1980) and 3.3 km2 (in 1977). The number seen per day’s fieldwork (again, considering only days on which TABLE 1 Number of mountain hares Lepus timidus seen (a) in the 10 km squares (of the national grid) contributing to the Peak District (b) the frequency distribution of 1 km squares by the number of hares seen in them (a) 10 km grid squares (b) 1 km grid squares Number of hares Number of squares Square Hare numbers per square with that many hares SD 90 4 1 112 SJ 99 1 2 58 SE 00 59 3 18 SE 10 39 4 16 SK 09 104 5 12 SK 08 56 6 5 SK 19 274 7 5 SK 18 61 8 6 SK 29 115 9 2 SK 28 22 13 3 Total 735 hares 14 4 15 1 19 1 22 1 23 1 24 1 Total 246 squares at least one Mountain Hare was seen, and only years with at least ten days' fieldwork) varied between 2.2 (in 1980) and 9.4 (in 1975). with an overall average of 4.9 (fable 2). Multiplying the average number seen per square (2.1) by the total range (246 squares) gives a population estimate of 517, ie roughly 500. This method of estimating the population can also be used to give some idea of the variation from year to year. In 1979, only 45 hares were seen in 32 squares, a density of 1.4, suggesting a total population of 344 hares; conversely, in 1977, with a survey density of 3.3 hares/km2, the implied population was 819 hares (Table 2). The limited amount of fieldwork with the dog did not suggest that I was overlooking any large proportion of the hare population. Together we visited 48 squares, and saw a total of 84 Mountain Hares. The highest counts in those same 48 squares totalled 170 hares. 1 was only aware of the dog detecting three or four hares which I might have missed, an error of perhaps 5 per cent; conversely, my height allowed me to spot many more which she did not detect. Discussion Summing the maximum number of hares seen in each 1 km square suggests a total population of 735 Mountain Hares. Without attempting a mark-rclease-recapture programme, it is impossible 58 The Status of the Mountain Hare , Lepus timidus, in the Peak District TABLE 2 Variation in sightings of Mountain Hare Lepus timidus in the Peak District from year to year Year Total hares seen No. squares Mean No. per square No. days fieldwork Mean No. per day Popn. estimate 1967 1 1 1.0 1 1.0 246 1968 14 9 1.6 4 4.7 394 1969 15 11 1.5 7 1.7 369 1970 82 51 1.6 21 3.9 394 1971* 114 63 1.8 26 4.4 443 1972* 95 41 2.3 11 8.6 566 1973* 113 49 2.3 19 5.9 566 1974 35 18 1.9 13 2.7 467 1975* 178 55 3.2 19 9.4 787 1976 10 3 3.3 2 5.0 819 1977* 112 34 3.3 16 7.0 819 1978* 52 24 2.2 11 4.6 541 1979 45 32 1.4 17 2.6 344 1980 60 44 1.4 27 2.2 344 1981* 77 40 1.9 16 4.8 467 1982* 87 33 2.6 12 7.3 640 Total 1090 508 2.1 222 4.9 517 * years in which the eastern edges were surveyed adequately. to judge how reasonable this is as a population estimate, but one can attempt to evaluate the magnitude and direction of the errors. Firstly, this method must inevitably lead to an increase in the estimated population as fieldwork progresses; hares are seen in further squares, or higher numbers in some squares, so the estimate goes up, but never goes down. When I did the same analysis at the end of 1977, I estimated 622 Mountain Hares. Thus 735 is likely to be an overestimate. Furthermore, this method of estimation implies that the population is stable from year to year. It is also likely that the estimate is higher than it should be because the hares certainly move round in response to weather, particularly wind conditions, as Flux (1970) also reported. With a strong west wind, the hares are spread on the east side of the Derwent Edges, whereas in an east wind they gather in the rocks of the steep west-facing scarp slope. The same individuals could thus easily be counted in two, or more, squares; this could not happen undetected on any one day, but could easily happen between successive trips. It should also be noted that the figure refers essentially to the late winter population, because that is when the hares are easiest to count, and when the most successful fieldwork has been undertaken. Since most mortality occurs over winter (Flux, 1970), one would guess that the autumn population might be twice the size of the March one. The highest total of hares seen in any one year in my fieldwork was 178 (in 1975), and the highest count in the Sorby NHS counts was 161 (in 1977). Since it is not possible to cover the whole of their range in one year, these figures suggest that 200 is an absolute minimum population estimate. The median estimate, of about 500, suggested by multiplying the average survey density by the total number of squares in which the species has been seen, is perhaps the best one. This method of estimating the population also allows some judgement of the variation in population from year to year, but there are problems with this method too. Because one can only sample part of the range each year, the estimated density is bound to reflect the sampling. In particular, if the dense population on the eastern moors is not sampled, or only poorly sampled (as was the case in 1979), then the survey density, hence the population estimate, will be low. If the estimate of 735 is an overestimate because of the ‘stage-army’ effect, one might nevertheless expect the autumn population to be rather higher, perhaps 1000 individuals. The Status of the Mountain Hare, Lepus timidus, in the Peak District 59 Clearly, the species is much less common than the Red Grouse (10,000 pairs, perhaps 60,000 birds in August (Yalden, 1979)). As potential prey for various predators, and as consumers of heather, the two species differ in their ecological impact by an order of magnitude. The hares also have much less impact than sheep on the grazing. Summary The highest number of Mountain Hares seen in the Peak District in each of 246 one km squares over the period 1967-1982 has been summed to give a population estimate of 735 individuals. This number refers to the late winter, minimum, population. A minimum figure of 200, a median of 500, and an autumn, high, population of perhaps 1000 are suggested. Acknowledgements I thank Derek Whiteley for making available to me the detailed counts made on the Sorby Natural History Society’s ‘mountain hare watches’, and Mrs P. E. Yalden for accompanying me on much fieldwork. References Clinging, V. (1982). Report of a survey of colour changes of the Mountain Hare in the Peak District 1977-1981. Sorby Record. 20: 20-27. Clinging, V. and Whiteley, D. (1980). Mammals of the Sheffield area. Sorby Record, Special Series. 3: 1-48. Coward, T. A. and Oldham, C. (1910). The Vertebrate Fauna of Cheshire and Liverpool Bay. Vol 1. The Mammals and Birds of Cheshire. London. Flux, J. E. C. (1970). The life history of the mountain hare ( Lepus timidus scoticus) in north-east Scotland. J. Zool., Lond. 161: 75-123. Hewson, R. (1956). The mountain hare in England and Wales. Naturalist. 107-109. Stubbs, F. J. (1929). The alpine hare on the Pennines. Naturalist. 31-34. Whiteley, D. and Whiteley S. S. (1976). Mammal fauna of Sheffield. Part 1. Distribution and status. Sorby Record. 14: 4-31. Yalden, D. W. (1971). The Mountain Hare {Lepus timidus L.) in the Peak District. Naturalist. 81-92. Yalden, D. W. (1979). An estimate of the number of red grouse in the Peak District. Naturalist. 104: 5-8. Yalden, D. W. (1981). Sheep numbers in the Peak District. In: Moorland Erosion Study: Phase 1 Report (J. Phillips, D. Yalden and J. Tallis, eds.), Peak Park Joint Planning Board, Bakewell. BOOK REVIEWS Mammals of Britain and Europe by Richard Orr and Joyce Pope. Pp. 176 with numerous colour plates and b/w drawings in text. World Wildlife Fund & Pelham Books. 1983. £14.95. This is an account, species by species, of the biology of the mammals of Europe. The text provides information on such aspects as size, geographical distribution, feeding habits, behaviour, habitat, economic status, reproduction, and conservation status. In some instances several species are dealt with in one section, e.g. squirrels, bats, shrews, although the text clearly distinguishes between the species within the grouping. Finally, there is a short section on observing Europe’s mammals and tracks and droppings. The most striking feature of this book is the high quality and lavishness of Richard Orr’s excellent illustrations. There are many colour drawings of species which are supplemented by numerous small drawings, mainly but by no means exclusively in black and white. These illustrate details of anatomy, behaviour, habitats, locomotion, and general biology. This is artwork of a quality to be compared with Archibald Thorburn's illustrations to Millais’s work on British Mammals which appeared at the beginning of this century. Orr’s style is different, being more meticulous and detailed. The work as a whole contains within its relatively limited size a 60 Book Reviews profusion of illustration which gives the reader considerable insight into the behaviour and appearance of Europe’s mammals. The representation is generally very good, although here and there one has reservations. Are the fox and red squirrel really as red as this and don’t the wood mice faces look a little blunt? This is a work to be thoroughly recommended to anyone interested in or wanting to know more about European mammals. MJD The Northern Yellowstone Elk: Ecology and Management by D. B. Houston. Pp. xx + 474 with 85 figures. Macmillan. 1982. $48.00. The elk is better known to naturalists in this country as the red deer. The account is centred around ten years’ research by the author on the scientific management of elk in Yellowstone National Park. During the course of this work Houston reappraised some of the established ideas on elk management; he questions the need to cull as a method of preventing habitat deterioration, considers the past role of burning and advocates the reintroduction of the wolf into the Park. This is a refreshing, stimulating and scientifically penetrating account of scientific management of a wildlife resource. It is inevitably limited to a particular location, although some of the broader concepts put forward merit consideration in a broader context. The main text is a solid scientific study supported by no less than 245 pages of appendices. Surprisingly, there are few comparisons with the studies on the management of this species in Europe. A useful addition to the wildlife manager’s library. MJD Seabirds: an identification guide by Peter Harrison. Pp. 448 including numerous maps and line drawings, plus 88 pages of colour plates. Croom Helm. 1983. £15.95. This magnificent book, specifically designed for field identification of seabirds, is the result of the author’s researches over the past seven years, during which time he travelled extensively throughout the world, drawing, photographing and studying the world’s seabirds. In a brief introductory chapter, he discusses the key diagnostic features and main problems involved in identifying seabirds. He does not pretend that seabird identification is easy. The plates, which illustrate the main plumages of all species and distinctive subspecies, precede the main text, but are provided with facing captions summarizing the main features to be observed in the field; symbols indicate regions where the birds are likely to occur. The order is not strictly taxonomic, since similar-looking species are occasionally included on the same plate, and where this is not possible they are cross-referenced but not, unfortunately, by plate number. Some plates are over-crowded, e.g. Skuas, Plate 54, but nevertheless, they are still far superior to those provided in any other seabird guides. The Systematic section, which forms the main body of the text, is well arranged for use in the field, and contains such information as detailed descriptions of the various plumages, flight, habits and jizz, distribution and migration, and similar species. More awkward groups and identification problems are most competently handled. The maps, in a separate section at the end of the book, are well drawn and use different scales according to the range of a species. There are few criticisms to be levelled at this book, but perhaps the inclusion of some non-seabirds (e.g. Grebes resident in the Andes and Central Madagascar) is unnecessary, and the somewhat skimpy section on seaducks, which are well covered elsewhere, could have been omitted altogether. A few typographical errors have been detected, such as the text-reference on the plate illustrating Bulwer’s Petrel which is mis-referenced to Herald Petrel. However, these are only minor criticisms and the book will undoubtedly become a standard reference work, essential for anyone interested in seabirds. TSC A Key to the Adults of the British Ephemeroptera with notes on their ecology by J. M. Elliott and U. H. Humesch. Pp. 101, including numerous figures and tables. FBA Scientific Publication No. 47. 1983. Available from: Freshwater Biological Association, The Ferry House, Far Sawrey, Ambleside, Cumbria LA22 OLP. £4.50. As well as detailed keys to forty-seven mayfly species, complemented by helpful line drawings, there are sections on the general characters, classification, collection, ecology, behaviour, egg development, etc. and a useful reference list (145 items). 61 LICHEN FLORA OF THE WEST YORKSHIRE CONURBATION — SUPPLEMENT III (1981-83) M. R. D. SEAWARD School of Environmental Science, University of Bradford and A. HENDERSON Department of Plant Sciences, University of Leeds The programme of fieldwork continues, making the West Yorkshire conurbation one of the most intensively studied urban lichen floras in the world. Detailed monitoring is called for, since the flora is at present critically affected by changing environmental conditions brought about by atmospheric amelioration as noted by Seaward (1981). A complexity of factors is involved: changes in pollution regimes affect a wide variety of important lichen substrates in terms of their acidity. Implementation of the Clean Air Acts of 1956 and 1968 on the one hand has resulted in the occurrence of lower acidic levels in urban substrates of long in situ standing, brought about by rainwash and leaching (substrates of recent origin being influenced to a lesser degree), whilst on the other hand, outer suburban and rural substrates have experienced reverse effects due to acid rain caused by the more widespread dispersal of gaseous pollutants (Likens et al, 1979; Press et al. 1983). Acid rain needs fuller consideration in future analyses of lichen floras. Numerous examples of lichen recolonization of the ameliorated urban environment can be cited; for instance, foliose plants ( Hypogymnia physodes , Parmelia saxatilis and Physcia spp.), not seen on trees in inner Leeds for many decades, are now colonizing mature Fraxinus within the University campus (44/2934; see Henderson, 1984); and Lecanora muralis, here as in many other localities within the conurbation, is now colonizing siliceous substrates after generations of confinement to calcareous habitats. In many suburban roads and streets L. muralis extensively occupies the moister microhabitat provided by paving immediately below walls, fences, gates, and hedges. Its invasion is almost certainly assisted by diaspore propagule trapping on walls, etc. and by bird-lime enrichment of such sites, with bird transport of diaspores and subsequent downward rainwash. Young plants of Xanthoria parietinal aureola (the two species difficult to differentiate when young) can now be found as very loosely scattered chains of generally dull yellow-gold thalli. at numerous calcareous sites within the inner conurbation, penetrating much further into this area than they previously did, even as isolated occurrences (see Fig. 1). Improvements like these are not evident on substrates in areas which suffered most intensely from pollution in the past, for instance, trees and exposed walls in the eastern and southern parts of the conurbation. The geology of the conurbation (Seaward, 1975: p. 153) presents substrates vulnerable to acid rain with consequent effects on natural drainage systems and, hence, on aquatic lichens. It is of interest, therefore, to note the record of Verrucaria elaeomelaena (see below), which brings the number of aquatic lichens recorded within the conurbation in recent years to four; only one of these, V. hydrela, has been recorded from the inner conurbation. The record of Leptogium turgidum (see below) brings the number of gelatinous species known in the conurbation today up to four. Interestingly, all are on anthropogenic substrates: on old garden walls ( Collema crispum, Leptogium plicatile and L. turgidum) and on park or garden paths ( Collema crispum, and C. tenax vars ceranoides and tenax). Distributional data listed below refer to recording units given in Seaward (1978. Fig. 1 and Table 1). They include additions to the flora made during the past three years, together with additions and corrections to former records provided mainly by Mr P. M. Earland-Bennett and Dr C. J. B. Hitch. We are grateful to these and other fieldworkers far too numerous to mention, and to Dr B. J. Coppins for his confirmation/identification of more critical material. Naturalist 109 (1984) 62 Lichen Flora of the West Yorkshire Conurbation — Supplement III ( 1981-83) Acarospora fuscata (Nyl.) Arnold Add Q. A. smaragdula (Wahlenb.) Massal. Add G. Arthonia cf. exilis (Florke) Anzi Henderson, 1981. M. On shaded sandstone wall. (For a description of this plant, see Coppins (1983: p. 196), where it is listed as Catillaria melanobola.) ‘It is not A. exilis; it is usually on basic bark, is rather common in Britain and is also known from Denmark' (B. J. Coppins, in litt.). First record of this genus in the conurbation; A. spadicea is known from just outside the conurbation’s north-west boundary, at Hetchell Wood (44/3742; see Henderson, 1982). Bacidia melaena (Nyl.) Zahlbr. See Micarea melaena (Nyl.) Hedl. Bacidia subfuscula (Nyl.) Th.Fr. Earland-Bennett, 1976. E, M. On decayed moss, soil and stones of siliceous walls. Uncommon. Not dissimilar externally from the plant listed as B. sp. in Seaward (1978: p. 71), but having quite different spores. At Copley, Halifax, this plant is growing in a garden path on the vertical surface of a siltstone flag immediately below mortar. Two colonies occur in inner Leeds, one on the capstones of a millstone grit wall, Meanwood Ridge, the other on border edgestones in a nearby garden. B. sp. (See Seaward, 1978: p. 71.) Add M. Probably overlooked. Cladonia chlorophaea (Florke ex Sommerf.) Sprengel Add S. C. macilenta Hoffm. Add E. C. polydactyla (Florke) Sprengel Add W. ’ Evernia prunastri (L.) Ach. Delete (U), add U. On dead stump in forestry plantation. Hypogymnia physodes (L.) Nyl. Add O. Gradual recolonization of suburbia (cf. Seaward, 1981, Fig. 2): several thalli on Fraxinus (44/1435, 44/2934, 44/2835, and 44/2836), and on single Quercus (44/267246) and Ulmus (44/291353), and on thin peaty soil over spoil heap (44/264247). See also under Parmelia saxatilis. Lecania erysibe (Ach.) Mudd Add D. The form sorediata is noticeably more numerous throughout the conurbation. Lecanora albescens (Hoffm.) Branth & Rostrup Bolton, 1775. (A), (G), M, (T), (V). On calcareous gravestone (44/2740). Uncommon. L. atra (Huds.) Ach. Add M. L. dispersa (Pers.) Sommerf. First record = Shackleton and Hebden, 1893. L. muralis (Schreber) Rabenh. Autecological studies of this species, based mainly on interpretations of its performance in the West Yorkshire conurbation, continue. Recent work has been concerned with recolonization following air pollution amelioration (Seaward, 1982). L. saligna (Schrader) Zahlbr. Add M. L. stenotropa Nyl. Add B. In some habitats forms of L. polytropa can resemble this species extremely closely in external characters. Microscopic examination of apothecia is frequently necessary to particularize field determinations. Lecidella carpathica Korber Earland-Bennett, 1974. G. On asbestos-cement roofing of Holmefield Barracks, now demolished. Possibly extinct. 63 Lichen Flora of the Wes/ Yorkshire Conurbation — Supplement III (1981-83) Leptogium turgidum (Ach.) Crombie Add U. First modern record: on decayed moss and soil in niches of old roadside wall. Rare. Several small colonies, including fruiting thalli, along 20 m stretch of wall at Farnley Hall (44/3147). Micarea botryoides (Nyl.) Coppins Add G, H. There is also a terricolous record of this plant growing amongst Lepraria incana at Danefield (44/2144). M. melaena (Nyl.) Hedl. Add B. Fruitless plants with black-stalked pycnidia, most probably referable to this species, occur occasionally on deeply shaded rock overhangs and vertical faces (e.g. in an overgrown derelict quarry near Bramhope, and on a large outcrop boulder, the Hollies, Leeds). ‘There are, however, other species that produce such pycnidia, e.g. M. botryoides and M. misella' (B. J. Coppins, in litt.). M. prasina Fr. A mesoconidial state of this plant (with white pycnidia) has been found in woodland at Elland Park (44/1022), and at the Hollies, Leeds (44/2738). The microconidial and fruiting states occur quite frequently in the conurbation. Mycoblastus sanguinarius (L.) Norman Add M. FIGURE 1 West Yorkshire conurbation: distribution of Xanthoria parietina/aureola showing major inner limit (equivalent to c. 70 p,g/m3 mean winter sulphur dioxide level) and disjunct recent sitings within the urbanized area. Each spot does not indicate merely isolated occurrences, but more extensive colonization reflecting the general invasive progress of these plants following atmospheric amelioration. 64 Lichen Flora of the West Yorkshire Conurbation — Supplement III (1981-83) Parmelia glabratula (Lamy) Nyl. subsp. fuliginosa (Fr. ex Duby) Laundon Add M. P. saxatilis (L.) Ach. Add O. More frequent now on corticolous substrates (Acer, Alnus, Fraxinus, Salix, and Ulmus) on the edge of the conurbation (cf. Seaward, 1975, Fig. 28). With Hypogymnia phy socles on Fraxinus at the inner Leeds sites mentioned above; rarely saxicolous in the same neighbourhood. When young or depauperate, difficult to differentiate from the next species. P. sulcata Taylor Several small thalli on single Salix (44/124382). Peltigera spuria (Ach.) DC. Add H. Undoubtedly the commonest member of the genus in the conurbation. Phaeophyscia orbicularis (Necker) Moberg Numerous further records from corticolous substrates, particularly Acer. Physcia dubia (Hoffm.) Lettau Add V. Placynthium nigrum (Huds.) Gray Add U. On stone of old roadside wall. Ramalina farinacea (L.) Ach. Interesting occurrence on millstone grit wall in an industrial area of central Halifax reported by Henderson and Stewart (1983). Strangospora pinicola (Massal.) Korber Add Q. Thelidium incavatum Nyl. ex Mudd Delete (T), add T. An isolated occurrence of this calciphile on petrified moss in the acidic surroundings of Ogden Clough, near Halifax (see Ackroyd, 1897; Henderson, 1983). Trapelia coarctata (Sm.) Choisy Add D. On the increase throughout the conurbation. Verrucaria elaeomelaena (Massal.) Arnold Add U. First modern record. Aquatic, on siliceous boulder in shaded woodland stream. Rare. V. viridula (Schrader) Ach. Add Q. Xanthoria parietina (L.) Th. Fr. Increasing number of reports of this species (including X. aureola ) from within the conurbation (see Fig. 1). As a consequence of this work, the lichen flora of the West Yorkshire conurbation can be summarized as follows: 322 lichen taxa have been reported from the area within 20 km of the centre of the conurbation, of which 5 are doubtful in the absence of supporting herbarium material, at least 32 are extinct in the area, and 185 have been recorded during the present survey (October 1967-December 1983). References Ackroyd, W. (1897). Petrified moss at Ogden Clough. Halifax Nat. 2: 15. Coppins, B. J. (1983). A taxonomic study of the lichen genus Micarea in Europe. Bull. Br. Mus. Nat. Hist. (Bot.) 11: 17-214. Henderson, A. (1982). Hetchell Wood survey. I. Lichens of Hetchell Wood and environs. Leeds Nat. Club Sci. Assoc. Newsletter, no. 5, 78-85. Henderson, A. (1983). Three interesting Yorkshire lichens. Leeds Nat. Club Sci. Assoc. Newsletter, no. 8, 139-140. Henderson, A. (1984). Lichen Report. Leeds Nat. Club Sci. Assoc. Newsletter, no. 9. in press. Henderson, A. and Stewart, P. R. (1983). The occurrence of Ramalina farinacea (L.) Ach. on millstone grit in central Halifax. Naturalist 108: 109-110. Likens, G. E., Wright, R. F., Galloway, J. N., and Butler, T. J. (1979). Acid rain. Sci. Amer., no. 241, 39-47. Press, M., Ferguson, P. and Lee, J. (1983). 200 years of acid rain. Naturalist 108: 125-128. Book Reviews 65 Seaward, M. R. D. (1975). Lichen flora of the West Yorkshire conurbation. Proc. Leeds Phil. Lit. Soc. (Sci. Sect.) 10: 141-208. Seaward, M. R. D. (1978). Lichen flora of the West Yorkshire conurbation — Supplement I (1975-78). Naturalist 103: 69-76. Seaward, M. R. D. (1981). Lichen flora of the West Yorkshire conurbation — Supplement II (1978-80). Naturalist 106: 89-95. Seaward, M. R. D. (1982). Lichen ecology of changing urban environments. In: Urban Ecology (R. Bornkamm, J. A. Lee and M. R. D. Seaward, eds): 181-189. Oxford. BOOK REVIEWS The Ferns of Britain and Ireland by C. N. Page. Pp. xii + 447, with numerous b/w figures, maps, etc. Cambridge University Press. 1983. £40 hardback, £15 paperback. Despite several editions of the very successful Hyde and Wade’s Welsh Ferns, there has been no comprehensive treatment of the British pteridophytes as a whole this century. C. N. Page’s book is therefore most welcome in that it provides a modern and highly informative text with detailed accounts of all species of British ferns and fern allies; hybrids are covered, but to a iesser extent. Only two skeletal keys (chart and multi-access) are provided: identification data is therefore contained within the individual species accounts, which is only of help to those with a certain amount of prior taxonomic knowledge. The text description to each taxon contains detailed notes on identification, drawing attention to possible areas of confusion in identification, and excellent field notes, often based on personal observation. Each taxon description is supplemented by a full page of frond silhouettes, a useful feature in that they show a range of variation, but sadly for the most part they are poorly reproduced (not necessarily the fault of the printer, since photocopies were used as originals). Each description is also provided with a small distribution map, but where the distribution is scattered or localized a larger scale map would have been more helpful, especially as some excellent maps showing environmental factors which influence native pteridophyte range are provided in the introductory matter. (Unfortunately the key to the atmospheric pollution map is reversed, and the map does not incorporate Irish data which is available.) Other chapters and sections in the book contain: a taxonomic list of native species and hybrids, a glossary, botanical subdivisions of Britain and Ireland, altitudinal distribution of native pteridophytes, growing ferns from spores, conservation, further studies most needed, a bibliography (several citations in the text have been omitted), and an index, as well as the keys and environmental maps referred to above. Despite the shortcomings described above, and the high price of the hardback edition, the wealth of information contained in the book will make it a most valuable aid for the botanist. MRDS Biology of Nonvascular Plants by Hayden N. Pritchard and Patricia T. Bradt Pp. x + 550. with numerous line drawings and b/w plates. Times Mirror/Mosby College Publishing. St. Louis. 1984. £23.20. For many years, there has been no adequate single account of nonvascular plants for college and university students, and it was necessary to consult numerous w'orks to appreciate their range and diversity. Doyle’s Nonvascular Plants: Form and Function (entitled Nonseed Plants: Form and Function in later editions) showed a breadth of study, but although an excellent introduction, did not provide the necessary depth, and in any case is now' out of print. It is remarkable that two books filling this gap. that under review' and Nonvascular Plants: An Evolutionary Survey by R. F. Scagel el al, should be published during the past two years; not only do they cover almost identical ground but they also treat their material in a similar manner and are published in a similar (typically American) format. (Furthermore, the recent new edition of Bell and Woodcock's The Diversity of Green Plants provides yet another alternative.) A particularly good feature of Pritchard and Bradt’s book are the sections on economic importance and ecology, which accompany their accounts of each major plant group. The work, which is well illustrated throughout, also contains selected references at the end of each chapter as well as a long list of cited literature (nearly 1000 items) and a useful glossary and index. MRDS 66 Book Reviews Wild Orchids of Britain and Europe by Paul Davies, Jenne Davies and Anthony Huxley. Pp. xii + 256, plus 64 pages of full colour plates. Chatto & Windus. 1983. £9.95. Consistently excellent colour photographs of each species (alas, without scales), com- plemented by clearly presented species descriptions, form the major part of this book; the descriptions, which also include details of flowering, habitat and distribution, are concise but highly informative. This part is preceded by interesting general accounts of morphology, pollination, development, and ecology, as well as a key to genera, and is followed by two sections on searching for and photographing orchids. Under the section ‘In search of orchids’ the authors draw on their considerable knowledge gained from fieldwork throughout Europe to provide useful general guides to the orchids of Britain, western European countries, Scandinavia, Canary Islands, Madeira, the Mediterranean islands, Turkey, Syria, Lebanon, and Israel, but in order to thwart unscrupulous collectors exact locations of rare species have been omitted. In the case of Spain, one of the outstanding sites recommended (viz. Carratraca, p. 204) has sadly disappeared: the reviewer witnessed major roadworks here following the disastrous fires three years ago. Despite numerous good orchid books on the market, this work can hold its own with the best of them. Thoroughly recommended to accompany amateur and professional botanists on their travels at home and abroad. MRDS Dimensions of Darwinism: Themes & Counterthemes in Twentieth-Century Evolutionary Theory edited by Marjorie Grene. Pp. 336, including diagrams and tables. Cambridge University Press and Editions de la Maison des Sciences de L'Homme, Paris. 1983. £15. The names of the twelve contributory essayists, Richard M. Burian; Stephen Jay Gould; Antoni Hoffman Wiejska; William C. Kilmer; Bernard Norton; D. S. Peters; William B. Provine; Wolf-Ernst Reif; Bernhard Rensch; Rupert Riedl; John Maynard Smith; John R. G. Turner (‘biologists, historians and philosophers’) are indicative of the quality of this book. The material originated from a conference held in Bad Homburg, West Germany, in 1981, on the topic of twentieth-century evolutionary theory. The book is divided into four parts with between two and four essays in each: Part I. The Developing Synthesis; Part II. Mimetic Theory: Its Relation to the History of Evolutionary Biology; Part III. The German Paleontological and Morphological Tradition; Part IV. Some Contemporary Issues: The Synthesis Reconsidered. The diversity of the work makes detailed comment difficult; Grene’s thoughtful fifteen-page Introduction makes it largely superfluous. The book is well-produced and clearly laid out, and illustrates the diversity of evolutionary thought from the morphological ‘metaphysic’ of Riedl to the ‘neo-Darwinian orthodoxy’ of Maynard Smith. DJH Spires of Form: Glimpses of Evolution by Victor B. Scheffer with drawings by Gretchen Daiber. Pp. viii -I- 152; illustrated in black and white. University of Washington Press, USA. 1983. £11.85. Spires of Form is a book on animal evolution and natural history for the interested layman or young naturalist. Diversity, adaptation, resource partitioning, animal navigation, defence, sex, interspecific and intraspecific interactions, sociobiology, and the speed of evolution are among the topics presented briefly (e.g. sociobiology in forty-two lines), and often anecdotally: ‘When I once visited Chitty at his laboratory in England he was weighing the tiny adrenal and thymus glands of field mice . . The illustrations, which include line drawings and photographs reproduced from various sources, share pages with the text; there are forty-two, serving a decorative rather than an informative function. A portrait of Charles Darwin, mandatory in popular books on evolution, occupies pride of place. A bibliography for each chapter is provided under Reference Notes, and a list of thirty-one books with Scheffer’s assessments and recommendations (‘A pleasureful book’; ‘For scholarly readers’; ‘High-school or college undergraduate reading levels’ . . ., etc.) is provided For Further Reading. Bold headings within chapters and printers’ embellishments compartmentalize the text, and make this a book easy to leaf through profitably in leisure moments. niH 67 CHANGES IN THE STATUS OF THE STONECHAT IN THE SOUTH-WEST PENNINES DURING THE 1970s J. B. SYKES Department of Environmental & Geographical Studies, Manchester Polytechnic Introduction Throughout the first seventy years of this century, the Stonechat (Saxicola torquata) was an uncommon species in the south-west Pennines. In the earlier part of the century, Parker (1928) reported only three passage records in twenty years in Rossendale, while Coward (1910) regarded it as a small-scale passage migrant and occasional breeding species in the hills of east Cheshire. The position was very similar in later years; Oakes and Battersby (1939), Oakes (1953), Bell (1962), and Spencer (1973). During the 1970s, there was a marked increase in the number of Stonechats recorded, at all seasons, in the Pennines and Pennine fringes of Lancashire and Greater Manchester (Wolstenholme, 1975; Spencer, 1978). The present paper examines this increase and the subsequent decline and briefly discusses some possible reasons for these changes in status. The Study Area This is depicted in Fig. 1 and includes the western Pennine hills between the rivers Ribble, in the north, and Mersey, in the south. Topographically, the area is very varied, with altitude ranging from 15 metres asl in the south-west corner up to 570 metres asl at Pendle Hill to the north of Burnley. It includes many large industrial towns and for approximately the last 150 years it has suffered the effects of relatively high levels of air pollution. Methods Stonechat records for the years 1971-79 inclusive were obtained from three types of source: (i) records published in the various county and local bird reports covering the study area; (ii) correspondence and discussion with many individual ornithologists in the area: and (iii) extensive field survey of known and likely breeding localities, particularly from 1977 onwards. Results The available records are plotted in Figs. 2, 3, 4, and 5 in which each rectangle represents the study area shown in Fig. 1. Records on Fig. 2 are for April to September inclusive. This period spans the breeding season of the species in the study area but it may include a few passage birds, particularly in September. Fig. 3 includes all October and November records, this being the main period of autumn passage (Spencer, 1977). Fig. 4 shows all December and January records, which are here deemed to represent wintering individuals. Fig. 5 includes all records for February and March which, according to Spencer (1977). is the period of spring passage through the district. Some birds remained in one locality for parts of both the autumn and winter periods, as defined above; these birds are recorded on both Figs. 3 and 4. Likewise, some individuals are plotted on both Figs. 4 and 5. April to September Records Fig. 2 shows only occasional, mainly lowland, breeding before 1974 but then a rapid increase to a peak of seventeen confirmed breeding pairs in 1976. This level of population was maintained through the 1977 and 1978 breeding seasons, but there was a marked decline to only two confirmed breeding pairs in 1979. If the other birds recorded at this season are regarded as possible breeding birds, then the number of pairs possibly breeding in the study area rose to a peak of twenty-six in 1976. These figures are plotted in Fig. 6. which shows the same pattern for both the confirmed breeding and possible breeding curves. In view of the likely underestimation inherent in the survey methods used, the higher total of possible breeding pairs is considered to be a more realistic estimate of the population size. Naturalist 109 (1984) 68 Changes in the Status of the Stonechat during the 1 970s 0 10 km. FIGURE 1 Map of the study area. Changes in the Status of the Stonechat during the 1970s 69 Comparison with Fig. 1 indicates an association of breeding records with the 305 metre (1000 ft) contour, with some concentration across the centre of the study area from Anglezarke Moor in the west to the Burnley, Rossendale and Rochdale hills in the east. This contrasts with the data for the whole of the British Isles, presented by Fuller and Glue (1977), which shows 91 per cent of Stonechat nests lying below 122 m (400 ft). Fuller and Glue also showed that, nationally, the related Whinchat (Saxicola rubetra) has a clear tendency to nest at higher altitudes than the Stonechat. Observations made during the present study, however, have repeatedly shown that where the two species are breeding in the same locality, the Stonechat is at a higher altitude than the Whinchat. October and November Records Fig. 3 shows an increase in records at this season from 1973; before the increase in breeding season records of 1974. There was then a steady increase in numbers to a peak in 1978, followed by a sharp decline in 1979, when the species reverted to its 1971 status at this season. The records in Fig. 3 also show interesting variation in geographical pattern. In 1974, 1977 and 1978 there was a noticeable concentration of birds in the lower-lying south-west comer of the study area, particularly along the Mersey valley. In 1973, 1975 and 1976, however, this concentration was not apparent. In an area with so many active ornithologists, this difference is unlikely to be due to large changes in recording efficiency. Rather, it may represent the response of the dispersing Stonechat population to between-years variation in autumn food supply and availability in different parts of the study area. Further, Fig. 3 shows fewer autumn records in 1975 than in 1974, although there were approximately twice as many breeding season records in 1975 than in 1974. This seeming paradox can be similarly interpreted if the dispersing population in 1975 found much of its autumn food supply outside the study area. It may be significant in this connection that Frost (1978) noted over 100 reports of Stonechats in Derbyshire during the autumn and winter of 1975. December and January Records Fig. 4 shows a generally similar pattern of distribution to that given for October and November in Fig. 3. The increase in the number of winter records began, however, in 1972-73, a year before the autumn increase noted above and eighteen months before the 1974 increase in breeding season records (Fig. 2). As in autumn, there is between-years variation in the distribution of winter records and it is particularly interesting that the cold winter of 1978-79 produced more records from the north-eastern half of the study area than any other winter during the 1970s, with the exception of 1976-77. February and March Records The records for these months are plotted in Fig. 5, which displays several points of difference from Figs. 2, 3 and 4. Firstly, the increase in numbers was not noticeable at this season until 1976; two years after the increase in breeding season records (Fig. 2) and three years after the winter increase (Fig. 4). Secondly, in all the years studied, except 1976, there was a distinct concentration of records in the western half of the study area. Thirdly, although there were fewer spring records in 1979 than in 1978, the number of 1979 records was not markedly lower FIGURES 2. 3, 4, and 5 (overleaf) Location of Stonechat records within the study area. Each rectangle represents the study area shown in Fig. 1. x confirmed breeding record # male © female •P pair O sex not recorded. 70 Changes in the Status of the Stonechat during the 1970s G» O 1971 - • 88 ‘X XX XX - o 1 1 1 1974 - •