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Ce are ee Care ie mee eM gweds WH i i ee ht OT a Ty CT ce i ce Smears cena oe Led whi HARVARD UNIVERSITY LIBRARY OF THE GRAY HERBARIUM ca a ee eee -_ Digitized by the Internet Archive In 2014 https://archive.org/details/watsonia113bota ig an WATSONIA Botanical Society of the British Isles Patron: Her Majesty Queen Elizabeth the Queen Mother Applications for membership should be addressed to the Hon. General Secretary, c/o Department of Botany, British Museum (Natural History), Cromwell Road, London, SW7 5BD, from whom copies of the Society’s Prospectus may be obtained. Officers for 1976-77 Elected at the Annual General Meeting, 22nd May 1976 President, E. L. Swann Vice-Presidents, Mrs H. R. H. Vaughan, Dr W. T. Stearn, Dr C. T. Prime, Mrs B. H. S. Russell Honorary Generai Secretary, Mrs M. Briggs Honorary Treasurer, M. Walpole Honorary Editors, Dr G. Halliday, Dr N. K. B. Robson, Dr C. A. Stace Honorary Meetings Secretary, Mrs J. M. Mullin Honorary Field Secretary, Miss L. Farrell Honorary Membership Secretary, Mrs R. Hamilton All enquiries concerning sales of, or advertising space in, the Society’s publications should be addressed to D. H. Kent, 75 Adelaide Road, West Ealing, London, W13 9ED _ Watsonia, 11, 189-194 (1977). 189 Presidential Address 1976 NORFOLK AND BOTANY E. L. SWANN Today there falls to me both the privilege and the pleasure as a mere amateur to give you the Presidential Address. Since I joined this Society 37 years ago I have witnessed a five fold increase in membership with an enormous growth in technical skill and professional competence. Our publica- tions have brought us a well-deserved reputation and today we have attained international status as was clearly shown at the Cambridge Conference (1974) when more than 20 visitors and speakers came from continental Europe. Among the many disciplines, bewildering at times, frightening by their complexity, an amateur has to pick his way very carefully. Thinking about this Address I decided there was only one subject on which I could perhaps claim some familiarity and that was my own county of Norfolk in which I have botanised for some fifty years. When I attend meetings of this Society at Burlington House I see the walls of the Linnean Society adorned with pictures of famous botanists, several of whom were born in Norfolk. But for one, there would probably have never been a Linnean Society, certainly no Linnean collections in London. I refer, of course, to Sir James Edward Smith; others are Sir William Joseph Hooker, Director of Kew Gardens for 24 years, and John Lindley, the father of orchidology, secretary to the Royal Horticultural Society for nearly 40 years and the man mainly responsible for saving Kew Gardens for the nation. If you will bear with me for a time I should like to try and find a reason, some explanation for the outstanding reputation of early Norfolk botanists and why they were leaders of scientific thought in the eighteenth and nineteenth centuries. It may be a flight of fancy but I believe the answer is sheep. It was during the reign of Edward III (1327-1377) that Flemish weavers were persuaded to set up their looms in and around Norwich. The abundance of local supplies of wool laid the founda- tion of the staple industry and Norwich became the second city of the realm. According to repute, the Flemings and the Walloons introduced many new or little-known plants such as the lilac, Provence rose, asparagus, carrot, celery and cabbage. Their gardens were walled-in enclosures but it is surely not difficult to picture them bringing in other plants from the country around Norwich when they were conscripted each year to help with the harvest; nor is it surprising that the taste for gardening spread to the local inhabitants and with it the study of wild flowers. Proof of this view comes from Ray’s Flora of Cambridgeshire, recently translated (1975) by Mr A. H. Ewen and Dr C. T. Prime: ‘Among we English the people of Norwich excel in their enthusiasm for the cultivation of beautiful flowers: they celebrate each year with a Festival of Flowers and award a prize of a floral wreath for the most neatly arranged flowers. For this reason they boast that Norwich should be called either a town 1n a garden or a garden in a town.’ It is to Sir Thomas Browne, the famous physician, mathematician, theologian and alchemist — an intellectual giant if ever there was one — that we owe two of our earliest plant records. He spent the last 45 years of his life in Norwich and in 1668 sent a box of the Sweet Flag (Acorus calamus) to one of his London friends. This plant, by reason of its aromatic leaves, was used for strewing on the floors of churches to counteract the somewhat less pleasant smells of the crowds of worshippers. Large quantities were sent to London and here must be the first awakening of conservation because the raiding of local supplies aroused the anger of the natives. Sir Thomas also recorded the Sea Holly (Eryngium maritimum) from Great Yarmouth in the same year, writing that ‘Ringstones, a small black and white bird like a wagtayle . .. common about Yarmouth sands. They lay their egges on the sand and shingle about June, and as the eryngo diggers tell mee, do not set them flat, but upright like egges in salt’. Of Benjamin Stillingfleet, R. W. Ketton Cremer (1944) in his Norfolk Portraiis writes: A 190 E. L. SWANN ‘A few people in every century are destined, whatever their real achievements and distinctions, to be remembered by posterity for some irrelevant reason. A peculiarity of manner, speech or dress, the responsibility for some popular innovation, a spectacular success or mishap—every trick of association has contributed to swell the ranks of this goodly company. This was the case with Benjamin Stillingfleet, a man of learning and cultivation who advanced the progress of knowledge and taste in a variety of modest ways, like all good botanists, but he is remembered because he wore blue stockings of worsted made from Norfolk wool in the drawing rooms of learned ladies and was the unconscious originator of an universal phrase.’ It was whilst staying with his friend Robert Marsham at Stratton Strawless that he kept careful records of natural history, which he published in 1761 as his Calendar of Flora and it ran to four editions. It contains no fewer than 250 early plant records. It was Sir James Edward Smith who became the most famous of Norfolk botanists. Born in Norwich in 1759 and living until 1828, he acquired a European reputation. From his earliest years he was interested in botany; in his letters he records ‘I can just remember tugging ineffectually with all my infant strength at the tough stalks of the wild succory on the chalky hillocks about Norwich’. At the age of 18 he began the study of botany as a science and the only book he could then procure was John Berkenhout’s Outlines of the Natural History of Great Britain and Ireland, published in 1770 in three volumes. Then he was shown the works of Linnaeus and writes ‘Never shall I forget the feelings of wonder excited by finding his whole system of animals, vegetables and niinerals contained in three octavo volumes’. This obviously refers to one of the early editions of Systema Naturae, but it was the first edition of Species Plantarum (1753) which gave added impetus to his career. Although his father wanted him to join the family business of woollen and silk merchants, Smith was determined to make botany his life’s work and the only way possible was to take up medicine. He went to Edinburgh because it was then the only place where medicine was combined with botany. He studied under Dr John Hope, the first teacher of the Linnean system in Scotland. Two years later, in 1783, Smith went to London to continue his medical studies and he spent many happy hours with Sir Joseph Banks; it was Banks who persuaded him to buy the collec- tions of the great Linnaeus. Smith’s father advanced the money, the total cost being £1088.5/-. So, the possessions of Linnaeus were brought to England by a Norwich man and purchased with Norwich money. In 1788 Smith and some of his friends founded the Linnean Society and Smith remained its President until his death in 1828, fourteen years after being knighted. When he died, the Linnean collections were still at his house in Norwich. It had been hoped that the President would bequeath them to the Society, but this was not to be. After long deliberations they were sold to the Society for 3,000 guineas, and as a result the Linnean Society was very short of funds for years afterwards. Smith’s help was sought in many ways and he did not always get the credit he deserved. In 1790 Sowerby asked him to help in publishing coloured plates of British plants and Smith wrote: ‘I readily undertook the letter-press of this work, and it came out in monthly numbers, under the title of British Botany. . .. The flippancy with which everybody quotes ““Sowerby”’ whom they know merely as the delineator of these plants, without adverting to the information of the work or the name of the author, leads me to the mortifying conclusion that all I have done is of little avail, except to the penetrating eye of the scientific few, who stand less in need of such assistance. But with their approbation I am conscious I ought to be content.’ Smith had the gift of encouraging others from all walks of life to take up botany: Dawson Turner, banker, who became an authority on algae; John Pitchford, surgeon and apothecary, whose favourite genera were Carex and Mentha (like most of the Norwich school of botanists he con- tributed to English Botany and it was he who discovered Holosteum umbellatum and Peucedanum palustre) and James Crowe, landowner of Lakenham near Norwich, who was closely associated with Smith for many years. Together they paid particular attention to the willows, many of which were grown in Crowe’s large garden. The foundation for the classification of this puzzling genus was laid by Smith, and salicologists following him were very unwilling to differ from one who had devoted so much time to their study. Unfortunately, he did not believe in the existence of hybrids, writing ‘seedlings innumerable, springing up all over the ground, were never destroyed till their species — were determined, and the immutability of each verified by our joint inspection. This was the more ——— aa -_ oe ™—— ee PRESIDENTIAL ADDRESS 1976 191 material, to set aside the gratuitous supposition of the mixture of species, or the production of new, or hybrid ones, of which, no more than of any changes in established species, I have never met with an instance’. Smith’s influence extended over many years and as late as 1926, in the Botanical Exchange Club Report of that year, James Fraser stated ‘In many cases today, Smith’s species remain where he left them at the beginning of the nineteenth century except that the “x” denotes that they are hybrids and not species’. It must be one of life’s ironies that Smith’s name should be commemorated by one of our most frequent hybrids, Salix x smithiana, the cross between S. cinerea and S. viminalis. Other names come to mind such as Lilly Wigg, in turn shoemaker, schoolmaster and bank clerk (his was the first record for Trifolium suffocatum in the British Isles), and the Rev. Charles Sutton, for whom the Linnean Society published the monograph on the genus Orobanche in 1789. Another clergyman was the Rev. Joseph Forby, who was also interested in willows, and Smith acknowledged his indebtedness to him by describing one of Forby’s discoveries as Salix forbyana, the Fine Basket Osier. What would Smith have said were he to be told that this willow seems likely to be a ternary hybrid, a cross between S. cinerea, S. purpurea and S. viminalis! The most famous of the Norwich school was undoubtedly Sir William Joseph Hooker, who was born in Norwich, and it was from Smith that he received the bias which determined his career, ultimately becoming Director of Kew Gardens and the author of some 50 volumes of descriptive botany. These are but a few, a very few, of the many Norfolk botanists and it cannot be denied that we have a very rich heritage. What of the county itself? It is a self-contained unit for there is but one place where you may enter without having to cross a river bridge, and that is at South Lopham where the River Waveney rises on one side of a small road and the Little Ouse on the other. The series of ecologically important habitats provide a rich flora ranging from the shingle ridges, sand-dunes and salt- marshes bordering its 87 miles of coastline, to the Broads, and to that part of Breckland to be found in south-west Norfolk. There are also sandy heaths and commons, semi-natural woodland, cal- careous valley fens (for chalk is the backbone of the county), bogs, a few remaining chalk downs, rivers, dykes and ponds. But man has so dominated and exploited his environment that it is almost unbelievable that the greater part of Norfolk was formerly one vast sheep-walk. The vegetation is almost all artificial and the only remaining natural habitats are the sea-coast and its bordering salt-marshes and sand-dunes. Human activities have been the greatest factor for change. Fields are no longer allowed to lie idle, arable land replaces much of our woodland and scrub, hedges are uprooted, fens drained and dykes filled in. Large areas of salt-marsh have been reclaimed whilst new roads, airfields and building operations have all taken their toll. It would appear that the lot of the present-day botanist is a SOrry one. There are still rewarding places to be found in Breckland, although much of its area is given over to dense conifer plantations. It is in this area that neolithic man came to settle and, as W. G. Clarke, the naturalist who first coined the word ‘Breckland’, points out in his classic In Breckland Wilds (2nd ed., 1937): ‘The important neolithic flint mines at Grime’s Graves; the flint implements scattered by the million over the surfaces of its heaths and arable fields, its extensive mileage of primitive trackways; its numerous barrows and other relics of early settlers, all indicate that in some of the prehistoric periods it was one of the most thickly populated districts in the country. These invaders of early man from the Continent initiated forest clearance of the native pine and birch and by a slow process stock-keeping by nomad hunters evolved into some primitive agriculture.’ Then followed a long period of change and decay with the displacement of the primitive people. Pollen analyses have shown that this was probably due to climatic changes and for many hundreds of years Breckland remained a vast sandy heath. The greatest numbers of Norfolk Enclosure Acts received the royal assent between the years 1799 and 1815. Land was enclosed; trees were planted, chiefly Scots pine, many of which, gnarled and deformed by ‘the wind on the heath’, are still to be seen. Thus Breckland has been subjected to a complete cycle of changes: first native forest, then clearance, and today we have one of the largest of modern forests created by the Forestry Com- mission who started their planting in 1922. 192 E. L. SWANN It was suggested that the lot of the present-day botanist might be thought to be a sorry one, but is this really true? We have become so accustomed to the publicity about losses in plant-life, and loud have been the laments about the effects of pollution. Maybe we have been more fortunate in Norfolk but our losses of truly native species do not exceed 5%, whilst the number of species recorded has shown a remarkable increase. To assess the changes in our flora it is necessary to have comparisons, and fortunately we have three Floras published at approximately 50 years’ intervals together with four Supplements. The first Flora of Norfolk was published in 1866 and listed 1,030 species ; it was the work of the Rev. Kirby Trimmer, who held the livings of Burnham Norton and Stanhoe in West Norfolk. The second appeared in 1913 and represented the labours of W. A. Nicholson, a bank clerk, who worked for some 30 years on this volume, which was published by the Norfolk and Norwich Naturalists’ Society and contained 1,208 species. In 1968 the third Flora was © published to coincide with the centenary of that Society. Aided by a keen team of contributors, and — with the result of some 50 years of fieldwork, Dr C. P. Petch and the writer were able to raise the | number of species to 1,563, whilst the recently published Supplement has brought our total to | 1,701. Strangely enough we lost more native species prior to 1914 than since that date. No two botanists will ever agree about what constitutes a native species; there are some who consider that any plant which arrives and establishes itself by natural means in natural or semi-natural habitats without allowance for the time factor is a native. Thus, the American Willow-herb (Epilobium adenocaulon) would be called native, but this is absurd. My estimate of native status is based on distribution, both within and outside our area, nature of the habitat, and historical evidence. We appear to have lost five since Nicholson’s day. The Fen violet (Viola stagnina) was last seen in 1936; the Northern Shorewort (Mertensia maritima) disappeared from the shingle ridge at Blakeney in 1931; Mudwort | (Limosella aquatica) was last seen in 1916 and the Greater Broomrape (Orobanche rapum-genistae) in 1920. In spite of careful searching each spring we can no longer find Veronica verna in the Norfolk Breckland; there is a specimen in J. E. Lousley’s herbarium dated 1938 and he saw it at - Santon for a few years after. Three species thought to be extinct in 1968 have recently been refound: they are the Common Clubmoss (Lycopodium clavatum), the Bladderwort (Utricularia neglecta, now to be called U. australis) and Field Southernwood (Artemisia campestris). New discoveries have been the Dutch rush (Eguisetum hyemale), Wayfaring Tree (Viburnum lantana), Saw-wort (Serratula tinctoria) and Meadow Saffron (Colchicum autumnale). | The influx of aliens accounts for the greatest number of changes and in 50 years they have risen from 194 to 485. Whilst some are only casua's, others are becoming integrated into the flora. Already a few have shown an explosive increase such as the Oxford Ragwort (Senecio squalidus), American Willow-herb (Epilobium adenocaulon), Slender Speedwell (Veronica filiformis), and ‘Gallant Soldier’ (Galinsoga parviflora). There have been two main sources of introduction. During the Second World War our autumn botanising was considerably enlivened by the many carrot- field aliens from American seed sources under Lend-Lease arrangements. Of the 32 weeds recorded, the most frequent were species of Amaranthus (American Pigweed) and Cock-spur grass (Echinochloa crus-galli). Owing to the low winter temperatures few have persisted but, in 1976, Amaranthus bouchonii appeared in thousands along the peaty road verges of one of the Breck fens. The other source is the very rich rubbish-tip at Harford Bridges, near Norwich. Bird-seed aliens figure prom- inently and Canary Grass (Phalaris canariensis) now occurs on most of the tips throughout the county. Towards the end of its use as a tip, Harford was more or less dominated by Kochia densi- flora, related to the garden ‘Burning Bush’. The four-fold increase in hybrids from a mere 26 to 109 calls for some explanation. It would be invidious to claim that we are better botanists than our predecessors; a far more likely explanation is the considerable disturbance of the habitat for, as the American botanist Edgar Anderson points out in his book Jntrogressive Hybridization (1949): ‘The environment exacts a powerful control over the results of natural hybridization.’ Disturbed ground may well apply to the greater part of Norfolk today. Two species hitherto occupying their own separate ecological preferences may be brought together so that they more readily hybridize and the first generation of hybrids can be expected to survive if there are zones in which conditions as a whole are somewhat intermediate between that of the previous habitats. An example of this is the cross between the Red and White Campions. It usually has pink flowers and is fertile. The Red Campion is a true native of woodland on the better soils whilst the White Campion is more strictly a naturalised alien of arable land and waste PRESIDENTIAL ADDRESS 1976 193 places. The hybrid frequently occurs between woodland and arable land, for example road verges and hedgerows, an intermediate taxon occupying intermediate habitats. As observed by Dr P. F. Yeo in Hybridization and the flora of the British Isles (ed. C. A. Stace (1975)), ‘When any two species of Euphrasia alike in chromosome number occur together, hybrids are likely to be found’. Euphrasia confusa is the most abundant species in Norfolk, occurring on basic grassland, particularly in Breckland, chalk pits and forest rides, whilst E. nemorosa grows on damp soil where the vegetation is lush. On Foulden Common, a calcareous fen in West Norfolk, the hybrid is found in an intermediate zone between the chalky grassland and the fen proper. Individuals have been found to vary considerably suggesting introgressive hybridization; the 1975 Supplement deals more fully with this genus. Thanks to the help and encouragement given by Dr C. E. Hubbard, another Norfolk man, the grasses have been intensively studied as will be seen both in the 1968 Flora of Norfolk and its 1975 Supplement. Many hybrids occur, some supporting Anderson’s observations. In the loose sand along the foreshore there are extensive colonies of the Sand Couch (Agropyron junceiforme), whilst the Sea Couch (A. pycnanthum; A. pungens auct.) occurs on shingle-banks, sand-dunes and salt- marshes. The Hybrid Sea-Couch (A. x obtusiusculum) ‘frequently occupies a zone in the fixed dunes intermediate between that of A. j:mceiforme on the loose sand and of A. pycnanthum fringing the salt-marshes’ (C. E. Hubbard, Grasses, 2nd ed., 1968). When Couch or Twitch (A. repens) grows in arable land near the sea, the hybrid between this and A. pycnanthum is found occasionally along sea-banks. Much as we deplore the loss or reduction in numbers of many of the Breckland specialities, there are compensations by way of new plants. There would appear to be good evidence that the dense conifer plantations have brought about an increase in humidity and along some of the tracks and in the more open woodland new plants have arrived. At Cranwich there is a flourishing colony of Creeping Lady’s Tresses (Goodyera repens) growing in pine litter; in 1973 no fewer than 70 flowering stems were counted. In 1969, the Isle of Wight variety of the Green-flowered Helleborine (Epipactis phyllanthes var. vectensis) was discovered at Santon and in 1948 Dr A. S. Watt found a rare alien Skullcap (Scutellaria hastifolia) in a wood near Brandon; the colony has since increased considerably. In addition to the land utilised by the Forestry Commission, several thousand acres are owned by the War Department. This so-called ‘Battle Area’ has unwittingly become a large nature reserve, for no trees are planted, public pressure of necessity is reduced to a minimum, and flocks of sheep keep the grass well grazed, for these ‘four-legged grass mowers’ are the best and only way to conserve chalk grassland. In 1967, a rare grass, Matgrass Fescue (Nardurus maritimus) was found growing in two chalk pits on Sturston Warren, and from a similar habitat in 1975 Dr C. P. Petch detected it near West Harling. There must surely be other species awaiting discovery along the several hundreds of miles of tracks and rides in this region. Maybe, too, other species hitherto considered to be extinct will be refound like the Dutch rush (Equisetum hyemale). At the time of its discovery in 1974, Dr E. A. Ellis wrote a note in the Eastern Daily Press (16.2.1974) in which he expressed surprise that it had not been detected before, as it was so abundant for some 200 yards in a hedgerow. The reason is that, although conspicuous in winter and early spring when the associated shrubs are leafless, in summer and autumn it is completely hidden by a prolific growth of the Common Horsetail (E. arvense). A count of the shrubs occurring in every 30 yards’ length of the hedge showed that these averaged between five and six, so that, using the Hooper system for dating hedges, the result Suggested that it originated somewhere between 1200 and 1400, long before the Enclosures but coinciding with medieval expansion. The ground flora was characteristic of woodland and early maps show woodland present, the hedge being part of a boundary. In Smith’s day the Dutch rush was known as the Greater Rough Horsetail or Shavegrass and there is a note in his English Flora (1828): ‘That eminent chemist Sir Humphry Davy first detected a quantity of pure silex, or flinty earth, in the furrowed cuticle of this plant, which accounts for its power, as a file in polishing wood, ivory or even brass. This purpose it has long served in England, under the name of Dutch Rushes, being usually imported from Holland.’ In 1972, whilst looking for Potamogeton hybrids in the water of Langmere, one of the Breckland meres, a plant was found which had both flowered and was fruiting submerged in some 45 cm of 194 E. L. SWANN water. This was the very rare Alisma gramineum subsp. gramineum, the third British record; a week later it was found near Manea in Cambridgeshire in a similar habitat. Common to both these places are the large numbers of wild-fowl and there is little doubt that it was brought in by wild-fowl on migration from Denmark and the Baltic States where this species is native. There is evidence that seeds of many aquatic species can remain viable in the intestinal tracts of wild-fowl and the hard pericarps of this species of Water Plantain are admirably adapted for transport over long distances. It has been shown that in the course of 50 years the number of flowering plants and ferns recorded from Norfolk has grown from 1,208 to 1,701. Admittedly, modern segregates have added their quota, for example, we now have 85 microspecies of Taraxacum and vie with Kent for first place, but there are certainly no grounds for pessimism or dejection. In January 1976 we suffered from terrific gales and many thousands of trees were uprooted, and it will take several years for the Forestry Commission to clear away the damage; available supplies of timber far outstrip the demand. There are, too, many elms suffering from Dutch elm disease. To all this may be added the fact that landowners are having second thoughts about replanting, for they claim that under the prevailing Capital Transfer Tax the government’s demands amount to three out of every four trees planted. However, even this setback has its compensations for botanists for, with the opening of a hitherto closed canopy and larger areas of open woodland, there is every likelihood of dormant or long- buried seeds germinating thereby giving a richer ground flora. What of the future? The Norfolk Naturalists’ Trust was the first of its kind to be formed and we now have some 30 Reserves covering more than 5,000 acres. We enjoy the enlightened policy of many landowners, particularly the Earl of Leicester who has created a coastal ‘Green Belt’ at Holkham. There are many Sites of Special Scientific Interest, the responsibility of the Nature Con- servancy. With a rapidly expanding population and industrial development there will be additional claims for space, but doubtless we shall be keeping a close watch and employing every means avail- able for the conservation and preservation of our rich heritage. Watsonia, 11, 195-198 (1977). 195 Atriplex praecox Hulphers: a species new to the British Isles P. M. TASCHEREAU Department of Botany, University of Manchester ABSTRACT Atriplex praecox Hiilphers, a native species new to the British Isles, is described. This species is known from Greenland, Iceland, northern Norway and adjacent U.S.S.R. and on the Baltic coasts of Sweden, Finland and Estonia. Collections from Kylestrome and Ullapool in north-western Scotland confirm its presence in the British Isles. Its relationship to other members of the A. prostrata group is defined. A key to the A. prostrata group is provided and the presence of A. /ongipes Drejer in the British Isles is confirmed. THE ATRIPLEX PROSTRATA GROUP The Atriplex prostrata group comprises a number of partially interfertile and morphologically similar taxa found on the coasts of western Europe and elsewhere. It is represented in the British Isles by the following species: A. prostrata DC. (A. hastata auct.), A. glabriuscula Edmondston, A. longipes Drejer, and A. praecox Hiilphers. Atriplex prostrata, the most widespread member of the group, is commonly known as A. hastata L. (Tutin 1962, Aellen 1964, Jones 1975b); this name is a nomen ambiguum (Taschereau 1972, Jones 1975a). The name 4A. triangularis Willdenow, used by Taschereau (1972), Gustafsson (1973a, 1973b, 1974) and Jones (1975a), refers to the same taxon. Gustafsson (1975) has shown that the legitimate name for A. hastata auct. is A. prostrata Boucher ex DC. It is common throughout much of the British Isles and is the only member of the group also occurring inland. It is a com- ponent of inland salt-marsh vegetation and a transient colonizer of freshly disturbed soil. Atriplex glabriuscula is common to abundant in sand or shingle around the coasts of the British Isles except in areas disturbed by excessive grazing or heavy human recreational use. This extremely variable species can be reliably distinguished only by the following combination of characters: Bracteoles thick-spongy with their margins united to the middle; seeds mostly dark brown to black, ovate to orbicular, irregularly biconvex; radicle median and + strongly ascending, diameter 2-0— 4-0 mm. Edmondston’s (1845) original collection from Baltasound, Unst, Shetland, v.c. 112 is not extant. The neotype selected by Gustafsson (1975) is: Keiss, near Wick, Caithness, v.c. 109, 19.9.1930, leg. M. L. Wedgewood (K). Atriplex praecox is a coastal halophyte restricted to meadows with short vegetation and to shingle on exposed coasts, above the high-water fucoid zone but below the zones occupied by most other terrestrial halophytes. Its distribution is disjunct, with populations in Greenland, Ice- land, northern Norway and adjacent U.S.S.R., and on the Swedish, Finnish and Estonian coasts of the Baltic Sea. It is now known from two localities in north-western Scotland and is described in detail below. The identity of the British specimens has been confirmed by M. Gustafsson. Atriplex longipes is a coastal halophyte occurring in tall vegetation communities dominated by species such as Scirpus maritimus and Phragmites australis. Its area of distribution is primarily in southern Fennoscandia. Atriplex longipes was first reported in the British Isles by Hulme (1957), and later by Aellen (1964) based on data provided by Hulme. Jones (1975a) briefly described A. longipes and comparable plants from the upper zones of sandy shores in S. Somerset, v.c. 6, and northern W. Norfolk, v.c. 28. She tentatively referred these to A. /ongipes, noting that this species was less common in Britain than suggested by Hulme. A. longipes, now known with cer- tainty from a locality in south-western Scotland, is discussed in detail below. The above four species may be separated by the following key: 196 P. M. TASCHEREAU 1. Bracteoles spongy-inflated, margins united up to the middle; radicle up- pointing os . A. glabriuscula 1. Bracteoles thin, herbaceous or membranous, margins united only at the base: radicle out-pointing 2. Axillary bracteoles with stalks 5-20 mm _... aa a ae A. longipes* 2. Axillary bracteoles sessile or with stalks up to 3 mm 3. Lower leaves triangular-hastate, subcordate to truncate or broadly obtuse at base; basal angle exceeding 160° oe : A. prostrata 3. Lower leaves lanceolate to rhombic-ovate, cuneate to attenuate at base; basal angle less than 140° .. fs nel ea ay. - A. praecox * Conspicuously stalked bracteoles also commonly occur in A. longipes hybrids. The following diagnostic characters require explanation. Leaf size and outline show extreme variation from the base to the apex of the plants in this group. The most constant and characteristic leaves are those on the central axis in the middle portion of the plant. Leaves in this region, about half way between the base and the terminal inflorescence, are called ‘lower leaves’. Such leaves frequently drop off before the bracteoles and seed are fully mature. The position of the seed radicle and its direction are important taxonomic characters. The fruit is a nut enclosed by two herbaceous, thick or thin bracteoles. Surrounding the seed is a loosely or firmly attached membranous pericarp with the vestiges of the style situated at the top. The outline of the radicle becomes visible on the seed margin when the lower portion of the pericarp is teased away. The radicle may emerge from the middle portion of the seed margin (‘median’) or from the base of the seed (‘basal’). The radicle apex may be directed upwards towards the style vestiges (‘up-pointing’) or outwards at a right angle to the style axis (‘out-pointing’). ATRIPLEX PRAECOX HULPHERS Atriplex praecox can be separated from related species and hybrids by the following combination of characters: Plants small, commonly 8-10 cm high, erect or ascending, sometimes with the lowermost branches elongating and spreading, often reddish. Lower leaves lanceolate to rhombic-ovate, usually with one or a pair of small divergent to ascending basal lobes, otherwise mostly entire, cuneate to attenuate at base. Bracteoles rhombic-ovate, thin-herbaceous or membranous, entire, free to the base, often barely exceeding the frequently visible, bulging nut, sessile or with a minute stalk up to 3 mm. Seeds black, lustrous, biconvex, 1-2—2-:5 mm wide, with a sub-basal, out-pointing radicle; pericarp translucent, firmly attached to the seed. The following collections from Scotland represent the first records of this species from the British Isles: Kylestrome, W. Sutherland, v.c. 108, 4th September 1975, Taschereau 75-85 (MANCH) Ullapool, W. Ross, v.c. 105, 5th September 1975, Taschereau 75-89 (MANCH) The plants at Kylestrome occurred on shingle. A. praecox was the only species present in the small area examined and about 90% of the plants consisted of the reddish spreading form; the remaining plants, also spreading, were green. The plants at Ullapool were smaller and more erect. Here they formed a distinctive reddish zone amongst shingle just above the high-water fucoid zone in a region devoid of other terrestrial species. M. Gustafsson (in /itt. 1975) noted that the Ullapool specimens are very similar to those collected by himself in the northernmost parts of Norway. The fruiting period is August to September in north-western Scotland. Turesson (1922a, 1922b) regarded A. praecox as an ecotype of A. Jongipes and later (Turesson 1925) treated it as a subspecies: A. longipes Drejer subsp. praecox (Hiilphers) Turesson. Gustafsson (1975) followed Turesson in ranking A. praecox as a subspecies of A. longipes, but I prefer to treat it as a separate species. Throughout most of its distribution it is morphologically distinct and occupies a characteristic ecological niche. In the British Isles, at least, it is as distinctive as other taxa in the A. prostrata group. Gustafsson (1972) recorded that about 30% of his artificial hybrids between A. longipes and ATRIPLEX PRAECOX HULPHERS: A SPECIES NEW TO THE BRITISH ISLES 197 A. praecox were almost fully fertile; only populations of A. praecox from Norway gave rise to semi-sterile hybrids with A. longipes. Natural hybrids between A. praecox and A. longipes are reported as rare in the Baltic area of coastal Sweden (Gustafsson 1972, 1975), but they have not been reported from the British Isles. ATRIPLEX LONGIPES DREJER A. longipes can be separated from related species and hybrids by the following combination of characters: Plants erect to ascending with elongated lower lateral branches, up to about 80 cm high in tall vegetation. Lower leaves narrowly triangular with a pair of weakly developed out-pointing basal lobes, otherwise mostly entire, cuneate at the base. Bracteoles elongate-triangular to rhombic, thin, herbaceous, entire except for distinctly developed pointed basal lobes, surface often strongly reticulate-veined especially toward the base, margins united only at the base, the axillary bracteoles with conspicuous stalks 5—20 mm long. As mentioned above, this species has been reported from the British Isles by Hulme (1957) and Jones (1975a). I have examined Hulme’s collection in LIV, and Jones’s specimen in OXF. The material, particularly that of Jones, is strongly reminiscent of A. longipes but not sufficient to permit a certain identification. The presence of A. /ongipes in the British Isles has been confirmed, however, by my collection from a stand of Phragmites australis in the north-east of Wigtown Bay, Kirkcudbright, v.c. 73, 9th September 1975, Taschereau 75-97 (MANCH). The specimens were compared with Scandinavian material in LD and my determination has been confirmed by M. Gustafsson. The distribution of A. /ongipes in Britain is still to be resolved. ATRIPLEX GLABRIUSCULA x A, PRAECOX AND OTHER HYBRIDS Gustafsson (1975) produced artificial hybrids of varying fertility between A. praecox and each of the members of the A. prostrata group. In crosses between A. glabriuscula and A. praecox the mean male fertility value (F:) was 60-80% (Gustafsson 1973a). Gustafsson (1975) noted that the occur- rence of natural hybrids between these two species was rare in northernmost parts of Norway. Specimens presumed to be A. glabriuscula x A. praecox have been collected from Scotland: Tongue, W. Sutherland, v.c. 108, Ist September 1975, Taschereau 75-83B (MANCH); and Ullapool, W. Ross, v.c. 105, 5th September 1975, Taschereau 75-89B (MANCH). The specimens agree with Gustafsson’s (1975) description: stems prostrate to ascending, larger than in A. praecox, green or reddish; leaves + similar to those of A. praecox; bracteoles rhombic, much thicker than in A. praecox, margins united to the middle; radicle out-pointing, sub-ascending. These specimens represent the first record of this hybrid in the British Isles. Other natural hybrids in the A. prostrata group reported from Scandinavia by Gustafsson (1975) are: A. glabriuscula x A. longipes (rare in the western parts of Scandinavia with one record from Bodo in Norway); A. glabriuscula x A. prostrata (relatively rare in the western parts of Scandi- navia); A. longipes x A. prostrata (common along the west coast of Sweden northwards to the southern parts of Norway; relatively rare in Denmark; rare in the Baltic area. The taxon called A. longipes subsp. kattegatense Turesson is one nothomorph of this hybrid (Gustafsson 1973b)); A. praecox x A. prostrata (common in the northern parts of Norway). None of these has yet been found in the British Isles. ACKNOWLEDGMENTS This work was done while the author was the holder of a Postgraduate Scholarship from the National Research Council of Canada. I should also like to thank Dr M. Gustafsson for the valuable help he has given me. 198 P. M. TASCHEREAU REFERENCES AELLEN, P. (1964). Atriplex in TuTIn, T. G. et al., eds. Flora Europaea, 1: 95-97. Cambridge. EDMONDSTON, T. (1845). A Flora of Shetland, p. 39. Aberdeen. GusTAFSSON, M. (1972). Distribution and effects of paracentric inversions in populations of Atriplex longipes. Hereditas, 71: 173-194. GusTAFSSON, M. (1973a). Evolutionary trends in the Atriplex triangularis Group of Scandinavia, 1. Hybrid sterility and chromosomal differentiation. Bot. Notiser, 126: 345-392. GUSTAFSSON, M. (1973b). Evolutionary trends in the Atriplex triangularis Group of Scandinavia, 2. Spon- taneous hybridization in relation to reproductive isolation. Bot. Notiser, 126: 398-416. GuSTAFSSON, M. (1974). Evolutionary trends in the Atriplex triangularis Group of Scandinavia, 3. The effects of population size and introgression on chromosomal differentiation. Bot. Notiser, 127: 125-148. GusTAFsson, M. (1975). Evolutionary trends in the Atriplex prostrata Group of Scandinavia, 4. Taxonomy and morphological variation. Lund. (Prepublication of Opera Botanica, 39 (1976)). Hug, B. A. (1957). Studies on some British species of Atriplex. Ph.D. thesis, University of Edinburgh. JONES, E. M. (1975a). Taxonomic studies of the genus Atriplex (Chenopodiaceae) in Britain. Watsonia, 10: 233-251. Jones, E. M. (1975b). Atriplex, in STAcE, C. A., ed. Hybridization and the flora of the British Isles, pp. 185- 186. London. TASCHEREAU, P. M. (1972). Taxonomy and distribution of Atriplex species in Nova Scotia. Can. J. Bot., 50: 1571-1594. TURESSON, G. (1922a). The species and the variety as ecological units. Hereditas, 3: 100-112. TURESSON, G. (1922b). The genotypical response of the plant species to the habitat. Hereditas, 3: 211-350. TuRESSON, G. (1925). Studies in the genus Afriplex. Lunds Univ. Arsskr., N.F. Adv. 2, 21 (4): 1-15. TuTIN, T. G. (1962). Atriplex, in CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F. Flora of the British Isles, 2nd ed., pp. 281-283. Cambridge. (Accepted July 1976) Watsonia, 11, 199-204 (1977). 199 Lychnis alpina L. in Britain J. PROCTOR and W. R. JOHNSTON Department of Biology, University of Stirling ABSTRACT Soils from two British sites of Lychnis alpina L. were analysed and shown to have very different chemical composition. Plants from the two sites were grown on both soils under controlled conditions and the differing responses indicated that each site has a distinct edaphic race of the species. The restricted occurrence in Britain of L. alpina is discussed in relation to soil factors. INTRODUCTION Lychnis alpina L. (Viscaria alpina (L.) G. Don fil.) has two main sites in Britain, at Hobcarton Crag in the English Lake District (Cumberland, v.c. 70), and near the hill called Meikle Kilrannoch at the head of Glen Doll in Forfar, v.c. 90, Scotland. (A second Lake District site, near Coniston, Furness, v.c. 69b (Potter 1880), requires confirmation and, on the island of Rhum, Mid Ebudes, v.c. 104, plants recorded by Heslop-Harrison et al. (1943) need further investigation of their indigeneity and indeed their survival). There are about 150 plants at Hobcarton (Ratcliffe 1960) whilst there are very many more than this at Meikle Kilrannoch. It has been suggested (Raven & Walters 1956) that the Hobcarton site is associated with a pyritic vein, and the data of Ratcliffe (1960) support this to some extent. The site at Meikle Kilrannoch is a well-known serpentine area (Proctor & Woodell 1971). For both sites it was felt desirable to collect more soil data on both exchangeable and total quantities of elements in the hope that they would provide some explanation of the species’ restricted occurrence in Britain. SOIL ANALYSES Soil samples were collected from as close as possible to established L. alpina plants without dis- turbing them. Seven samples (HI-H6, H10) were taken at Hobcarton in August 1969 and five samples (MK1—MKS) from Meikle Kilrannoch in August 1975. Analyses were carried out on air- dried samples. pH was measured in a 1:1 soil:deionized water mix. Exchangeable cations were extracted with 1 M ammonium acetate adjusted to a pH value similar to that of the soils, i.e. pH 4-2 for the Hobcarton samples and pH 7 for the Meikle Kilrannoch samples, and analysed by atomic absorption spectrophotometry. In addition, subsamples were sent to the Macaulay Institute for Soil Research, Aberdeen, for semiquantitative spectrographic analyses of total quantities of elements. The analyses (Tables 1-4), which include three samples (H7—H9) from Ratcliffe (1960), clearly show that the British sites of L. alpina bear two very different soils. Judging from the data of Swaine (1955), there is little abnormal in the total quantities of elements in the soils at Hobcarton apart from the high manganese with associated cobalt in samples H1 and H2. The exchangeable cation analyses, however, suggest some further local soil abnormalities. In Ratcliffe’s samples (H7—H9), copper levels are very much higher than the ‘Up to c 1 ppm’ which Swaine (1955) gives as being water or dilute acid extractable in normal soils (Ratcliffe surprisingly regarded these copper levels as being ‘only in average concentration compared with most soils’). Our own analyses show exchangeable copper levels up to 3:5 ppm, and presumably indicate dif- ferent soil sample collecting points from those of Ratcliffe. Exchangeable manganese ranges from J. PROCTOR AND W. R. 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SI OI> SI OZ 0009 08 O1> 007 I EaIW OD = POOTS Gr-S: = 00e* O00'S Hct: Eok>c coz cor “ag F'009't. 1 OOl'T O% O1> SE O€ O009T 08 O1> OST I> ZW OOl O0O'T> OF O0€ 0009 OF O1> O£€ OF OOF 008 I> 006 Oc OI> O02 OF O00'%] 09> OI> OS™ I> IAW eee eee a ees ES Se IZ UZ xX A LL 2S bs os AE -4W IN OW (GN TT -¥T © tO59OS-313 2-2: Sa we ae an ojdueg eee eee eee a a ee eee HOONNVYTIIA ATAHIAW WOU “Tl VNIdTY SINHOXAT JO ANOZ ONILOOU AHL WOU STIOS NI SNOILVULNAONOD LNAWATA AOVUL IVLOL ‘'€ AIAVL 202 J. PROCTOR AND W. R. JOHNSTON 16 to 214 ppm, exchangeable iron from 5 to 254 ppm. These levels do not seem outstandingly high. For example, Gupta & Rorison (1975) found up to about 80 ppm exchangeable manganese and about 280 ppm exchangeable iron in a brown podzolic soil supporting undistinctive grassland in Derbyshire. The analyses for Meikle Kilrannoch show the characteristic features of serpentine soils (Proctor & Woodell 1975) and have high total levels of nickel and chromium and exchangeable nickel. The exchangeable magnesium/calcium ratio at this site is amongst the highest of British serpentine soils and has been demonstrated by Proctor (1971la, b) to be a major cause of the extremely toxic con- ditions there. ARE THERE TWO RACES OF LYCHNIS ALPINA IN BRITAIN? An important question is whether L. alpina is a plant of wide tolerance or whether it has evolved separate races in response to the differing edaphic conditions at each of the known British sites. This was investigated by growing plants from seed collected from both sites on both soil types. Soil and a small quantity of seeds were collected from Hobcarton and Meikle Kilrannoch in August 1975. The seeds were stored in laboratory conditions for approximately six weeks and then germinated, without pre-treatment, on filter papers. Seedlings were transferred to 8 cm diameter plastic pots of soils and two plants from the same site were grown in each pot. A randomized block experimental design was used, with five blocks each with one replicate of each treatment. John Innes Potting Compost No. 2 (J.I.) was used as a reference treatment. Plants were grown in the soils from 7th October 1975 to 12th December 1975 at 16°C and with 16 h supplementary light daily. The aerial parts were then harvested, dried at 85°C and weighed. The results (Table 5) were subjected to three separate analyses of variance because of very large differences in variance between plant weights on the three soil types. TABLE 5. MEAN DRY WEIGHT (mg)+S.E. OF TWO PLANTS OF LYCANIS ALPINA L. GROWN ON DIFFERENT SOILS Soils Seed source J.1. Meikle Kilrannoch Hobcarton Meikle Kilrannoch 292:8+ 116-0 4:2+0-42 5:2+6:4 Hobcarton 540-2+ 116-0 0-8+0-42 34-3+6°4 Both races of plants grew vigorously on J.I., without any symptoms of nutrient deficiency. Differences in dry weight between the two races fell short of statistical significance at the 5% level. In the Hobcarton soil, the Hobcarton plants grew well, whilst the Meikle Kilrannoch plants were very small, stunted and in some cases showed both chlorosis and reddening. Dry-weight differences were significant at the 5% level. In the Meikle Kilrannoch soil the Hobcarton plants were very small and chlorotic, in contrast to the larger Meikle Kilrannoch plants, which were healthy in appearance. Dry-weight differences were significant at the 1 % level. DISCUSSION The two known populations of Lychnis alpina in Britain occur on soils of very different chemical composition and the species has apparently evolved separate races in response to these. The soil samples from Hobcarton show some unusual features but none which would not be © expected at pyritic outcrops elsewhere in Britain. The flora associated with L. alpina there is of © commonplace species with the exception, noted by Ratcliffe (1960), of Grimmia atrata,a rare moss — which is reputedly particularly associated with copper-bearing rocks (Martenssen & Berggren 1954). — By contrast with the unremarkable Hobcarton soils, the samples from Meikle Kilrannoch reveala — LYCHNIS ALPINA L. IN BRITAIN 203 very unusual chemical composition at a site which is at a higher altitude (c 850m) than most other British serpentines. The flora associated with L. alpina there includes a number of very local species (Proctor & Woodell 1971). The abundance of L. alpina at Meikle Kilrannoch and its well-documented occurrence on serpentines elsewhere (e.g. Knaben 1950) suggested that tissues of this race of the species should be analysed to investigate if the plants accumulated abnormal quantities of any unusual elements. A small number of leaves of L. alpina (from soil sample collecting points MK 1-3) were digested in concentrated nitric acid and analysed by atomic absorption spectrophotometry. The results (Table 6) show no extreme accumulations of elements compared with some examples discussed by Proctor & Woodell (1975), and the concentrations are all within the ranges measured for other species at this site (Johnston unpublished). TABLE 6. ELEMENT CONTENT OF LEAVES OF LYCHNIS ALPINA L. FROM MEIKLE KILRANNOCH m-equiv. per 100 g dry tissue Sample No.* Ca Mg K Na MKI 57-7 137-1 187-2 17-3 MK2 42-0 147-7 174-5 11-5 MK3 57-7 216-2 148-8 23-0 p.p.m. dry tissue Sample No.* Ni Cr Co Fe Cu Mn Zn Al MKi 88-4 89-7 26-3 256°1 19-1 35-2 50-0 63-2 MK2 64:9 68-3 12-3 206-4 10-5 a7 47-0 66-6 MK3 70-8 93-3 16-9 270-7 6-3 42-2 47-0 63-2 * Numbered as in Tables 3 and 4. At Meikle Kilrannoch there is a puzzling difference in the numbers of L. alpina plants on the two main serpentine outcrops, which are less than 1 km apart. L. alpina is common on the northern outcrop, yet Proctor & Woodell (1971) reported the species to be absent from the southern outcrop. A search of the latter in June 1976 revealed a single specimen. The soil analyses in this present paper all refer to the northern outcrop. Preliminary comparisons of the soils of the southern with those of the northern outcrop (Proctor & Woodell 1971) revealed no gross differences in chemical composi- tion although we are currently subjecting the soils of the southern outcrop to a more critical Teinvestigation. It is curious that, in Britain, L. alpina should be restricted to certain pyritic and serpentine out- crops since elsewhere the species grows on a wide variety of preponderantly neutral to basic soils. Dr G. Halliday (pers. comm. 1976) pointed out, for example, that in southern Greenland the species is widespread and apparently shows no particular preference for heavy metal/ultrabasic soils. In view of this, and taking into account other observations such as the unremarkable nature of at least some of the Hobcarton soil samples, we believe that there are many sites in Britain which appear to provide suitable soil chemical conditions for L. alpina and that the factors limiting its distribution here are at present unknown. ACKNOWLEDGMENTS We should like to thank Dr R. L. Mitchell and staff at the Macaulay Institute for Soil Research, Aberdeen, for their analyses of total levels of elements in the soils, and Professor C. D. Pigott for his assistance at the Hobcarton site. The Nature Conservancy Council is acknowledged for per- | mission to work at Meikle Kilrannoch. 204 J. PROCTOR AND W. R. JOHNSTON REFERENCES GupTA, P. L. & Rorison, I. H. (1975). Seasonal differences in the availability of nutrients down a podzolic profile. J. Ecol., 63: 521-534. HESLOP-HARRISON, J. W., HARRISON, H., CLARK, W. A. & Cooke, R. B. (1943). Vascular plants from the Isle of Rhum (v.c. 104) and the Isle of South Uist (v.c. 110). J. Bot., 80: 113-116. KNABEN, G. (1950). Botanical investigations in the Middle Districts of Western Norway. Bergen. MARTENSSON, O. & BERGGREN, A. (1954). Some notes on the ecology of the ‘copper mosses’. Oikos, 5: 99-100. Potter, R. (1880). In Lees, F. A., ed. New county records. Reps botl Record Club, 1879: 53. Proctor, J. (1971a). The plant ecology of serpentine, 2. Plant response to serpentine soils. J. Ecol., 59: 397-410. Proctor, J. (1971b). The plant ecology of serpentine, 3. The influence of a high Mg/Ca ratio and high nickel and chromium levels in some British and Swedish serpentine soils. J. Ecol., 59: 827-842. Proctor, J. & WoOopDELL, S. R. J. (1971). The plant ecology of serpentine, 1. Serpentine vegetation of England and Scotland. J. Ecol., 59: 375-395. Proctor, J. & WOoDELL, S. R. J. (1975). The ecology of serpentine soils. Adv. Ecol. Res., 9: 255-366. RATCLIFFE, D. A. (1960). The mountain flora of Lakeland. Proc. bot. Soc. Br. Isl., 4: 1-25. RAVEN, J. & WALTERS, S. M. (1956). Mountain flowers. London. SwAINE, D. J. (1955). The trace-element content of soils. Tech. Commun. Commonw. Bur. Soils, 48. (Accepted July 1976) Watsonia, 11, 205-210 (1977). 205 A hybrid swarm between the diploid Dactylorhiza fuchsii (Druce) Soo and the tetraploid D. purpurella (T. & T. A. Steph.) Sod in Durham R. M. LORD and A. J. RICHARDS Department of Plant Biology, University of Newcastle upon Tyne ABSTRACT A study was made of the pollen grain meiosis and floral characters of a random sample of a population of diploid Dactylorhiza fuchsii (Druce) Soo, tetraploid D. purpurella (T. & T. A. Steph.) Sod and apparent hybrids in a limestone quarry in Co. Durham. Eudiploids, eutetraploids and a predominance of eutriploids were found, but about a quarter of the population were aneuploid, with chromosome numbers 2n = 44, 48, 52 and 72. A hybrid index showed that a wide range of hybrids occurred. Eutriploids, presumed to be F, hybrids, were very variable. They showed a wide range of univalents at meiosis, with a (rare) maximum of 20. The appearance of aneuploids was consistent with an F,, backcross, or subsequent origin. It is suggested that population differentiation in tetraploid Dactylorhiza may be due to polytopic allopolyploid origin. Viability of aneuploid gametes and hybrids may be due to ancestral polyploidy. The situation is unusual because extensive F, and backcross hybridization seems to occur across a diploid/tetraploid barrier. INTRODUCTION It has long been recognized that many of the taxonomic problems associated with the genus Dactylorhiza in the British Isles arise from hybridization (Stephenson & Stephenson 1922). Popula- tions containing many and varied morphological intermediates between the diploid D. fuchsii (Druce) Soé (2n = 40) and the tetraploids D. purpurella (T. & T. A. Steph.) So6é and D. praeter- missa (Druce) So6 (2n = 80) have been studied by Heslop-Harrison (1953, 1957). His main con- clusions (cited by Roberts (1975)) were that hybrids are eutriploids (2n = 60), although very variable, and show a high level of seed-sterility. He suggested that the few embryos which are produced arise parthenogenetically, being themselves eutriploid, and that backcrossing to parents or crossing between triploids is unlikely to occur, although the possibility is not ruled out. No aneuploids were discovered. A more recent study (Richards 1963) of mixed populations of D. fuchsii and D. purpurella in recently abandoned magnesian limestone quarries in Durham, v.c. 66, suggested that the situation there was a more complex one. These populations contained a wide range of phenotypes, many of which showed varying degrees of intermediacy between the two species, and were morphologically consistent with extensive backcrossing of the F; hybrid to both parents. Furthermore, out of nine reliable chromosome counts which were made from pollen grain mitosis, two proved to be aneu- ploid with n = 22 and 37. It also seemed that other populations occurring in long-disused magnesian limestone quarries and consisting of robust plants with broad, shallowly trilobed labella resembling T. and T. A. Stephenson’s Orchis purpurella ‘Form B’ (Stephenson & Stephenson 1922), may have arisen as a result of introgression from D. fuchsii into D. purpurella. The present work was undertaken to establish whether aneuploids of presumptive backcross origin were important in Durham populations of Dactylorhiza. MATERIALS Buds were fixed from 50 individuals in a mixed population of parent species and apparent hybrids at Quarrington, Durham, v.c. 66 (G.R. 45/330.364), on the 14th June 1974. Plants were chosen using a Im grid and a table of random numbers. The latter generated pairs of numbers to indicate | B 206 R. M. LORD AND A. J. RICHARDS grid intersections, and buds were fixed from the nearest flowering spike with only the bottom one or two flowers open. The labellum of the most open flower was preserved on card under transparent adhesive tape. METHODS Buds were fixed in 3:1 absolute ethanol/glacial acetic acid overnight and then kept in the deep- © freeze. After hydrolysis in N hydrochloric acid at 60°C for 15 minutes, they were stained in Feulgen | stain for 1 hour. Pollinia were excised and squashed in acetocarmine. Slides showing meiotic | stages were made permanent in ‘euparal’. As pointed out by Heslop-Harrison (1953), meiosis is highly synchronized; in suitable preparations large numbers of dividing cells can be observed. The following characters (Fig. 1) were scored from the preserved labella: . Maximum width . maximum length . vertical length of central lobe (if any) from apex to sinus . shape: deltoid, intermediate, trilobed . colour: white to pink, deep-pink, red-purple . markings: spots, intermediate, rings. NanAbhWwWNe FicureE 1. Labellum types in Dactylorhiza. A, D. fuchsii, trilobed labellum with spots; B, labellum of putative hybrid with intermediate shape and markings; C, D. purpurella, deltoid labellum with rings. | Measurements: 1, maximum width; 2, maximum length; 3, vertical length of central lobe. Individuals with eudiploid (2n = 40) and eutetraploid (2n = 80) chromosome counts were assumed | to be pure D. fuchsii and D. purpurella respectively. Somatic chromosome numbers were calculated from meiotic preparations by counting the || number of univalents and bivalents in a number (usually about 50) cells in each individual. Some | apparent variation (up to 3 chromosomes per cell) was noted within an individual plant. This is | thought to be due to chromosomes being obscured, or to the occasional misinterpretation of || univalents and bivalents, rather than actual variation. | RESULTS Meiotic stages (diakinesis to metaphase I) which allowed the accurate determination of somatic |) chromosome number were found in 33 individuals. Of these, 5 were eudiploid (2n = 40), 7 were | eutetraploid (2n = 80) and 13 were eutriploid (2n = 60). The remaining 8 were aneuploid, with | 2n = 44, 48, 52 and 72 (Table 1). In 13 plants, all triploids or aneuploids, a sufficient number of cells could be analysed for the! variation in the number of univalents within one plant to be counted (Table 2). Only univalents | HYBRID SWARM BETWEEN DACTYLORHIZA FUCHSII AND D. PURPURELLA 207 TABLE 1. SOMATIC CHROMOSOME NUMBERS 2an'= No. plants Ploidy level 40 5 Eudiploid 44 y 48 2 52 1 60 13 Eutriploid 72 3 80 7 Eutetraploid and bivalents were seen, so counts apparently of odd numbers of univalents are probably instances of experimental error. Eudiploids and eutetraploids invariably showed regular formation of bivalents and regular disjunction. It will be seen that a single individual may show a wide range in the number of univalents occur- ring in a meiotic cell. Among the triploids, individuals do not differ greatly as to mean number or range of univalents found, although the small range in plant number 2 and the high mean number in plant number 30 seem to differ from the rest. The number of univalents in an individual does not follow a normal distribution, but shows distinct peaks at 9 and 12, with perhaps a sub- sidiary peak at 15. 20 univalents, as constantly recorded by Heslop-Harrison (1953), rarely occurred. As might be expected, the numbers of univalents are less in subtriploid aneuploids, but here also there is a suggestion of a double peak in the distribution. The fate of the univalents is not known for certain. Limited studies of anaphase I and the second meiotic division suggest that most, if not all, migrate to one pole at anaphase I and are incorporated in one daughter nucleus. Bivalents in triploids behave normally, with regular disjunction (of 20 or more chromosomes) to each pole. Pure populations of the parent species are rare in Co. Durham, and therefore eudiploid and eutetraploid individuals from Quarrington were used to define the morphological limits of the two species. This has the advantage of removing bias arising from phenotypic differences between populations of the same species. However, introgressive hybridization at the euploid level would not be detected. It was found that the length, width and the length/width ratio of the labellum showed no signi- TABLE 2. 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K. MORTON DISCUSSION CHROMOSOME NUMBERS AND TAXONOMY Most of the chromosome numbers reported in this account confirm those already reported for the same species in other parts of their range. The only species whose chromosome numbers were previously unreported are Cirsium dissectum and C. tuberosum. Those of several varieties also appear to be new, viz. Antennaria dioica var. hyperborea, Aster tripolium var. arcticum, Cirsium arvense var. incanum, Crepis capillaris var. glandulosa, Leontodon autumnalis var. pratensis and Solidago virgaurea var. cambrica. In addition, chromosome numbers for 53 of the taxa are here reported for the first time from material from the British Isles. Most of these counts call for no comment, but the following are worthy of note: Aster tripolium var. arcticum Fries. This is a dwarf northern variant not recognized in most British Floras. It occurs in salt-marshes on several of the Hebridean islands (Heslop-Harrison et al. 1946, 1951). It grows from 5 to 15 cm high and bears only a few capitula, which are of full size. Material transplanted from North Uist to a garden in northern England grew well for several years and retained its dwarf habit. Leucanthemum vulgare Lam. The material from Kynance Cove was of a dwarf maritime ecotype about 10-15 cm high growing in the short turf near the top of the wind-swept cliffs. It retained its dwarf habit in cultivation. Senecio vulgaris L. Seed from radiate plants found on a cindery roadside near Pirbright produced plants both with and without conspicuous ray florets. All had 2n = 40. The radiate plant is referable to var. hibernicus Syme on the basis of Allen’s (1967) account of variation in this species. However, Crisp & Jones (1970) and others have suggested that var. hibernicus may represent stages of intro- gression between S. vulgaris and S. squalidus. Serratula tinctoria L. The small population growing on the steep, grassy slopes atop the magnesian limestone cliffs at Marsden consists of a distinctive maritime ecotype of shorter, more stocky habit with larger capitula in compact inflorescences. In cultivation it made an attractive rock-garden plant and retained its characters, though after several years it increased in stature from about 30 cm to nearer 40 cm. Solidago virgaurea L. This is a very variable species found in a wide range of habitats. Over a period of many years I kept plants showing much of this variation under observation in cultivation in a garden in northern England. These observations demonstrated that material of this species from the British Isles behaves in a very similar manner to Continental material studied many years ago by Turesson (1925, 1930) and more recently by Bj6rkman and co-workers (Bjorkman 1966, 1968; Holmgren 1968; Bjorkman ef a/. 1960). All this work indicates that in this species there is a wide range of genetically fixed variants, each closely adapted in physiology, phenology and morphology to particular habitats. Two major variational trends are recognizable; one of dwarf plants from open exposed habitats in the mountains, and the other of taller plants of wood- land and heaths at lower elevations. The two differ in many ways though they are not wholly discrete and various intermediates are encountered. The differences are sufficient to warrant taxo- nomic recognition and, in view of the nature of this variation, subspecific status is appropriate. British botanists have for many years referred to the dwarf variant as var. cambrica (Huds.) Sm., but it is doubtful whether our material is different from that occurring on the Continent, where Wagenitz (1964) referred to it as subsp. minuta (L.) Arcangeli. However, Turesson (1925), in a discussion of the systematics of this variation, suggested that the Scandinavian material may not be the same as S. minuta L. from the Alps, but be referable to var. alpestris (Waldst. & Kit. ex Willd.) DC., which Hegi included in subsp. minuta. Pending a resolution of these nomenclatural and taxonomic problems it is probably better that we continue to refer to our material as var. cambrica, whilst recognizing that it may not be endemic to the British Isles. Var. cambrica occurs in most of the mountainous areas of the British Isles—in Wales, the Pennines, the Lake District, mainland Scotland, the Hebrides and Ireland, where it also comes down to low altitudes on sea-cliffs. All the material that I grew retained its characters in cultivation. In cultivation var. cambrica flowers from mid June to the end of July, about 6 weeks earlier than the lowland variety—a difference which is also noticeable in the wild. Var. cambrica is characterized by its short, stocky habit, from 5 to 20 cm in height; its larger capitula, which range from 8 to 10 mm long in dried material; its compact, few-flowered inflorescences, in which the capitula are A CYTOLOGICAL STUDY OF THE COMPOSITAE OF THE BRITISH ISLES 221 borne more or less singly in a compact racemose arrangement on the main axis; its rounded, obtuse and spathulate basal leaves; and its bracts of the main axis, which are large and leaf-like. Usually the phyllaries are very narrowly lanceolate and sharply acute. Var. virgaurea, though very variable, is characteristically a lowland species of open woodland, heaths and hedgebanks. In cultivation it flowered in August and September, with material from south-western England being several weeks later than the other material. It is characterized by its taller, less sturdy stature, from 60 to over 150 cm in height; its smaller capitula, 5 to 8 mm long, arranged in slender panicles or clustered in axillary branches along the tall, slender inflorescence; its acute, oblanceolate, petiolate basal leaves; and its small, narrow bracts or reduced leaves in the upper three-quarters of the stem. The phyllaries tend to be oblanceolate and more or less blunt in many, but not all, populations. Capitulum size tends to increase from south-western to northern districts. In the valleys of northern England and Scotland, populations combining the characters of the type variety and var. cambrica are sometimes encountered. That on the upper reaches of the River Tees is particularly well developed and extensive. A stunted ecotype occurs on the grassy slopes atop the sea-cliffs at Kynance Cove. Though compact, its other characters are those of the lowland variant. It retained its characters in cultivation though it increased in height from about 20 cm to 30 cm. POLYPLOIDY The spectrum of polyploidy for the Compositae native to the British Isles is as follows: Numbers of species Primary diploids 50 Triploids 2 Tetraploids 22 Hexaploids 6 41 Octoploids : 1 Secondary base numbers 10 % polyploidy 44-6 Only species and subspecies have been used in compiling these data, except where variants of a lower level have a different ploidy level. Three native genera (Cirsium, Petasites and Tussilago) have high base numbers which are almost certainly of secondary origin due to alloploidy, and these have been regarded as polyploids. The incidence of polyploidy in the Compositae (44:6%) can be compared with that in other families for which fairly complete data from the British Isles are available—41-2°% in the Caryo- phyllaceae (Blackburn & Morton 1957) and 69-8 °% in the Labiatae (Morton 1973). There appears to be no particularly convincing reason for the lower incidence of polyploidy in the Caryophyllaceae and Compositae than in the Labiatae, though the reason may in part be associated with the pre- dominance of vigorous, vegetatively reproducing rhizomes in the Labiatae, a mode of reproduction occurring much less frequently in the other two families. ACKNOWLEDGMENTS I am very much indebted to Mr D. M. Cranston for information on previously published and unpublished chromosome counts; to the several workers who have permitted me to use their unpublished data (their names appear in Table 1); to Dr P. L. Pearson for permission to quote from his thesis; to Professor D. A. Webb, who many years ago supplied me with living material of Inula salicina; and to Miss J. M. Venn, who has assisted in preparing the data for publication. Completion of this work was facilitated by a fellowship from the Nuffield Foundation and an operating grant from the National Research Council of Canada. REFERENCES ALLEN, D. E. (1967). The taxonomy and nomenclature of the radiate variants of Senecio vulgaris L. Watsonia, 6: 280-282. Cc 222 J. K. MORTON Barser, H. N. (1941). Spontaneous hybrids between Sonchus asper and S. oleraceus. Ann. Bot., n.s., 5: 375-377. BARLING, D. M. (1955). Tragopogon pratensis in the central Cotswolds. Watsonia, 3: 210-212. BJORKMAN, O. (1966). Comparative studies of photosynthesis and respiration in ecological races. Brittonia, 18: 212-224. . BJORKMAN, O. (1968). Further studies on differentiation of photosynthetic properties in sun and shade | ecotypes of Solidago virgaurea. Physiologia Pl., 21: 84-99. BJORKMAN, O., FLORELL, C. & HOLMGREN, P. (1960). Studies of climatic ecotypes in higher plants. The temperature dependence of apparent photosynthesis in different populations of Solidago virgaurea. K. LantbrHogsk. Annir, 26: 1-10. BLACKBURN, K. B. & MortTon, J. K. (1957). The incidence of polyploidy in the Caryophyllaceae of Britain and Portugal. New Phytol., 56: 344-351. BOcHER, T. W. & LARSEN, K. (1955). Chromosome studies on some European flowering plants. Bot. Tidsskr., 52: 125-132. BoOcuer, T. W. & LARSEN, K. (1957). Cytotaxonomical studies in the Chrysanthemum leucanthemum complex. Watsonia, 4: 11-16. Burtt, B. L. (1950). Lapsana intermedia in Britain. Watsonia, 1: 234-237. CLAPHAM, A. R. (1962). Compositae, in CLAPHAM, A. R., TuTin, T. G. & WARBURG, E. F. Flora of the British Isles, 2nd ed., pp. 800-931. Cambridge. Crisp, P. & JONES, B. M. G. (1970). Senecio squalidus L., S. vulgaris L. and S. cambrensis Rosser. Watsonia, 8: 47-48. CURRAN, P. L. (1968). Chromosome numbers of some Irish plants. Jr. Nat. J., 16: 7-9. EpMonps, J. M., SELL, P. D. & WALTERS, S. M. (1974). Some British chromosome counts in the Compositae subfamily Cichorioideae. Watsonia, 10: 159-161. ‘ ELKINGTON, T. T. & MIDDLEFELL, L. C. (1972). Population variation within Centaurea nigra L. in the | Sheffield region. Watsonia, 9: 109-116. ELLioT, E. (1950). A new phase of amphiplasty in Leontodon. New Phytol., 49: 344-349. Fincyu, R. A. (1967). Natural chromosome variation in Leontodon. Heredity, 22: 359-386. Frost, S. (1958). The geographical distribution of accessory chromosomes in Centaurea scabiosa. Hereditas, — 44: 75-111. HASKELL, G. & Marks, G. E. (1952). Chromosome ecology of British Galinsoga species. New Phytol., 51: 382-387. . HESLOP-HARRISON, J. W., HESLOP-HARRISON, J., CLARK, W. A. & COOKE, R. B. (1946). Further observations | on the vascular plants of the Outer Hebrides (v.c. 110). Proc. Univ. Durham Phil. Soc., 10: 358-367. HEsLop-HarrIson, J. W. & Morton, J. K. (1951). Botanical investigations in the islands of Raasay, Rhum | (v.c. 104), Lewis and Harris (v.c. 110) in 1951. Proc. Univ. Durham Phil. Soc., 11: 12-23. HOLMGREN, P. (1968). Leaf factors affecting light-saturated photosynthesis in ecotypes of Solidago virgaurea | from exposed and shaded habitats. Physiologia Pl., 21: 676-698. HOLMGREN, P. K. & KEUKEN, W. (1974). Index Herbariorum, Part I, 6th ed. Regnum veg., 92. Kay, Q. O. N. (1969). The origin and distribution of diploid and tetraploid Tripleurospernum inodorum (L.) | Schultz Bip. Watsonia, 7: 130-141. Kay, Q. O. N. (1971a). Anthemis cotula L., in Biological flora of the British Isles. J. Ecol., 59: 623-636. Kay, Q. O. N. (1971b). Anthemis arvensis L., in Biological flora of the British Isles. J. Ecol., 59: 637-648. | Kay, Q. O. N. (1972). Variation in sea mayweed (Tripleurospermum maritimum (L.) Koch) in the British | Isles. Watsonia, 9: 81-107. MaAubpegE, P. F. (1939). The Merton Catalogue. A list of the chromosome numerals of species of British flowering plants. New Phytol., 38: 1-31. MAupgE, P. F. (1940). Chromosome numbers in some British plants. New Phytol., 39: 17-32. MILLs, J. N. & Stace, C. A. (1974). Chromosome numbers of British plants, 2. Watsonia, 10: 167-168. Morton, J. K. (1962). Cytotaxonomic studies on the West African Labiatae. J. Linn. Soc. (Bot.), 58: 231- | 283. Morton, J. K. (1973). A cytological study of the British Labiatae (excluding Mentha). Watsonia, 9: 239- 246. Morton, J. K. (1974). Chromosome numbers of British plants, 3. Watsonia, 10: 169. PEARSON, P. L. (1967). An experimental taxonomic study of Chrysanthemum leucanthemum L. Ph.D. thesis, University of Durham. | Picott, C. D. (1968). Cirsium acaulon (L.) Scop., in Biological flora of the British Isles. J. Ecol., 56: 597- | 612. | RICHARDS, A. J. (1972). The Taraxacum flora of the British Isles. Watsonia, 9 (Supplement). Rosser, E. M. (1955). A new British species of Senecio. Watsonia, 3: 228-232. Roy, B. (1937). Chromosome numbers in some species and hybrids of Centaurea. J. Genet., 35: 89-95. RUTLAND, J. P. (1941). The Merton Catalogue. Supplement No. 1. New Phytol., 40: 210-214. A CYTOLOGICAL STUDY OF THE COMPOSITAE OF THE BRITISH ISLES 223 SORENSEN, T. & CHRISTIANSEN, H. (1964). Contribution to the chromosome cytology of Petasites. Bot. Tidsskr., 59: 311-314. Stace, C. A., ed. (1975). Hybridization and the flora of the British Isles. London. STEBBINS, G. L., JENKINS, J. A. & WALTERS, M. S. (1953). Chromosomes and phylogeny in the Compositae, tribe Cichorieae. Univ. Calif. Publs Bot., 26: 401-429. STEARN, W. T. & BRIGHTON, C. (1969). The history and cytology of double daisies. J. R. Hort. Soc., 94: 403-408. TiscHLER, G. (1935). Pflanzliche Chromosomen-Zahlen pt. 2 (1). Tabul. Biol., 11: 281-304. TISCHLER, G. (1937). Pflanzliche Chromosomen-Zahlen pt. 2 (2). Tabul. Biol., 12: 57-115. TIsCHLER, G. (1950). Die Chromosomenzahlen der Gefasspflanzen Mitteleuropas. The Hague. Turesson, G. (1925). The plant species in relation to habitat and climate, 15. Ecotypes of Solidago virgaurea L. Hereditas, 6: 204-219. TurReEsson, G. (1930). The selective effect of climate upon the plant species. Hereditas, 14: 99-102, 130-133, 146-152. WAGENITZ, G. (1964). Solidago, in Illustrierte Flora von Mitteleuropa, 2nd ed., V4(3,1): 16-29. Munich. (Accepted April 1976) Watsonia, 11, 225-228 (1977). 225 Botany of the lower Weaver valley, north Cheshire T. EDMONDSON 42 Shepherds Lane, Chester The River Weaver rises near the River Gowy on the eastern slopes of the Peckforton Hills in mid-west Cheshire. While the Gowy flows north and then, without much meandering, north-north- west to its confluence with the Mersey near Ellesmere Port, the Weaver follows a tortuous course east, south-east, north, and finally north-west to enter the Manchester Ship Canal below Weston Hill. Long before that canal was constructed, however, and while the Weaver still flowed into the Mersey, it was canalized downstream of Northwich, some sharp meanders were cut off and locks were constructed at Dutton; the tidal reaches were largely discarded in favour of a new canal from Sutton to the Runcorn docks at Weston Point. This account is concerned with the five-mile stretch of the lower valley between the A56 (War- rington to Chester) and the A49 (Warrington to Whitchurch) road bridges. By absolute standards the natural history of this area might not seem outstanding but its value is enhanced in a region which some would like to see become a vast conurbation and where the density of population is already very high. Its conservation is therefore very desirable. It is clear from the recent Flora of Cheshire (Newton 1971) that the plants of the Weaver valley form an important component of the county’s flora. Earlier records are surprisingly scanty. It is evident that neither Lord de Tabley nor the collaborators in his Flora (de Tabley 1899) were very familiar with the valley, for they made no mention of Equisetum sylvaticum, Galium odoratum, Lathraea squamaria, Thalictrum flavum and Phyllitis scolopendrium. The reason is not far to seek, for, as de Tabley observed, woods and private estates were rigorously preserved for game. John Harrison, a St Helen’s miner, contributed numerous important first records for the area. Harrison who, according to de Tabley, ‘was lost sight of in 1855’ was self-taught, but, although de Tabley had reservations about his records of plants such as Hypericum hirsutum and Lathyrus sylvestris, these have since been substantiated. It is not unlikely that Harrison’s origins helped him to gain access to the Aston Estate through the goodwill of the gamekeepers. The Frodsham botanist, J. F. Robinson, contributed several interesting records from the then accessible parts of the valley. Though several of his records published elsewhere are demonstrably in error, there is no reason to doubt those accepted by de Tabley. Although published in 1899, a few years after his death, de Tabley’s manuscript was completed about 25 years earlier so that a century elapsed before a more detailed survey of the area was made. The present notes are derived from walks with Frank W. Clarke in the 1960s. These walks were recreational as well as exploratory and no attempt was made to check older records, of which we were then ignorant. Access was readily obtained and expressions of gratitude are due particularly to Mr Brian Talbot of Aston, Mr Ieuan Jones of Catten Hall and Mr S. Sproston of Belleair. The north-eastern side of the lower Weaver valley lies in the parishes of Sutton, Aston and Dutton in the old Bucklow Hundred, and the opposite side in Frodsham, Kingsley and Acton in the old Eddisbury Hundred in 10 km square 33/5.7. The uppermost solid formation is Keuper Marl, which lies against Keuper Waterstone outcrops of the Overton Fault by Frodsham Bridge. Underlying Triassic sandstones are exposed in those lateral wooded cloughs which have been more deeply incised. The region was affected by the four main glaciations and there are deposits of fluvioglacial gravel flanking the alluvial flood plain, and extensive but discontinuous deposits of sand and boulder clay on the higher land, which rises to about 50 m. The river plain varies in width from about 200 to 800 m and includes fairly dry fields, damp summer pastures and two base-rich marshes below heavily flushed slopes. There are disposal grounds for dredged sludge near Frodsham and there is now an embanked lagoon for the same purposes below Crewood. In places the canalized river banks are somewhat above the sea-level 226 T. EDMONDSON plain and some temporary flooding occurs in prolonged wet weather. A considerable amount of brine effluent enters the river from salt industries at Northwich and this salinity could account for the considerable inland extension of estuarine plants such as Apium graveolens and Scirpus mari- timus. Before the Ship Canal was built, exceptionally high tides almost lapped Frodsham Main Street, and Mrs Bates, a centenarian who died only a few years ago, could remember such an event. Woodland. Altogether there are about 30 woodland sites, including incised valleys, often heavily flushed within, wooded banks above the old river course and old plantations and parkland on more level ground around the former site of Aston Hall. The woods have been periodically thinned of the best timber and Big Wood was completely felled some years ago. As might be expected Acer pseudoplatanus predominates, with Alnus glutinosa in wetter situations, among Betula pendula, Fraxinus excelsior, Sorbus aucuparia, Quercus robur and Ulmus glabra. Aesculus hippocastanum, Pinus sylvestris and Tilia x vulgaris are evidently planted and there are a few fine specimens of Fagus sylvatica. The woodland contains well-grown Crataegus monogyna, Ilex aquifolium and Prunus spinosa. In the spring Prunus avium, which is not valued as timber, is a visual delight with Malus sylvestris and Viburnum opulus, the latter prevalent in the understorey with Corylus avellana and Sambucus nigra. A small group of Tilia cordata in a species-rich locality appears to be a relic of the ancient woodland. If the trees are not in themselves remarkable they provide a canopy for a colourful herb layer which has all but disappeared in all the developed areas near by. Some of the main members are: Athyrium filix-femina Dryopteris filix-mas Anemone nemorosa Ranunculus ficaria Cardamine amara Silene dioica Moehringia trinervia Geranium robertianum Oxalis acetosella Geum urbanum Chrysosplenium oppositifolium Epilobium montanum Circaea lutetiana Conopodium majus Mercurialis perennis Lysimachia nemorum Veronica montana Lamiastrum galeobdolon Glechoma hederacea Ajuga reptans Galium odoratum Adoxa moschatellina Lapsana communis Endymion non-scriptus Allium ursinum Arum maculatum Some species are scarcer than one might expect, e.g. Carex sylvatica, Mycelis muralis, Myosotis sylvatica, Orchis mascula, Ranunculus auricomus and Sanicula europaea. However some specialities of the area are plentiful. They include Campanula latifolia, Carex pendula, Equisetum telmateia (often extensive at sand/marl junctions and up to four feet high), Lathraea squamaria (at six discrete stations extending, discontinuously, for 100 and 200 m in two localities, variously parasitic on Ulmus glabra, Prunus avium and Corylus avellana), Milium effusum, Phyllitis scolopendrium (in all the deeper, rock-exposed gulleys and abundant on the steep, friable slopes of two ravine woods), Polystichum setiferum (characteristic of six of the deeper cloughs usually near the preceding species). Luzula pilosa and Melica uniflora are widespread but seldom in quantity on drained, untrodden fibrous-loam, and Ribes nigrum, R. rubrum and R. uva-crispa are scattered in congenial sites. Luzula sylvatica occurs on two, sand-topped woodland shoulders. Although Convallaria majalis is known to have been introduced in a pleasant dingle at Dutton, its presence also on glacial sand in another remoter place is puzzling unless somehow taken there by badgers which inhabit the bank. In two places Polygonatum multiflorum is clearly adventive from nearby woodland cottages but in so suitable an area it could once have been native. Viola reichenbachiana is a frequent woodland border plant, especially on south-facing slopes, and opens well before V. riviniana in a normal spring. The remaining woodland plants are rare in the region: Carex laevigata in one clough; Dipsacus | pilosus below a fiushed bank; and Viola odorata on a shaded bank rich in humus. Although Carex | strigosa occurs in five sites its physical requirements seem to be exacting as it is found on unstable stream-side alluvium in the deeper gulleys where the water runs rapidly after heavy rain. | A narrow, small, wooded bank with bordering scrub on exposed calcareous red marl below | BOTANY OF THE LOWER WEAVER VALLEY, NORTH CHESHIRE 227 glacial sand is important as the only Cheshire station for Lathyrus sylvestris. This is clearly part of a relict community which includes Adoxa moschatellina, Allium ursinum, Arum maculatum, Carex pendula and Melica uniflora in the shade with Ballota nigra, Clinopodium vulgare, Hypericum perforatum, Inula conyza, Linaria vulgaris, Rosa sherardii and Tragopogon pratensis in the scrubby grassland. This appears to be the only natural site in the county for Jnula and one of the few remaining sites for Clinopodium. Water Meadows. Below Bradley Orchard Carex acutiformis and Carex riparia are plentiful in the perimeter ditches, with other plants which occur also in the main alluvial marshes below the wooded marl slopes of Beckett’s Wood and Blackamoor. The marshes are important for the extensive zones of Eleocharis uniglumis and Carex disticha and contain a rich variety of plants at the various micro-levels from Veronica beccabunga in shallow runnels through Veronica catenata on summer-exposed mud to Lysimachia nummularia on flattened tussocks. Excluding common grasses and rushes, important members of the wetland association are: Caltha palustris Valeriana officinalis Ranunculus trichophyllus V. dioica Cardamine pratensis Senecio aquaticus Lychnis flos-cuculi Triglochin palustris Stellaria graminea Potamogeton pectinatus Filipendula ulmaria Tris pseudacorus Epilobium palustre Lemna gibba Apium nodiflorum Eleocharis palustris Polygonum hydropiper Carex hirta Myosotis caespitosa C. nigra Scrophularia auriculata C. spicata Mentha aquatica C. ovalis The main site for Dactylorhiza praetermissa was in Crewood Marsh, now displaced by the sludge lagoon. Brackish conditions occur in the old tidal reaches below Sutton Weir and Frodsham Lock, where Phragmites australis and Scirpus maritimus are abundant in the dykes, Carex otrubae is prolific on ditch-sides, and Hordeum secalinum is locally common in the old rank meadows. These wetlands are regionally valuable since they are the last remnants of a formerly extensive zone which included the area between the sub-coastal reaches of the Gowy and the Weaver. This is now deeply dyked and maintained for farming by pumping stations on both eastern and western boundaries. Other Habitats. The third main topographical feature of the valley are the contoured field- hummocks which are too steep or too short for ploughing. There is a complex structure of sand, clay and marl with frequent small flushed zones and spring-heads and a few leached areas. The following composite list, which excludes some very common species, embraces several plant associations of different micro-habitats: Equisetum sylvaticum Myosotis discolor Polygala vulgaris Veronica officinalis Hypericum maculatum Pedicularis sylvatica Linum catharticum Betonica officinalis Geranium dissectum Teucrium scorodonia G. molle Succisa pratensis Genista tinctoria Solidago virgaurea Lathyrus montanus Dactylorhiza fuchsii Potentilla anglica D. praetermissa Agrimonia eupatoria Scirpus setaceus Alchemilla filicaulis subsp. vestita Carex pallescens Primula veris C. caryophyllea P. vulgaris C. echinata Anagallis tenella Briza media The low embankment bordering the river is popular with cattle and holds few plants of interest 228 T. EDMONDSON except, at intervals, some species which appear to be characteristic of upper estuarine margins in this region, e.g. Tanacetum vulgare, Conium maculatum, Dipsacus fullonum and, in damper situa- tions, a little Thalictrum flavum. There is a fine stand of Carduus tenuifiorus on an eroded sandy level. Geranium pratense grows plentifully in several fenced grassy corners and the summer leaves of a female colony of Petasites hybridus—a common species in the region—grow to a prodigious height at Pickering Lock. The higher fields bordering the valley are grazed or cultivated. The few good ponds are probably mineral-rich and spring-fed and have a typical Cheshire pond association (Edmondson 1967), which includes Berula erecta, Nymphaea alba, Oenanthe fistulosa and Typha angustifolia, with Pulicaria dysenterica by the margins. A Salix-Alnus swamp in a hollow below the Kingsley slopes is notable for Carex paniculata and Scutellaria galericulata and the now isolated meanders at Dutton have a varied flora which includes Butomus umbellatus, Myosotis scorpioides, Rumex hydrolapathum and Polygonum amphibium. The verges of lanes and roads above the valley show an abundance of Pimpinella major, Leontodon hispidus and Ononis spinosa, with Lonicera periclymenum and Tamus communis prominent in the hedgerows. In this district Blackstonia perfoliata is predominantly a plant of freshly exposed or constantly flushed, bare clay (Edmondson 1974). It is massed, with Carex spicata, on the creep ledges of a large clay slip at Dutton. Clay-pits soon acquire a colourful plant population but, given a suitable topography and associated moisture levels, a greater variety is achieved when localized leaching, weathering and humus accumulation has taken place. Some old Dutton pits, although now going over to scrub, still have a fascinatingly diverse association which, in addition to Blackstonia per- foliata, includes Centaurium erythraea, Senecio erucifolius and Sieglingia decumbens. This broad account neglects the more critical groups but should mention a few interesting localized plants. Hypericum hirsutum grows on banks bordering an old pasture and Malva moschata similarly but on a lighter soil. There are vigorous thickets of the introduced Polygonum sachalinense | in scrub near Crewood, Cruciata laevipes is surprisingly scarce, while Sedum acre has effectively invaded derelict boat hulks at Sutton Locks. Finally, the restriction of Scirpus tabernaemontani, in considerable quantity, to one flushed area suggests that it is associated there with saline springs. Like John Harrison I grew up in a coal-mining district in south Lancashire where only the ‘rosebay’ temporarily brightens the spoil-heaps, and I can image his intense pleasure in visiting the lower Weaver valley. More than a hundred years later the contrast between the two districts is all the greater. Surrounded by everything that expands—industries, new towns and village dor- mitories—is it too much to hope that this valley, a microcosm of all that the local countryside used to be, will be spared the fate that befell south Lancashire in the days of the first economic revolu- tion? : REFERENCES De TABLEY, LorD (1899). The flora of Cheshire, London. EDMONDSON, T. (1967). Pond flora in North Cheshire. Field Naturalist, 12 (n.s.): 30-35. EDMONDSON, T. (1974). Habitat preferences of Blackstonia perfoliata (L.) Huds. in Cheshire and adjoining | parts of North Wales. Watsonia, 10: 78-79. Newton, A. (1971). Flora of Cheshire. Chester. (Accepted April 1976) Watsonia, 11, 229-236 (1977). 229 Artificial hybridization between Quercus robur L. and Quercus petraea (Matt.) Liebl. B. S. RUSHTON School of Biological and Environmental Studies, The New University of Ulster, Coleraine, N. Ireland ABSTRACT The results of attempted interspecific crossings between Quercus robur L. and Quercus petraea (Matt.) Liebl. are presented and compared with the results of selfed and intraspecific crosses. Only 22 fruits (acorns) were produced from 3,934 flowers pollinated in interspecific crosses. This success rate of 0:56% compares with those of intraspecific crosses of 43-6% (Q. petraea) and 51:4% (Q. robur) and selfed success rates of 40-2% (Q. petraea) and 19-3 % (Q. robur). It is argued that large seed output over the long life of an indivi- dual tree, coupled with heterosis, could allow a relatively low success rate for interspecific crosses to produce a comparatively high percentage of hybrids and backcross derivatives. The acorns from intraspecific crosses were readily distinguished on shape, colour and longitudinal striping, whilst hybrid acorns were smaller and usually intermediate in respect of these characters, but with a comparatively wide range of variation. INTRODUCTION Morphological evidence for the existence of naturally occurring hybrids in Britain between Quercus robur L., the pedunculate oak, and Q. petraea (Matt.) Liebl., the sessile oak, has been presented by Carlisle & Brown (1965), Cousens (1963, 1965), Rushton (1974) and Wigston (1971, 1975). However, as Gottlieb (1972) observed, morphological data alone should not be considered suffi- cient to establish a case of hybridity. Supplementary evidence, such as an additive profile for biochemical characters (e.g. seed proteins) that are present in each parent but not in both, the fertility of intermediates, or the occurrence of suspected hybrids in intermediate habitats, is scanty in the case of Q. robur and Q. petraea, and attempts at producing the hybrid by artificial means have met with only limited success. Indeed, despite the large-scale populational surveys in Britain and in Europe, e.g. Becker (1972), only four authors have attempted to hybridize artificially Q. robur and Q. petraea. The accounts of Klotzsch (1854) and Geschwind (1876), which have been discussed by Dengler (1941), can both be disregarded since, although they recorded a modicum of success, their methods would probably not have met the carefully controlled conditions required of modern-day tree-breeding. Pyatnitski (1939, 1947, 1954) attempted an extensive breeding pro- gramme between many Quercus species but met with only marginal success. The attempts of Dengler (1941) appear to have been the most successful, since in one year 15% of Q. robur flowers pol- linated by Q. petraea pollen produced fruits (acorns) compared with only 6% in the reciprocal cross. In other years his attempts were less rewarding, a level of between 0 and 2% being recorded. Cousens (1965) and Gardiner (1971) have emphasized the need for more extensive crossing experiments between these two species. The present paper is concerned with the results of artificial hybridization between naturally occurring individuals of the two species, Q. robur and Q. petraea, in England. SITES, MATERIALS AND METHODS CROSSING EXPERIMENTS Attempts to hybridize the species were carried out over three seasons—the first year, 1969, at Wyre Forest, v.c. 37, GR 32/745.762, and Uffmoor Wood, v.c. 37, GR 32/950.815, both near Birmingham, and during 1970 and 1971 at Hetchell Wood, v.c. 64, GR 44/438.443, and the sur- rounding small woodlands, near Leeds. 230 B. S. RUSHTON The methods employed were identical in all three years. During the previous growing season, | trees of each species were identified and labelled. A hybrid index, based on 17 characters, was calculated (Rushton 1974) for each tree and only those trees scoring the median hybrid index | representative of each species were used, i.e. extreme trees, whether they were indicative of hybridi- zation or extreme for the species, were disregarded. In April the following year, pollination bags were attached to the terminal parts of branches at approximately 2 m from the ground. Each bag | was 45 cm x 20 cm and had a transparent window to allow photosynthesis of trapped, expanding | leaves to proceed normally. Bagging was completed before the buds had fully broken—a necessary 100 ~J ur Percentage pollen viability ur Oo Q. petraea 25 0 aS 0 20 40 60 80 Time (days) 100 = = 75 ‘5 = > = 2 ro) a 50 © Q. robur D © ~ = ® oO o 25 o. 9 L 0 20 40 60 80 Time (days) Ficure 1. Loss of viability of stored pollen of Q. petraea and Q. robur. Pollen viability measured using nitroblue tetrazolium. Storage conditions: in open phials stood in closed sandwich boxes containing a layer of silica gel, the whole being kept at room temperature. ARTIFICIAL HYBRIDIZATION BETWEEN QUERCUS ROBUR AND Q. PETRAEA 231 precaution, since in two of the three years Quercus pollen was detected on pollen traps around the experimental trees within three or four days after attaching the pollination bags. The bags were checked every day and emasculation carried out as the male catkins became exserted from the buds. A preliminary study, in which pollen viability had been measured using nitroblue tetrazolium (Hauser & Morrison 1964), indicated that pollen of both species could be kept under low humidity and room temperature for a period of up to 14 days without any substantial loss of viability (Fig. 1). Since there is great variation in the time of flowering of individual trees, it was necessary to devise a scheme that would allow for this. Consequently five branches on each tree were ‘bagged’. As the trees came into flower, they were grouped in fours, two Q. robur and two Q. petraea. Large numbers of male catkins were harvested from each tree and pollen was collected from these in the laboratory by suspending them on nylon netting over petri dishes. The viability of this bulk collection of pollen was measured using nitroblue tetrazolium and any tree with an abnormally low count was rejected. Of the five bags on each tree, the female flowers of the first to mature were pollinated by pollen from the same tree, the flowers of the second by the other tree of the same species (i.e. an intraspecific cross), and the flowers of the third and fourth by the two trees of the other species in the group (i.e. interspecific crosses). The bag on the fifth branch was a control, being opened for the same time as the others but no pollination was attempted. Pollen was trans- ferred using a camel-hair brush, the controls being ‘pollinated’ with a camel-hair brush but with- out pollen. No mature acorns were produced from these control bags. Pyatnitski (1947) noted that the stigmatic surface of Quercus species has a short period of maxi- mum receptiveness (up to six days), but that the female catkin remains receptive for a period of 10-14 days. The detection of a receptive stigma is difficult, but a glistening appearance of the stigmatic surface observed during development almost certainly indicates the commencement of the period of maximum receptiveness. The female flowers were pollinated as they became receptive and this was repeated every other day until nine days after the first pollination. At the time of pollination, the number of female flowers (5 to 48) in each bag was noted. This varied from 5 to 48. In all three years, no pollination was carried out before 27th April and none after 23rd May. The pollination bags were left in place until mid-July, when all danger of contamination had passed; the bags also provided some protection during the early development of the acorns. The bags were replaced towards the middle of September, before the acorns had fully ripened, to protect them from foraging animals. The acorns were harvested, counted and recorded in mid- October. Vandalism throughout the three years of these experiments proved to be a major problem and accounted for the loss of about 15% of the pollination bags. ACORN CHARACTERS The production of a large number of acorns of known parentage provided an excellent opportunity to study acorn characters. Three characters were used to distinguish Q. robur and Q. petraea acorns —the presence of olive-green, longitudinal stripes on the outside of the mature, fresh acorn in Q. robur but absence from Q. petraea; colour, pale fawn in Q. robur but dark brown in Q. petraea; and shape, the Q. robur acorn being heavier, longer and thinner than that of Q. petraea. Longitudinal striping, colour and shape were recorded on a random sample of the acorns produced from the artificial crosses. It was found possible to score the striping as well-developed, absent or intermediate and the colour as pale fawn, dark brown or intermediate. The shape was determined as the length: breadth ratio. The colour and striping characteristics of Q. robur were scored as 0, those of Q. petraea as 2, and intermediate states as 1. RESULTS CROSSING EXPERIMENTS The results for the different types of crosses are given in Table 1. Differences are clearly evident between the success rates of different years—1970 appeared to be a relatively poor year for the production of acorns. Such annual variations between years are fairly common: Ovington & Murray (1964) noted a particularly poor year in 1961, preceded and followed by good years. Analysis of variance (Table 2) showed clearly that for every type of cross except selfed Q. petraea there was a significant difference between the results for different years. B. S. RUSHTON 232 ee 99-0 00:1 9 66S 8-0 LL:0 S £9 9-€P c:8S SC vor PIs v-79 LtC p9E c-OP v-Ov 0cI L6C €-6l 8°ST 89 v9T 3181 9) eI poure}qo pojyeurjod ssooons ssaoons sulooe SIOMO[ % UPIIA % o1njVur JO ‘ON JO "ON IL6l ve-0 rd v6S £9-0 v OP9 £ vavdjad *G x& Angos *O €1-0 I LOL 65-0 v 189 £ angod *O x & vavajad *O 6ST Ort Ips 1-¢¢ ig IZ€ SOSsO1d Dav.Jad *O SyioedseayuUy ple III PSe 8-8¢ prT civ S9SSOID Angod ‘GE dyloodsesjU] 6-97 LS “itd CLP 981 Z6E vavajad ‘OE Pryj[2S§ v-Ol (43 LOE 8-77 89 867 ANGOd “OE P2JIPS 9} RI pourey}qo pojeurjyjod 9} e1 poure}qo poyeurjod ssooons SUIOOPB SIOMOT ssooons SUIOSR SIOMOL Yes on} eu JO ‘ON * oin}eUul JO ‘ON jo ‘ON jo ‘ON OL6I 6961 a a een VIVYLId ‘O UNV YAGON ‘O AO SASSOUD OIOAMSUALNI GNV -VULNI SO GNV “ONIATAS AO SLTNSAU “I ATAVL i 7 px 4 ARTIFICIAL HYBRIDIZATION BETWEEN QUERCUS ROBUR AND Q. PETRAEA 233 TABLE 2. ANALYSIS OF VARIANCE OF THE RESULTS OF SELFING, AND OF INTRA- AND INTERSPECIFIC CROSSES OF Q. PETRAEA AND Q. ROBUR: DIFFERENCES BETWEEN YEARS Degrees of Variance freedom Variance ratio r Selfed Q. robur Between years 2 1,634-6 9-89 < 0-001 Within years 64 165-2 Selfed Q. petraea Between years 2 2,358-3 2:57 0-05-0-10 N.S. Within years 58 918-1 Intraspecific Q. robur crosses — Between years 2 5,868-9 18-83 < 0-001 Within years 59 311-7 Intraspecific Q. petraea crosses Between years 2 7,967°-3 24-24 < 0-001 Within years 65 328-7 : Q. petraea 2x Q. robur 3 Between years 2 5-08 13-37 < 0-001 Within years 146 0-38 Q. robur 2x Q. petraea 3 Between years 2 5-71 10-20 < 0-001 Within years 151 0-56 N.S. Not significant. There are obvious differences between the different types of cross (Table 3). The results suggest that, although not self-sterile, the success rate of selfed Q. robur trees is significantly lower than that of intraspecific Q. robur crosses for all three years (Table 3). Generally, in such selfed Q. robur crosses, the success rate was about 35% of the corresponding intraspecific cross (Table 1). The same was not true for selfed Q. petraea and intraspecific QO. petraea crosses, although, in two years out of the three, the selfed crosses produced a lower success rate and in one year, 1971, this dif- ference became significant (Table 3). Generally, only rather low levels of success were obtained in the intraspecific crosses; with the exception of the poor year 1970, only about 50% of the flowers pollinated actually produced acorns. The probable reasons for this lie either with the pollination method or with some barrier between successful pollination and the production of the mature acorn. Although the detection of a ‘receptive’ stigma was difficult, it is felt that a more likely explanation lies in the abortion of maturing seed. Under natural conditions, rarely more than three acorns are ever produced on any one peduncle—this is particularly true of Q. petraea, where the short peduncle brings the acorns into close proximity with the stem and petioles, so that there is little space available for all the acorns to mature. In Q. robur the same is true, since the female flowers are clustered at the end of the peduncle. During pollination, many more than three flowers were commonly found on each peduncle, and it seems likely that factors have contributed to bring about the abortion of some TABLE 3. ANALYSIS OF VARIANCE OF THE RESULTS OF SELFING, AND OF INTRA- AND INTERSPECIFIC CROSSES OF Q. PETRAEA AND Q. ROBUR: DIFFERENCES BETWEEN TREATMENTS yi Analysis of variance between: 1969 1970 1971 Q. petraea selfed and Q. robur selfed <0-001 0:10-0:25 N.S. <0-001 Q. petraea selfed and Q. petraea intraspecific crosses 0-05-0-10 N.S. 0:10-0:25 N.S. <0-001 Q. robur selfed and Q. robur intraspecific . crosses <0-001 <0-001 <0-001 Q. petraea intraspecific crosses and Q. robur intraspecific crosses 0-50-0-75 N.S. 0-10-0-25 N.S. 0:50-0:75 N.S. Q. petraea as 2 parent and Q. robur as 2 parent in interspecific crosses 0:25-0:50 N.S. <0-001 0-10-0-25 NS. N.S. Not significant. 234 B. S. RUSHTON maturing seeds. Frequently, it is possible to see immature acorns on peduncles at the end of the summer alongside fully mature acorns. : Crosses between the two species were singularly unsuccessful, the level of success being approxi- mately 1% of that of intraspecific crosses (Table 1). Over the three years, 22 hybrid acorns were produced out of a total of 3,934 flowers pollinated, a success rate of 0:56%. Q. robur proved to be the more successful female parent in all three years, and was significantly better than Q. petraea in 1970 but not in 1969 and 1971 (Table 3). Such low levels of success are similar to those obtained by Dengler (1941) and Pyatnitski (1939, 1954). Attempts were made to germinate some of the 22 hybrid acorns. Two acorns from 1969 were dissected in order to determine whether or not the embryo was fully formed. These acorns were lighter and smaller both in length and breadth than those derived from intraspecific crosses, but as far as could be determined, the embryo was fully developed and appeared normal. The remaining 20 acorns were sown in John Innes Compost No. 2, one per 8 cm diameter black polythene pot. No mature seedlings have been produced, but some degree of success was obtained. Of these 20 acorns, five were infected by the fungus Sclerotinia pseudotuberosa (Rehm) Boud., which converts the whole of the acorn into a black sclerotium; three failed to show any signs whatsoever of germination; six produced a radicle but no further development took place; two produced an extensive root system, but no plumule emerged; two produced an extensive root system, but the seedlings died shortly after the plumule emerged; two produced weak seedlings, with fully emerged plumules, but these too died, one after 8 and the other after 14 weeks. The percentage of hybrid acorns infected by the fungus S. pseudotuberosa (nearly 25%) is very much higher than that of non-hybrid acorns. Acorns used for other experiments and collected from the same woodlands were also infected, but to a lesser degree (4%), suggesting that hybrid acorns may be more susceptible to attack by the fungus. ACORN CHARACTERS The results for acorn characters are presented in Table 4. Acorn shape, colour and striping dif- ferentiated the species very clearly but there were slight differences between selfed and intraspecific crosses within each species. This contrasts with population samples observed by Wigston (1971). He noted the colour and striping of naturally produced acorns in six woodlands. In both the Q. petraea and the Q. robur populations, he found that generally only 50% of the acorns were of the charac- teristic type; approximately 10° of the acorns resembled those of the other species, and the remaining 40% were intermediate. Jones (1959) regarded both colour and striping as useful charac- ters in distinguishing the species and the results presented here agree closely with such an assertion. However, it must be remembered that the present results are from artificially produced acorns whilst Wigston’s results were based on population samples. The hybrid acorns were generally intermediate for colour and striping, but showed a wide range of variation (Table 4). The differences in acorn shape are consistent with published accounts and, since they are from controlled crossings, could probably be regarded as ‘typical’ of the species. However, Wigston’s (1971) study showed that acorn shape had an anomalous distribution. In the Q. petraea woodland studied by him, the length: breadth ratio ranged from 1-0 to 1-4, whilst in the three Q. robur wood- lands studied the ranges were 1-3—1-7, 1-0-1-:3 and 1-0—1-4. Thus two of the Q. robur woodlands produced similarly shaped acorns to those of the Q. petraea woodland. Wigston’s results were taken from natural populations and, moreover, since shape is a function of differential growth, TABLE 4. MEANS AND STANDARD DEVIATIONS OF ACORN CHARACTERS OF ACORNS OF KNOWN ORIGIN. COLOUR AND STRIPING SCORED AS INDICES AS DESCRIBED IN TEXT Length: breadth Sample ratio (shape) Colour Striping size Selfed Q. petraea 1-49+0-19 1-:94+0-17 1-94+0-08 76 Intraspecific Q. petraea crosses 1-36+0-26 1-96+0-13 1-86+0-10 144 Selfed Q. robur 1-7140-12 0:44+ 0-24 0-39+0-43 67 Intraspecific Q. robur crosses 1-68+0-14 0-36+0-19 0-28+ 0-26 200 Interspecific crosses 1-57+0-25 0-95 + 0-43 0:744+0-71 22 ARTIFICIAL HYBRIDIZATION BETWEEN QUERCUS ROBUR AND Q. PETRAEA = 235 their appearance is likely to be markedly influenced by external factors. Jones (1959) indicated that acorns produced in southern England are much larger than those produced in Scotland and Ireland, due possibly to the longer, warmer summers in the former. Geographical differences might, therefore, account for the differences between Wigston’s (1971) results from populations in south-western England and those reported here. The shape of the hybrid acorns fell midway between those of Q. robur and Q. petraea, but again it was very variable and overlapped considerably the ranges shown by the parent species. It should also be noted that the acorns were generally smaller than in either species. DISCUSSION The low success rates (0-5-1 %) for artificially produced hybrids noted here and by other authors prompt the question whether or not they can create and maintain the high level of hybridity reported by Cousens (1962, 1963, 1965) and Wigston (1971). Jones (1959), for example, considered that the large numbers of intermediates recorded in natural populations were not of hybrid origin, but were ‘mostly based on imperfect understanding of the specific characters’. Cousens (1963), in discussing the results of Dengler (1941) and Pyatnitski (1939, 1947), argued that since Quercus species show so much intraspecific morphological variation, any crossing plan should attempt to include all morphological types—those at the very extremes as well as those showing characters of the other species. Although an admirable suggestion, lack of manpower and resources prevented such an inevitably large crossing pian to be undertaken in the present investigation. Cousens (1963) also noted the importance of attempting backcrosses. This is an important point, since, once established in a population, an F; hybrid may produce a whole range of backcross progeny and, consequently, once the barriers to production of an F; hybrid are overcome, the rate of introgres- sion depends mainly on the success of backcrosses. No information on this point is forthcoming from the researches presented here, since the exercise was solely to determine the success of inter- specific crosses using trees of known morphological purity. However, it has already been shown {Rushton 1974) that trees of suspected second and third generation backcross status had a high level of pollen viability when compared with suspected F;s and consequently the production of backcrosses might be relatively successful compared to the initial production of F; trees. One important point rarely stressed in hybridization studies within the genus Quercus is that, whilst large numbers of hybrid trees have been recorded, only a relatively small proportion of these could be regarded as Fs, the others being probably backcross derivatives. Therefore, the question should be revised—can such a low success rate as recorded for artificially produced hybrids account for the frequency of apparent F,; hybrids observed in nature? It has to be remembered that individual trees of Quercus species are generally long-lived and consequently, as Stebbins (1950) has pointed out, although the success rate of production of hybrids may be fairly low, the long period over which they may be produced means that, in absolute terms, the number of hybrid acorns may be quite high. For example, Jones (1959) recorded that 120—140-year-old trees produced on average 50,000 acorns per year, and if only 0:1% of these were of hybrid origin, this would represent 50 acorns per tree per year for a substantial period of time. Under natural conditions, an oak woodland must receive large quantities of foreign pollen (Stebbins et al. 1947) and consequently the possibility of hybrid production must always exist. Klotzsch (1854) reported heterosis in the F; hybrids, whilst in the present investigation it proved impossible to raise mature seedlings. However, Cousens (1965) argued that, since Q. robur is a very variable species, the degree of interfertility with Q. petraea might also be variable and might be dependent upon the parental genotypes. Gardiner (1970) has noted a comparable situation in Betula pendula Roth and B. pubescens Ehrh., where there is normally only a low interspecific crossing rate, but where certain trees of the two species will hybridize easily. Similarly, the degree of heterosis could also depend upon the parental genotypes and could account for the differences between previous work and the present study. If the hybrid were more vigorous than either parent, this might overcome some of the objections proposed by Jones (1959) to the existence of wide- spread hybridization among British oak species. Obviously, such negative results as presented here cannot be used to support the hypothesis of widespread hybridization between Q. robur and Q. petraea. However, the successful production of 236 B. S. RUSHTON hybrid acorns, even though they were apparently inviable, together with the fact that only a limited range of material of each species was used, indicate that a wider survey, both in terms of the num- bers of crosses attempted and the morphological types of the trees used, might produce comparatively large quantities of hybrid acorns. ACKNOWLEDGMENTS Mr A. S. Gardiner gave much advice in the early stages of this work and I am indebted to him for his help and interest. My thanks are also due to Dr M. C. Lewis for encouragement and stimulation during the progress of this work. REFERENCES BECKER, M. (1972). Quelques observations morphologiques chez le Chéne sessile et chez le Chéne pédoncle. Bull. Soc. bot. Fr., 119: 231-236. CARLISLE, A. & Brown, A. M. F. (1965). The assessment of the taxonomic status of mixed oak (Quercus spp.) populations. Watsonia, 6: 120-127. Cousens, J. E. (1962). Notes on the status of sessile and pedunculate oaks in Scotland and their identification. Scott. For., 16: 170-179. Cousens, J. E. (1963). Variation of some diagnostic characters of the sessile and pedunculate oaks and their hybrids in Scotland. Watsonia, 5: 273-286. Cousens, J. E. (1965). The status of the pedunculate and sessile oaks in Britain. Watsonia, 6: 161-176. DENGLER, A. (1941). Bericht tiber Kreuzungsversuche zwischen Trauben- und Stieleiche und zwischen europdischer und japanischer Larche. Mitt. H.-Géring-Akad. dt. Forstwiss., 1: 87-109. GARDINER, A. S. (1970). Pedunculate and sessile oak (Quercus robur L. and Quercus petraea (Mattuschka) ~ Liebl.). A review of the hybrid controversy. Forestry, 43: 151-160. GorTT ies, L. D. (1972). Levels of confidence in the analysis of hybridization in plants. Ann. Mo. bot. Gdn, 59: 435-446. Hauser, E. J. P. & Morrison, J. H. (1964). The cytochemical reduction of nitroblue tetrazolium as an index of pollen viability. Am. J. Bot., 51: 748-752. JONES, E. W. (1959). Quercus L., in Biological flora of the British Isles. J. Ecol., 47: 169-222. OVINGTON, J. D. & Murray, G. (1964). Determination of acorn fall. Q. J] For., 58: 152-159. PYATNITSKI, S. S. (1939). Gibridizacsiya dubov. Lesn. Hoz., 7: 38-43. PYATNITSKI, S. S. (1947). Ob opylenii u dubov i prorastinii pyltsy na ryl’tsah. Dokl. Akad. Nauk SSSR, 56: PYATNITSKI, S. S. (1954). In ALBENSKI, A. V., Methody Ulucsenija drevesnyh pored, p. 147. Moscow & | Leningrad. RusuTon, B. S. (1974). The origin and possible significance of variation of leaf structure in the two native | British oak species, Quercus robur L. and Quercus petraea (Matt.) Lieb/. D.Phil. thesis, University of | York. STEBBINS, G. L. (1950). Variation and evolution in plants. New York. STEBBINS, G. L., MATZKE, E. B. & EPLING, C. (1947). Hybridization in a population of Quercus marilandica | and Quercus ilicifolia. Evolution, Lancaster, Pa., 1: 79-88. WicsTon, D. L. (1971). The taxonomy, ecology and distribution of sessile and pedunculate oak woodland in | south-west England. Ph.D. thesis, University of Exeter. WIGSTON, D. L. (1975). The distribution of Quercus robur L., OQ. petraea (Matt.) Liebl. and their hybrids | in south-western England, 1. The assessment of the taxonomic status of populations from leaf charac- | ters. Watsonia, 10: 345-369. (Accepted May 1976) Watsonia, 11, 237-246 (1977). 237 A review of Rubus section Discolores P. J. Muell. in Britain A. NEWTON 11 Kensington Gardens, Hale, Cheshire ABSTRACT Amendments to the British list of Rubus section Discolores P. J. Muell. are proposed and discussed. Three new species: Rubus armipotens, R. rossensis and R. anglocandicans are described and a revised list of the Section is given. Distribution maps of the new species are included. INTRODUCTION The taxonomy of Rubus section Discolores in Britain is greatly complicated by hybrids of the diploid, outbreeding Rubus ulmifolius Schott. These may be sterile or fertile, are usually robust and showy, and occur frequently as isolated clumps or individuals in areas where R. ulmifolius comes into contact with ancient bramble populations, especially in southern and western England and coastal South Wales. Such plants are frequently collected and named from Watson (1958), but, since very few British examples of the Section have been found to match syntypes of the original authors, it is likely that most of these identifications are erroneous. It appears necessary, therefore, to establish a reliable British list which may serve as a sounder basis for future observa- tion and research. The twenty names discussed below are, with one exception, those included in this section by Dandy (1958); they were extracted from Watson’s manuscripts which were post- humously published (Watson 1958). 5 DISCUSSION 1. Rubus ulmifolius Schott Watson (1958) gave a satisfactory account of this species although the list of fertile hybrids could be amended and expanded not only for cases where the parentage is obvious but also where it is putative or uncertain. As it is a thermophilous species it avoids high ground in Wales, Scotland and northern England and, north of a Mersey-Trent line, is largely confined to the coasts as far as Ayr in the west and Montrose in the east. 2. R. pseudobifrons Sudre The widespread and well-marked British species given this name by Watson (1958), and earlier (1928) referred to as R. winteri P. J. Muell., differs from the holotype of R. pseudobifrons Sudre (coll. G. Genevier, 28/6/1867, Mortagne, Vendee, France, as R. discolor, herb. P. J. Muell., no 1318 in LAU) in several respects, but notably in the shape and toothing of the terminal leaflet and in stem and panicle armature; it is therefore described below as a new species. Rubus armipotens Barton ex A. Newton, sp. nov. Robustus. Turio altiarcuatus in umbrosis fuscoviridis in apricis rufoviolaceus, pruinosus, capillis appressis dense obsitus, obtusangulus superficiebus planis vel concavis, striatis; aculeis crebris ad angulos dispositis longis, validis, plerumque patentibus, acute acuminatis e basi lata paullatim contractis instructus. Folia 3- vel 5-nata atroviridia, margine undulata digitata vel subpedata quorum petioli aculeis falcatis nonnullis validis armata; foliola subimbricata superne + glabra inferne grisea vel albe- D 238 A. NEWTON scentia tomentosa, pubescentia; foliolum terminale obovatum vel rhomboideum late acuminatum basi emarginata dentibus divaricatis mucronatis haud profundis. Inflorescentia angusta ad apicem aphylla, ramuli superiores anguste, inferiores laxe adscendentes cymosi c 3-flori saepe inter se distantes; pedicelli breves. Rachis vix flexuosa supra tomentosa praeterea pilosa, aculeis nonnullis rectis declinatis vel patentibus longis tenuibus obsita. Sepala griseoalba tomentosa pubescentia interdum parum aculeolata, laxe reflexa. Petala sub- orbicularia pallide rosea margine capillata apice emarginata. Stamina alba stylos pallide luteos vix superantia. Carpella breviter pilosa. Robust. Stem high-arching, greenish in shade to brownish or reddish-purple in sun, closely felted with appressed tufted and simple hairs, pruinose, blunt-angled with sides flat to slightly concave, striate; prickles on the angles fairly numerous, long, strong, quickly tapered to a slender sharp point from broad bases, usually patent, a few sometimes obfalcate, straight, slanting or with a falcate tip on the same plant. Leaves 3- to 5-nate, dark green, with undulate margins, digitate or subpedate, petioles with a few rather large falcate prickles; leaflets usually subimbricate, + glabrous above, grey to white felted (suppressed in shade) and pubescent beneath. Terminal leaflet constant in shape, + narrowly obovate or rhomboid with wavy margins, broadly acuminate with usually shallow, open, divaricate, + mucronate teeth, and emarginate base. Panicle usually leafless at apex, composed of + narrowly ascending cymose usually 3-flowered branches often spaced out giving a narrow. + equal appearance; pedicels short; lower branches laxly ascending from axils of ternate leaves; simple leaves rarely more than one; leaves white-felted and pilose beneath. Rachis + straight or slightly flexuous, felted and moderately pilose above with spreading hairs, armed with several long, straight, declining, slender prickles, sometimes deflexed at base. Sepals greenish-white, felted, pubescent, smooth, sometimes minutely aculeolate, loosely reflexed. Petals suborbicular, very shortly clawed, pale pink, erose, ciliate on margins. White stamens + equal to yellowish styles. Anthers glabrous; carpels shortly pilose. HOLOTYPUS: Sibford Heath, v.c. 23, 17/8/1927, H. J. Riddelsdeil, as R. godroni, nos 1883/5 herb. Barton & Riddelsdell (BM) Numerous sheets of this species are scattered throughout the main British herbaria under a variety of names; early gatherings of open-ground specimens were attributed to one of the Con- tinental ‘discolorous’ taxa current at the time, e.g. R. discolor, R. thyrsoideus, R. robustus, and shade-grown examples were called R. rhamnifolius or R. villicaulis. Identification with Continental taxa has always presented problems. Rogers (1900) included it under R. argentatus P. J. Muell. and later (1905), at Focke’s instigation, adopted the name R. godronii Lec. & Lam. But in his MS notes in 1908 (BM) he admits ‘our godroni is very aggregate and our robustus ill defined’. Many gatherings made by H. J. Riddelsdell in the 1920s now in BM were labelled R. godronii but later W. C. Barton, after working through Continental material in Focke’s and Miiller’s herbaria — without finding a match, labelled many of the sheets with the MS name R. armipotens, but left no | description. This name is now taken up as a tribute to his extensive research into this critical group of brambles. The first British reference to R. pseudobifrons was by Watson (1929b), but he nowhere gave reasons for his choice. It is concluded that, as in many other cases, he was relying solely on the description in Sudre (1909) and misinterpreted it. The known distribution of this species is shown in Fig. 1 and includes v.c. 7, 8, 11-19, 21-23, 27, 33, 35, 37-40, 42, 55, 62. 3. R. chloocladus W. C. R. Watson (R. pubescens Weihe ex Boenn., non Rafin.) Watson’s specimens from Clophill, v.c. 30 (K), the only recorded occurrence of this plant in Britain, while superficially similar to R. chloocladus differ substantially in details, such as stouter stem prickles, stem pilosity and shape of terminal leaflet, from herbarium specimens from Germany and collections I have made in Weihe’s own district with H. E. Weber. R. chloocladus should there- fore be excluded from the British list. The identity of Watson’s specimens is not known. A REVIEW OF RUBUS SECTION DISCOLORES P. J. MUELL. IN BRITAIN 239 ign o WwW cate on af az NO PLOTTED ON UTM GRID © Ficure 1. Distribution of Rubus armipotens Barton ex A. Newton in the British Isles. 4. R. winteri P. J. Muell. ex Focke Syntypes of this taxon (incl. Wirtgen, Pl. Sel. Fl. Rhen. 1063, K) reveal a plant virtually identical with Set of British Rubi no 113 from Stydd, v.c. 57, issued as R. argentatus var. robustus, and some specimens collected by Bagnall and Bloxam and named R. discolor y macroacanthos from Hartshill, v.c. 38. The same species occurs also in v.c. 36, 55 and 56, but many of Watson’s determinations of specimens from other counties (OXF, SLBI) are incorrect and his description in Watson (1958) was widened to include some of these. 240 A. NEWTON 5. R. crassifolius Genev. Specimens of a plant frequent in parts of v.c. 34 and 36 were referred by Rogers (1900) to R. argentatus P. J. Muell. and later (1905) to R. godronii Lec. & Lam., which has priority. Sudre (1904) determined specimens as R. propinquus P. J. Muell. and was followed by Watson (1958). Since this was a later homonym of R. propinquus Richardson, Dandy (1958) replaced it in the British list by R. crassifolius Genev., which is given by Sudre (1909) as a synonym. R. crassifolius, however, as revealed by syntypes in CGE (coll. G. Genevier, Evrunes, 4 la Rigandiére, Vendée, France, 3/9/1866), is a densely hairy plant in all its parts and differs also in other respects from the British taxon. Since our plant is also different from R. propinquus P. J. Muell., as revealed by no. 1316 in LAU (syntype), and also from R. godronii Lec. & Lam. (Rub. praesertim Gall. no. 69 ex Nancy = R. argentatus P. J. Muell. (Bat. Eur. 332 in BM, MANCH)) and cannot otherwise be matched, it requires to be described as a new taxon. Rubus rossensis A. Newton, sp. nov. Turio in umbrosis viridis in apricis fuscus vel atrorufescens, pruinosus, capillis appressis dense obsitus, obtusangulus superficiebus planis striatis. Aculeis paucis falcatis vel curvatis vel rectis declinatis vel patentibus, nonnullis apice obfalcatis e basi valida exortis praeditus. Folia 3- vel 5-nata subpedata atroviridia quorum petioli pilosiores quam turio aculeis paucis curvatis vel falcatis obsiti. Foliola + imbricata; foliolum terminale ovale obovatum vel sub- orbiculare paullatim nonnunquam longe cuspidatum basi truncata dentibus crenatis haud pro- fundis superne + glaber subtus aspere pilosa viride- vel griseoalba tomentosa. Inflorescentia ad apicem aphylla pyramidata superne bracteis longis trifidis inferne foliis non- | nullis simplicibus vel 3-natis ornata ramuli inferiores multiflori saepe longi foliosi medii laxe ~ adscendentes subcymosi 3-5 flori superiores pedicellis brevissimis aculeolis curvatis ornatis congesti. Rachis vix flexuosa inferne tomentosa, pilosa superne villosa aculeis rectis (nonnullis curvatis) e basi media exortis instructa. Sepala griseotomentosa albomarginata crasse pilosa inermia reflexa. Petala suborbicularia rosea margine pilosa. Stamina rosea stylos pallide roseos superantia. Carpella glabra vel capillis sparsis obsita. Stem green (in shade) or brownish to maroon, closely felted with appressed simple and tufted hairs, also with a few long hairs, pruinose, blunt-angled with flat striate sides. Prickles few, on the angles, varying from falcate, slightly curved, straight sloping to exactly patent or obfalcate at the tip (more or less constant on a particular plant), from a stout but not usually long base. Leaves 3- to 5-nate subpedate, glossy dark green, petioles more pilose than stem with a few strong-based curved or falcate prickles. Leaflets + imbricate, the terminal oval, roundish-obovate or orbicular, long or short cuspidate, with entire or emarginate base, with shallow crenate teeth + glabrous above, roughly pilose and greenish or greyish to white-felted beneath (suppressed in shade). Panicle narrow pyramidal, + flat topped, lower branches long, spreading, many-flowered, often leafy, median branches loosely ascending, subcymose 3—5 flowered, uppermost with very short pedicels giving a congested appearance. Pedicels short with a few small curved prickles. Uppermost | quarter of panicle leafless but with long trifid bracts almost equalling branches, with several simple | and ternate leaves below. Rachis somewhat flexuous, felted, thickly pilose below, villose above with several mostly straight prickles (a few curved) tapering gradually from a moderate base. Panicle with a few minute glan- dular hai1s up to 0-5 mm, particularly on the bracts and stipules. Sepals grey-felted, with white margins, + strongly pilose, unarmed, reflexed. Petals suborbicular, short-clawed, bright pink with strongly pilose margins. Stamens pink, longer than pinkish styles. Carpels glabrous or with a few longish hairs. HOLOTYPUS: Hillside above Redbrook, v.c. 34, 11/8/1892, A. Ley & W. M. Rogers, as R. argentatus P. J. Muell., Set of British Rubi no. 30 (MANCH) | This bramble is particularly prominent in the vicinity of Ross and southward to the Forest of A REVIEW OF RUBUS SECTION DISCOLORES P. J. MUELL. IN BRITAIN 241 FIGuRE 2. Distribution of Rubus rossensis A. Newton in the British Isles. Dean, where it makes a fine show on wood margins with handsome rose-pink flowers and grey- white foliage. Some of the early gatherings received various names but, after Rogers’ visit to the area in 1892, it became the principal component of his concept of R. argentatus and was later known as ‘Herefordshire godronii’. A good series can be seen in herb. Barton & Riddelsdell (BM) nos. 648, 915, 4880/2-5S, 4892/4, 8147/8 (isotypi). The known distribution is shown in Fig. 2 and covers v.c. 33-37, 41, 45 (T. A. W. Davis no. 73/1359). 6. R. lamburnensis Rilstone This Cornish plant is constant over a limited area and should be retained in the list as a species. 242 A. NEWTON 7. R. carnkiefensis Rilstone Another constant but local Cornish plant whose affinities are rather, as the author states, with R. dumnoniensis Bab. and R. ramosus Bloxam ex Briggs than with the section Discolores. It should be included within the section Sylvatici, subsection Discoloroides. 8. R. pydarensis Rilstone Rilstone (1952) finally placed this in the Discolores where, in view of its constancy over a wide area and its combination of characters, it should be retained as a species. 9. R. stenopetalus Muell. & Lefév. The single British gathering of this plant by Watson from Everleigh Ashes, v.c. 8 (SLBI), has some affinity with R. armipotens and is distinct from R. stenopetalus Muell. & Lefév., as revealed by Bat. Eur. 342 (BM) (coll. Questier 7/1860, Buisson de Tillet nr Vaumoise, Valois, France, Joc. class.). Its inclusion in the British list cannot be sustained. 10. R. geniculatus Kalt. The single British specimen cited by Watson (1958) cannot be equated with gatherings of R. geniculatus Kalt. from the Aachen district, nor with any other named taxon known to me. 11. R. cuspidifer Muell. & Lefeév. Specimens so determined by Watson (K, SLBD cannot be matched with any named taxon known to me. They differ from one another and also from the holotype of R. cuspidifer Muell. & Lefeév. (coll. Lefévre 26/7/1855, Forét de Retz, Oise, France, herb. P. J. Miiller no. 1387 in LAU). The inclusion of this species in the British list cannot, on this evidence, be justified. 12. R. cornubiensis (Rogers & Riddelsd.) Rilstone As Rilstone (1950) suggested in his account, the association of this species by Watson with R. bifrons Vest ex Tratt., a widespread central European species which I have studied both in exsiccata and in the German Ardennes, cannot be maintained. The British species belongs rather in the section Sylvatici, subsection Discoloroides. 13. R. tresidderi Rilstone This local Cornish species should also, as its author suggests, be included within the section Sylvatici, subsection Discoloroides. 14. R. vulnificus Lefév. ex Genev. Lefévre’s original gatherings, as revealed by specimens in herb. P. J. Miiller (LAU), differ signi- ficantly from Watson’s gatherings from v.c. 14 and determined as R. vulnificus. These represent a local unnamed taxon. R. vulnificus Lefév. ex Genev. should be excluded from the British list. 15. R. procerus P. J. Muell. This is widely planted as ‘Himalayan Giant’ in gardens and allotments and frequently persists when these are abandoned. It is also bird-sown and, being extremely vigorous, it is soon established in hedges and on waste ground. It is now widely naturalized and spreading in Britain, particularly on the outskirts of towns; any gathering of a bramble of section Discolores encountered in such habitats should first be checked for this. A REVIEW OF RUBUS SECTION DISCOLORES P. J. MUELL. IN BRITAIN 243 16. R.x polioclados W. C. R. Wats. This taxon was said by Watson (1929a) to be abundant in parts of Epping Forest, v.c. 18, and to have an affinity with R. ulmifolius. It exhibits considerable glandular development in stem and panicle and is best regarded as a hybrid between R. ulmifolius and a glandular species. It was not included in Watson (1958). 17. R. neomalacus Sudre Sudre’s autographed syntypes (coll. G. Genevier, 1858 & 1860, Torfou & St Hilaire, Vendée, France, herb. P. J. Miiller, nos 1636/8-40, LAU) are indeed close to, if not identical with, some specimens so determined by Watson. The name should be retained provisionally for some Surrey and Hampshire gatherings, but care should be taken to distinguish it from R. rubritinctus and from R. imbricatus, and also from hybrid derivatives of R. armipotens which occur in the same area. I have collected examples from Bramshott Common, v.c. 12, and the following, from v.c. 17, can also be taken as representative: BM (herb. Barton & Riddelsdell) 6541 Wareham Hill, Witley W. M. Rogers 4.8.1894 6542 Haslemere W. M. Rogers 20.8.1900 6543 Ockham Common C. E. Britton 8.1901 6544 Farley Heath, Guildford W. C. R. Watson 15.7.1930 CGE Opposite Littleworth Common, NW side of A3 W. H. Mills 1.7.1953 Milford Heath W.C. R. Watson 30.6.1934 Smithwood Common W.C. R. Watson 14.7.1930 OXF Witley (no. 2050) E. S. Marshall 6.7.1897 SLBI Thursley Common E. S. Marshall 5.9.1891 Fairmile Common C. E. Britton 9.8.1902 near Newlands Corner C. Avery 7.7.1938 18. R. falcatus Kalt. Despite extensive enquiries to various European herbaria, no authentic specimen of Kaltenbach’s can be discovered. The specimen referred to by Watson (1958), collected by himself and N. D. Simpson at Neuenhof, near Forst, Aachen, no. 371137 in herb. Simpson (BM), appears to be R. goniophylloides Sudre, which I have seen myself in quantity in the Aachen district. Despite Watson’s assertion of the identity of this with the ‘Sandy (Beds) plant’ there are significant dis- parities and, since I am unable to match the British species with exsiccata of any eligible taxon validly named on the Continent, nor could it be identified by H. E. Weber (pers. comm. 1974), it is here described as a new species. R. anglocandicans A. Newton, sp. nov. Turio altiarcuatus fuscorufescens + fulgens subglaucescens + glaber acutangulus superficiebus sulcatis. Aculeis ad angulos dispositis e basi lata paullatim contractis rectis declinatis vel patentibus vel curvatis vel falcatis nonnullis uncinatis praeditus. Folia quinata subdigitata superne glaber inferne griseo- vel albo-tomentosa molliter pubescentia. Foliolum terminale late vel anguste obovatum vel ellipticum basi truncata longe acuminatum argute inciso-serratum. Inflorescentia saepe elongata thyrsoidea ad apicem aphylla ramuli pedunculis brevibus cymosi, pedicellis sat longis, utrisque tenuibus aculeolis curvatis instructis. Rachis vix flexuosa inferne glabrescens superne tomentosa pilosa aculeis validis uncinatis falcatisve e basi lata exortis obsitus. Petala alba late elliptica ciliata; sepala viride- vel griseoalba tomentosa pubescentia reflexa. Stamina alba stylos virides parum superantia. Carpella capillis longis barbata. 244 A. NEWTON Stem brownish-red, high-arching, + shining, subglaucescent, + glabrous, usually sharp-angled with grooved sides. Prickles few on the angles, gradually tapered from a broad base (quite wide — for most of length), straight, slanting, curved, or falcate. | Leaves quinate, subdigitate, glabrous above, velvety pubescent and thickly grey to white-felted | beneath. Terminal leaflet narrow to broad, obovate or elliptical, acuminate, with an entire base, : coarsely and sharply serrate. Panicle often long, cylindrical, the branches mostly short-peduncled cymes, the pedicels rather long. Apex usually leafless but with broad bracts, not, or only slightly, : trifid; lower portion with ternate, sometimes quinate, leaves, Peduncles and pedicels rather slender _ with curved pricklets. Rachis straight or slightly flexuous, glabrescent below, felted and pilose above, with several, often stout, strongly falcate prickles; those below often shorter but all very broad-based. PLOTTED ON UTM GRID Ficure 3. Distribution of Rubus anglocandicans A. Newton in the British Isles. A REVIEW OF RUBUS SECTION DISCOLORES P. J. MUELL. IN BRITAIN 245 Petals white, broad, elliptical, ciliate. Sepals greenish to greyish-white, felted, + acute or mucro- nate, reflexed. Stamens white, slightly exceeding the green styles. Carpels long-pilose, bearded. HOLOTYPUs: Frilford Heath, v.c. 22, 21/8/1895, W. M. Rogers, as R. thyrsoideus sp. coll., Set of British Rubi no. 116 (MANCH) (Isotypi in BM, CGE, OXF) This is the chief representative in Britain of the series Candicantes, which is widespread in areas of northern and central Europe where R. ulmifolius is absent. It is similar to R. candicans Weihe but differs in leaf-toothing, quality of armature, panicle outline and flower characters. It is also close to some specimens named R. thyrsanthus by Danish authors. A good series of specimens can be seen at OXF. It is noticeable that R. anglocandicans presents a ‘Continental-type’ distribution, being confined to a belt of country from Berks. to S.E. Yorks., where the terrain is similar to that of northern Germany, and chiefly on sandy soils; it is particularly frequent in the northern Cots- wolds and along the lower Trent Valley. The known distribution is shown in Fig. 3 and includes VC. 22,1235 29; 30... 32,533,358, 39,537, GL: 19. R. hylophilus Rip. ex Genev. (RX. brittonii Barton & Riddelsd.) British specimens of R. brittonii match well examples of R. hylophilus Rip. ex Genev. in K and Bat. Eur. no. 366 in BM and MANCH;; the use of the name R. hylophilus is therefore justified for the British plant. The account and figure in Watson (1958) give a satisfactory impression of this species. 20. R. thyrsanthus Focke Watson’s single example of putative R. thyrsanthus (SLBID appears to be a depauperate specimen or a recent hybrid derivative of R. armipotens. It is certainly different from R. thyrsanthus Focke, which I have collected from the Harz area in Germany (whence it was first described) and have verified with syntypes. CONCLUSIONS A revised British list of Rubus section Discolores is as follows: Series Gypsocaulones P. J. Muell. (incl. Hedycarpi Focke) R. ulmifolius Schott R. rusticanus Merc., R. discolor auct. R. armipotens Barton ex A. Newton R. pseudobifrons auct. R. winteri P. J. Muell. ex Focke R. rossensis A. Newton R. crassifolius auct. R. lamburnensis Rilstone R. pydarensis Rilstone R. procerus P. J. Muell. R. hedycarpus Focke, R. armeniacus Focke Series Candicantes Focke R. neomalacus Sudre R. anglocandicans A. Newton R. thyrsoideus auct. R. hylophilus Rip. ex Genev. R. brittonii Barton & Riddelsd. 246 A. NEWTON ACKNOWLEDGMENTS I am indebted to the Curators of the various herbaria cited for the loan of specimens, to E. S. Edees for his advice particularly on the treatment of Rilstone’s taxa and for some distributional data, and to the Biological Records Centre for production of the maps from data supplied by me. REFERENCES Danby, J. E. (1958). List of British vascular plants. London. RILSTONE, F. (1950). Some Cornish Rubi. J. Linn. Soc. (Bot.), 53: 413-421. RILSTONE, F. (1952). Rubi from Dartmoor to the Land’s End. Watsonia, 2: 151-162. Rocers, W. M. (1900). Handbook of British Rubi. London. Rocers, W. M. (1905). French and German views of British Rubi. J. Bot., Lond., 43: 198-205. Supre, H. (1904). Observations sur ‘Set of British Rubi’. Bull. Soc. Etud. scient. Angers, 33: 106-145. SuprE, H. (1909). Rubi Europae, (2). Paris. WaTSON, W. C. R. (1928). Brambles of Kent and Surrey, 1. Lond. Nat., 7: 12-17. Watson, W. C. R. (1929a). Bramble Notes. Rep. botl Soc. Exch. Club Br. Isl., 8: 782-788. Watson, W. C. R. (1929b). In TuRRILL, W. B., ed. Report of the distributor for 1928. Rep. bot! Soc. Exch. Club Br. Isl., 8: 904. Watson, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge. (Accepted June 1976) Watsonia, 11, 247-251 (1977). 247 The past and present status and distribution of Sea Pea, Lathyrus japonicus Willd., in the British Isles R. E. RANDALL Board of Extra-Mural Studies, Cambridge University ABSTRACT Lathyrus japonicus Willd. is a rare and declining species in the British Isles. From 1973 to 1975 beaches on which the species had been recorded in the past were visited and the present status recorded. There has been a marked reduction in the number of Lathyrus japonicus colonies and under half of those sites where it had once been recorded now contain the species. However, in E. Sussex, v.c. 14, and E. Suffolk, v.c. 25, the populations are expanding and at Chesil Beach, Dorset, v.c. 9, Rye, E. Sussex and Orford, E. Suffolk its continued existence seems assured. Elsewhere, human pressure or a change in physical conditions could rapidly eliminate the species. The range of a plant species and its distribution within that range are frequently dynamic. This may be the result of physical or climatic changes, successional sequence or, above all, human interference. The Sea Pea, Lathyrus japonicus Willd., is an example of a species easily disturbed by man and it illustrates the care with which he must monitor his effects on the flora even in non- agricultural landscapes. L. japonicus is a creeping or climbing perennial most commonly found on shingle beaches but occasionally recorded from dunes and other coastal habitats. It is fairly long-lived and once estab- lished it is not likely to disappear except where coastal changes or human pressure cause this to occur. Where beaches are accreting seawards and more closed vegetation enters as humus builds up, it disappears on older shingle but persists nearer the shore where the vegetation is open. This is particularly clear at Shingle Street, Suffolk, v.c. 25. A description of this species is given by Bright- more & White (1963), who consider that the British plant is L. japonicus var. glaber (Sér.) Fernald. However, a narrow-leaved variant, L. japonicus var. acutiformis Bab., is present in north-eastern Britain. This may well be a northern ecotype with distinct habitat preferences, since all the Scottish sites for the species are on sand-dunes and the dune plants in northern Jutland, Denmark, are similarly narrow-leaved. Conversely, the Danish plants growing on shingle are broad-leaved like the shingle plants of southern Britain (F. Rose, pers. comm. 1976). L. japonicus is a northern plant with a disjunct circumpolar distribution. It formerly extended south into northern France but it is now extinct there. It is common in Denmark, especially in north Jutland in the dunes of the Hanstholm Nature Reserve and on shingle beaches between Struer and Thisted. With the exception of southern and eastern England, it is rare over much of the rest of the European coast and this is probably explained by the lack of shingle beaches. In Britain it has been recorded most frequently on the southern and eastern coasts from Chesil Beach, Dorset, to Benacre, E. Suffolk, but isolated colonies occur elsewhere. Its seeds are avidly eaten by birds, and many animals, especially sheep, find the whole plant palatable. The plants cease flower- ing and soon die when heavily or frequently trampled. For many years L. japonicus has been regarded as a rare and declining species in the British Isles as excerpts from county Floras show: DORSET ‘Native; pebbly beaches; rare.’ (Mansel-Pleydell 1895) HANTS ‘On pebbly and sandy sea beaches; very rare; only found in I. of W., and has not been seen for many years.’ (Townsend 1904) SUSSEX ‘Native. Shingly shores: very rare. Now very nearly extinct, the specimen at Rye in 1931 being the last seen since 1878. . . . Its shyness in flowering and therefore inability to reproduce itself, and the rapacity of collectors when it does appear, sufficiently account for its disappearance.’ (Wolley-Dod 1937) 248 R. E. RANDALL KENT ‘Native. Shingly shores; very rare. In some stations it seems to be extinct; but it may still be found at Dungeness and Kingsdown, in small quantity.’ (Hanbury & Marshall 1899) SUFFOLK ‘Qn shingly shore; rare.’ (Hind 1889) However, population resurgences do occur and currently all these counties except Hampshire have colonies of some quantity. From 1973 until 1975 the population at Shingle Street, E. Suffolk, was much larger than that recorded in 1962 and Brightmore & White (1963) suggest that an exceptional spread of L. japonicus at Rye Harbour from 1962 until 1964 resulted from dispersal of seeds by flocks of stock dove, Columba oenas L. This colony has remained large to the present day. LATHYRUS JAPONICUS Willd. @ var.-glaber @ var. acutiformis @ Extinct PLOTTED ON UTM GRID FicureE 1. Distribution of Lathyrus japonicus in the British Isles. Map compiled by the Biological Records Centre, Monks Wood. SEA PEA, LATHYRUS JAPONICUS WILLD., IN THE BRITISH ISLES TABLE 1. PLANT SPECIES FREQUENTLY ASSOCIATED WITH L. JAPONICUS IN THE BRITISH ISLES Based on data collected by the author and within 5m square quadrats Agropyron junceiforme Ammophila arenaria Arrhenatherum elatius Atriplex sp. Bellis perennis Beta vulgaris Bromus mollis Cirsium arvense Crambe maritima Echium vulgare Epilobium angustifolium Erodium cicutarium Festuca rubra Lupinus arboreus Medicago arabica Medicago minima Montia perfoliata Plantago lanceolata Plantago coronopus Rumex crispus Sedum acre Senecio jacobaea S. viscosus Silene maritima Solanum dulcamara Tamarix anglica 249 Geranium robertianum Trifolium pratense Glaucium flavum T. repens Hieracium pilosella T. suffocatum Honkenya peploides Tripleurospermum maritimum Lathyrus nissolia Vicia hirsuta Leontodon autumnalis V. lathyroides Lolium perenne V. lutea Because of the uncertainty of the present distribution of this species and the scant knowledge of the population size at any of the recorded sites, a study was undertaken from 1973 until 1975 to assess the distribution and status of L. japonicus on the coasts of the British Isles. Enquiries were also made of many shingle researchers to discover their knowledge of the species’ present distri- bution. Full data from each site have been lodged with the Biological Records Centre, Monks Wood. The following records describe the current situation and trends. Fig. 1 illustrates the present distribution of L. japonicus in the British Isles and Table 1 shows the more important associated species. With the exception of a new site first recorded in 1972 on St Mary’s, Isles of Scilly, v.c. 1, L. japonicus is now absent from all four of its former sites west of Chesil Beach, Dorset; these are Penzance and St Keverne, W. Cornwall, v.c. 1, St Austell, E. Cornwall, v.c. 2, and Charmouth, Dorset, v.c. 9. This area of England has become one of the major tourist regions in the post-war period and a marked reduction has occurred in the quality and quantity of beach vegetation in general. The first record of L. japonicus from Chesil Beach, Dorset was in 1799. Mansel-Pleydell (1895) and Good (1948) mentioned that it grew only intermittently along the beach. In 1973 a zone of L. japonicus was found along much of Chesil Beach just landwards of the crest of the shingle ridge and extending for great distances along the beach. In places it disappzared for some distance only to reappear a few hundred metres further on. This is certainly the largest population of L. japonicus on the south coast of Britain and is rivalled only by the extensive recent spread at Rye Harbour. Chesil Beach has been scheduled as a Site of Special Scientific Interest but it is not within a nature reserve. From Chesil eastward to Brighton, E. Sussex, v.c. 14, L. japonicus has been recorded in seven local- ities but in 1973 the only site found was within the fenced enclosure of Volk’s Railway on Brighton beach. It is now absent from Poole, Dorset, Hurst Castle spit, S. Hants, v.c. 11, East Cowes and Sandown, Wight, v.c. 10, Kingston and Worthing, W. Sussex, v.c. 13. Again, this is an important holiday area but considerable stretches of the coast have also been altered for sea-defence purposes. A small patch, about 1 m2, occurred at Pagham Harbour, W. Sussex, in 1972 but timber dumping had eliminated it in 1973. F. Rose (pers. comm. 1976) reports, however, that it has re-appeared in 1976. East of Brighton, it had been recorded at Seaford, Pevensey, Bulverhythe (until 1960s) and Hastings, all in E. Sussex, v.c. 14, but it is now extinct at each site. In eastern E. Sussex, v.c. 14, and E. Kent, v.c. 15, L. japonicus is much more abundant. In the former it is present in great quantity for over 3 km of shingle from Winchelsea Beach to Rye Harbour 250 R. E. RANDALL and on to Camber Sands, where it becomes a dune species. It is most abundant on new shingle ridges but persists locally as these become covered with blown sand east of the estuary of the River Rother. East of Dungeness power station, E. Kent, there is a colony which has increased considerably between 1946 and 1973, though two other colonies nearby known in the 1940s have now apparently disappeared. Along this stretch of coast, especially at Winchelsea and Dungeness, pressure from tourism and fishing have made some colonies poor or non-flowering, but at Pal- marsh, west of Hythe, a colony first recorded in 1954 on the rifle-range is expanding. Elsewhere in E. Kent, L. japonicus is now absent from its old sites at New Romney, Kingsdown and Reculver, although it still persists at Walmer and at Whitstable, where it was first recorded in the 1970s. Walmer beach is heavily used by holiday-makers but there is a wooden track across the shingle to which most visitors keep. This has reduced the pressure on the beach vegetation and the colony of L. japonicus is expanding. Perhaps this augers well for future conservation of the species and suggests a satisfactory means of reducing trampling on the species at other sites. F. Rose (pers. comm. 1975) saw L. japonicus ‘in local plenty’ about 2 km west of Minnis Bay, Reculver in the early 1960s but inroads by the sea and the erection of a new sea-wall have eliminated it. The only two N. Essex, v.c. 19, sites are both recent ones. Jermyn (1975) mentioned that it occurred at Walton-on-the-Naze in 1964 and that it was still present in 1971; it was also recorded just north at Little Oakley in 1970. All eight sites from which L. japonicus has been recorded in E. Suffolk still contain the species and in several! places, for example Orford Ness, Shingle Street and Benacre, it is increasing. At Landguard Point and Thorpeness the populations are dwindling rapidly because of human pressure. The other sites are Felixstowe Ferry, Minsmere, Dunwich and Walberswick. Records suggest that until about 1850 Aldeburgh was its northern limit in East Anglia but now it seems to be spreading very considerably north to Benacre. The Orford Ness/ Shingle Street area has vast mats of L. japonicus over the apposition banks (Randall 1973); this is the largest population in the British Isles and is only exceeded on the Continent by those along the west coast of Jutland. The Suffolk coast is much less used by tourists than the south coast of Britain, although pressure has increased markedly in recent years. Orford Ness is part of a National Nature Reserve and is virtually inaccessible except by boat, but Shingle Street is unprotected. Caius (1570) noted that between the towns of Orford and Aldeburgh, in the autumn of 1555, peas grew of their own accord in such great abundance as to be sufficient even for thousands of people. This is the earliest record for L. japonicus in the British Isles. L. japonicus seems not to have been part of the indigenous flora of Norfolk in historic times and several attempts to establish it there this century have failed after a few years. At present a small patch about 2 m? still exists on Cley beach, Norfolk, v.c. 27, just west of the coastguard lookout. In Lincolnshire, it is now extinct but there are new records in the 1970s at Warkworth and Birling Links, Cheviot, v.c. 68. Scotland has had three isolated populations of L. japonicus: one by the Firth of Tay, Forfar, v.c. 90, one on Unst, Shetland, v.c. 112, and one by Loch Ewe, W. Ross, v.c. 105. At the first of these populations it disappeared near Monifieth as a result of industrial activity but it still exists in two small areas, one near Carnoustie and the other near Arbroath. Both sites are on golf-links. In Shetland, the population has disappeared at one site through gravel extraction and recreation and at the other it has been reduced to under 2 m2 by similar pressures. The Loch Ewe colony has also disappeared. Hodgson (1899) recorded L. japonicus from Harrington and St Bees on the Cumberland coast, v.c. 70, but these colonies have now disappeared, as has also that at West Aberthaw, Glamorgan, v.c. 41, as a result of the construction of a power station and heavy use of the area for fishing. However, a single plant was reported at Seascale, 15 km south of St Bees, in 1975. For many years L. japonicus was recorded at Rossbeigh, S. Kerry, v.c. H1, and also across the bay at Inch. It has not been seen at the latter site since before 1916 and although collections have been made from Rossbeigh in recent years, it was not found in 1973 (Scannell per. comm. 1975). Thus it can be seen that there has been a drastic reduction in the number of L. japonicus sites although the species has certainly increased locally on the Sussex and Suffolk coasts. The dis- appearance of L. japonicus from many of the remaining areas can be attributed in part to shyness of flowering (Lousley & McClintock 1951) but above all to human use of the beaches concerned. It seems that the present climate in southern and eastern England would enable it to grow very vigorously were it allowed to do so, but this may not be the case further north or west. It seems SEA PEA, LATH YRUS JAPONICUS WILLD., IN THE BRITISH ISLES 251 that shingle extraction is totally responsible for its elimination from its only French locality, on shingle south of the mouth of the River Somme. Lack of grazing in the twentieth century may well have-helped the species considerably in southern and eastern England. Where trampling does not occur, such as at Walmer, even heavily-used beaches retain good populations. As it is a plant of the mobile foreshore, there are bound to be natural changes or fluctuations in its distribution, and some extinctions can be attributed to local land-erosion. However, the most stable sites physically are those where the vegetation is more closed and where there is greatest risk from human activity. Thus the long-term outlook for the species in many areas is bleak. The populations at Chesil Beach, Rye Harbour and Orford Ness/Shingle Street are by far the largest and are the only ones in which its continued existence seems assured, unless deliberate protective measures are taken else- where. ACKNOWLEDGMENTS This research was financed by the World Wildlife Fund. I should like to thank Dr F. Rose and Dr F. H. Perring for their criticism of this study and for help in obtaining data. REFERENCES BRIGHTMORE, D. & WuirteE, P. H. F. (1963). Lathyrus japonicus Willd. in Biological Flora of the British Isles. J. Ecol., 51: 795-801. Catus, J. (1570). De rariorum animalum et stirpum historia. In De canibus britannicus. London. Goon, R. (1948). A geographical handbook of the Dorset flora. Dorchester. HANBURY, F. J. & MARSHALL, E. S. (1899). Flora of Kent. London. HIND, W. M. (1889). Flora of Suffolk. London. Hopcson, W. (1898). Flora of Cumberland. Carlisle. JERMYN, S. (1975). Flora of Essex. Fingringhoe. Lous Ley, J. E. & McC.uintock, D. (1951). In WALLACE, E. C., ed. Plant Records. Watsonia, 2: 41. MANSELL-PLEYDELL, J. C. (1895). Flora of Dorsetshire, 2nd ed. Dorchester. RANDALL, R. E. (1973). Shingle Street, Suffolk: an analysis of a geomorphic cycle. Bull. geol. Soc. Norfolk, 24: 15-35. TOWNSEND, F. (1904). Flora of Hampshire, 2nd ed. London. WoLLEyY-Dop, A. H. (1937). Flora of Sussex. Hastings. (Accepted June 1976) b: + 157 oer) er err 1 rr L208 tas 21 Te HeITISM BUT WP “pam apMeN \e 7 10. VRIOL dom M Ito 21h ATWO, ROL Usninoite etl 202 ake ; r Vos i) 190 fl puta ti ast i peasy 6" eee ] a oH HOGI. 10 ore. sib a atio Tint od Ww mil ble AP ag fst) ba Teoh . | . ads ot | ako in trary? & af feed rons Tilodans) vie bu nleiste os Aiba. is LYE aH reve : ee Five ih. (Puce 271 ae anos bout 4 1 te) Joan) Bega ; ‘ ‘ rd Reet il q eatie videre Renny St avaroHt aoieor- -bont iol ott i) 10h 2 ibs oe pe of ivGse npttiod COT! dal? Ted) ti NPI FU oily fits aed Shc re toi T ay uy : ; Héietiog Sik iss aj as Y A Sart At Baga | ar 40%. on he > ure, = rs brie eda PGT 38} if wie! Tesla Sanide\as ah ( Bi0T °) Aer AgY St. Sotuesem SVviId9I8 Sis) sion gaat, Ds vilen asi an: ip Roden. ee meats a Naw yer ‘i sciibe mh elt : tye he inch vegeta tit vigth : { out 5 as vu) a ir te auecrwation 0: . The Py . >. ~Y" if A a may » > Ww tar ycy i LT eae ee aA DAE Oe Lens ihe rd, “> at ther 2 “a4 re Weeld { Mibiyais May: Senay vee Ty ry Aigote T ba ABV BAOW Bay} Le hai Sa rai ye iS ase 01 ah ret, N00) (SG YOLDge Vous i 6 petal ii F g bi ey , r ts “~; jenpormtic ts hans NOGUL TO Lit ‘ y =) “ee Lhe pe dag Hai ae tris | m, ' at ifrt LO fi i CAbTe oy. DHLVY sake Subba, fas te iy ft bal a8 ana: ; , | | oh As iaeabesy ee, Sycws.D ects ipt ied Ns SEL: bas AON bl wore \s Witonnls yeti Lined sea tery i Twetaerhi pad], gy quan HAithis Agvet listewh bro-itelelaieg. bw cetety i) phalndlianad bee: yal O08) (Asana Mie bY Age st fn Moonod pki’ Na seek, i 06 ro e ajuk 1.) 7, tent Luger ‘ee aS) Rw woh CH) Wi, 48 ra « epodamizani | iene mca | od bt MOAI ol Spent r AenTIY 1 be ine ae I 6 gt ve th pt ABS. . WS Sy WHE Shoes SaNTIGMOIg a TO ie ny 1k 1B 7 one. yee ape eee ia (ah ARH arta 547 We hae alata Yoo aantoisoenys j 1s ORR iets re OAT (URC EECA . ; np “> vic. Bis juat of ibe = : » SONCL w wh hrarnosahd how recede: int we 19708 4 We at . 7 a ye 2 ° 8 | poinshetiors’ af aa pons One by te "hee sdb Ty Biwey, Ww, ‘Rookwa ‘a Mant = we fe ts aah of induste’ activated wine tre t ew pear Arbroath,’ Bis be wh “ perarey. ai ca sy es a) Urtcagla ur: s a a or) fe pale bee th i uit ivy sear pn we MSs: he, ook. & ‘red uy a“ hho ae oa iit ee. sone (1396) poured Bod ss! Ct 1 Beerinagoo:an in > fat e rmoner han aw ee dy glee abt Ve a), Ries a of the eenatrustion ol a POWe - x0 CIvweve ey a Let 9 rengrye “cf ie Sami . oe oa ed yoart f fee ae Witt ieee sorabst x fh eas agit. Damopemnadh: Kvecin ees, Ht thon he a betes iy ses | » CONN. Row bes MF hay Pero’ P bsiesdh . VIRATAR bmSlILA i aa a aie ot the! tintre hig 9 nies ni atchinat's is Syecien tras centage a TERY rin | Latest of ares Spon a any O88 hi ee | ws ’ -1 es 1 ytd # ‘ i, aa t i me hyn he ihe Tits y be ions rita ¥. ie Pin ar | wy 48 a caan , Watsonia, 11, 253-257 (1977). 253 Short Notes RUBUS SPRENGELII WEIHE IN SCILLY After many years of combing Cornwall and western Devon for brambles, Rilstone (1952) was able to report R. sprengelii Weihe—perhaps the most distinct of all the British species—in only a single locality: near Penzance, W. Cornwall, v.c.1. There was also one old record from Valency in the literature that he believed correct. In view of this great rarity in the Cornish peninsula it was with some surprise that in a fortnight’s visit to the Isles of Scilly in June 1975 I came across the species in no fewer than four places. Three of these were on Tresco: small patches by two successive cross-paths on Middle Down and a patch bordering bracken by the south tip of the Great Pool. On St Mary’s, which I sampled much more cursorily, it was frequent under bracken round Innisidgen burial chamber. Similar habitats occur in plenty almost throughout the islands and there seems no reason why the species should not prove widespread. On the other hand it was not among the Rubi that F. Townsend collected on his major exploratory visit in 1862—the only extensive sampling of the Scilly brambles hitherto (Lousley 1971). There are other plants, such as Ornithopus pinnatus (Mill.) Druce, that occur widely in Scilly but are missing from the Cornish mainland, but it is unexpected that a member of a genus so readily dispersed by birds should exhibit this pattern. REFERENCES LousLey, J. E. (1971). Flora of the Isles of Scilly. Newton Abbot. RILSTONE, F. (1952). Rubi from Dartmoor to the Land’s End. Watsonia, 2: 151-162. D. E. ALLEN CAREX POLYPHYLLA KAR. & KIR. AND C. LEERSIANA RAUSCHERT Since the writing of David’s paper (1976) on the nomenclature of the Carex muricata L. aggregate, we have, through the kindness of the curator of the herbarium of Moscow University, been able to examine the type material of C. polyphylla Kar. & Kir. in M“W. There are two sheets, both carrying the tickets of Karelin and Kirilov, 1841, and both determined as C. polyphylla Kar. & Kir. by Kreczetowicz in 1937. They bear no localities, but can be assumed to originate ‘in sylvaticis ad torrentem Tscheharak-Assu, circa montes Tarbagatai, 1840’ in E. Kazakhstan. These are the only specimens collected by Karelin and Kirilov that are extant and that can be considered type material. One, numbered 2075 and 227279, is labelled ‘Isotypus’ in recent hand-writing, and contains one specimen so incomplete that we cannot be certain what it is. The other, numbered 2081 and 227274, is labelled “Typus’ in the same recent hand-writing, and contains two very adequate specimens, in excellent condition, which we here take the opportunity of designating as Lectotype. (Alekseev (1973) commented that one must reckon as the type the specimen in MW bearing both Karelin and Kirilov’s and Kreczetowicz’s labels, but did not mention that there are two sheets to choose from.) The lectotype material corresponds in every respect with C. leersiana Rauschert (C. leersii F. W. Schultz, non Willd.). Further, more recent, collections in MW, determined as C. polyphylla by Kreczetowicz and others, from the southern parts of the U.S.S.R. ranging from the Crimea to Lake Baikal, exactly match the lectotype and indicate that this taxon is widespread in the area. Thus, if C. leersiana Rauschert is considered as a distinct species, its correct name is C. polyphylla Kar. & Kir. The extreme form of the taxon (as in the type) is indeed very different from the extreme form of C. divulsa Stokes, but these extremes are linked by a series of intermediates so that no clear morphological dividing line can be drawn. The same conclusion was reached by Medovié (1960), E 254 SHORT NOTES who, in a comprehensive series of statistical analyses, failed to separate C. leersiana from C. divulsa. In our opinion, the only possible course, unless further studies succeed in establishing some means of separation, is to regard C. polyphylla Kar. & Kir. as a subspecies of C. divulsa, and at this rank the correct name of the taxon remains C. divulsa subsp. leersii (Aschers. & Graebn.) W. Koch. REFERENCES ALEKSEEV, YU. E. (1973). Carex polyphylla Kar. et Kir. i blizkie ej vidy iz vostochnoj Evropy. Vest. mosk. Univ., nauch. Zh., ser. 6, 28 (5): 66-71. Davip, R. W. (1976). Nomenclature of the British taxa of the Carex muricata L. aggregate. Watsonia, 11: 59-65. Mepovicé, J. (1960). Suborny druh Carex muricata L. na Slovensku. Acta Fac. Rerum nat. Univ. comen., Bratisl. (Bot.), 5 (4-2): 1-116. R. W. Davin & A. O. CHATER CHROMOSOME NUMBERS OF BRITISH PLANTS, 5 Chromosome Grid reference and locality number Hippocrepis comosa L. n=7 21/89.76 St Bride’s Valley, Glamorgan, v.c. 41 2=7 21/57.87 Pwlldu, Gower, Glamorgan, v.c. 41 ne 20/92.62 Daddyhole Plain, Torbay, S. Devon, v.c. 3 Juncus articulatus L. 2n = 80 28/50.57 Loch Ussie, E. Ross, v.c. 106 2n = 80 35/83.31 Sand Sike, Upper Teesdale, Durham, v.c. 66 Carex paupercula Michx. n= 2 35/81.29 Cow Green, Upper Teesdale, Westmorland, v.c. 69 n= 29 26/68.32 Glengavel, Lanark, Ve Ty Alopecurus alpinus Sm. n = 50 35/70.33 Little Dun Fell, Westmorland, v.c. 69 2n = 117 37/16.76 Glas Maol, Forfar, v.c. 90 G. M. FEARN PHYTEUMA SCHEUCHZERI ALL. NATURALIZED IN OXFORD Phyteuma scheuchzeri All., a native of the southern Alps from Piedmont and Valais to Croatia, and first recorded in cultivation in Britain in 1816 (Sims 1816), has in recent years become estab- lished on the wall of St Johns College gardens, Oxford. A specimen collected by me in 1960 is in OXF. Allied to P. tenerum R. Schultz, it may be distinguished as follows: Basal leaves (in the Oxford plant) ovate to lanceolate, long-petioled, cordate to truncate at base; stem-leaves few, linear-lanceolate, distinctly stalked; leaves all with shallow, distant teeth, or the uppermost subentire. Inflorescence capitate. Bracts linear-lanceolate, conspicuous, much exceeding inflorescence, often curving downwards. Corolla deep blue, straight in bud; stigmas three. Flower- ing late May to June, the withered heads persisting. From enquiries made, P. scheuchzeri seems to have first appeared on the wall about 1951. It evidently originates from St Johns College gardens to which, as Mr K. E. Bull has suggested to SHORT NOTES 255 me, it may have been introduced by Rev. H. J. Bidder (1847-1923), a former bursar of the college who collected many choice plants in the Alps and Pyrenees for the rockeries (Druce 1924). But it has now spread to the outer side of the wall, both in Parks Road and the court of the School of Agricultural Science, and to pavements nearby, and seems at least as worthy of record as such aliens as Chondrilla juncea L. and Melica ciliata L., which temporarily established themselves on Oxford walls in the past (Druce 1886). REFERENCES Druce, G. C. (1886). The flora of Oxfordshire, pp. 179, 342. Oxford. Druce, G. C. (1924). Obituary of Rev. H. J. Bidder. Rep. botl Soc. Exch. Club Br. Isl., 7: 148-149. Sms, J., ed. (1816). Curtis’s Botanical Magazine, 43: t. 1797. R. C. PALMER WHITE-FLOWERED BLUEBELLS (ENDYMION NON-SCRIPTUS (L.) GARCKE) Despite the wide distribution and abundance of Endymion non-scriptus (L.) Garcke in Britain, very little is known of the white-flowered form, which appears to be equally widespread although sporadic and never common. The following observations on this form have been prompted partly by Harborne’s (1967, p. 257) statement that a general ‘characteristic of white flower mutants of coloured forms is their unthriftiness as plants’. A population of white-flowered plants in a wood north-west of Cronton, S. Lancs., v.c. 59, was studied between 1967 and 1971. Although the wood contains an estimated 60,000 blue-flowered plants, the number of white-flowered individuals seen annually ranged from 16 to 22. Populations with this proportion of white-flowered plants appear to be fairly general. Over the above five-year period counts were made of the number of flowers, capsules and seeds per inflorescence. It was found that white-flowered plants had a mean of 4-7 flowers (blue-flowered 5-5) 2-1 capsules (3-7) and 23 (45) seeds per inflorescence, the average number of seeds per capsule being rather similar in each. It was not possible to follow all the original flowering plants through to seeding, so the sample-size progressively decreased. Four plants of each colour, transplanted to a garden from the Wrekin, Salop, v.c. 40, produced the same number of flowers per inflorescence, although the white-flowered plants had fewer cap- sules. It may be that the latter plants flower less freely when in competition with blue-flowered ones in the wild. Germination of seeds collected in the wild from white- and blue-flowered plants and sown in a garden was similar, mostly 20-30% after eight months from collection. All 26 plants raised from seeds collected from white-flowered individuals proved ultimately to be blue-flowered. These observations confirm Harborne’s suspicions on the generally lower reproductive efficiency of albino plants and they also demonstrate a high level of outbreeding in the bluebell. REFERENCE HARBORNE, J. (1967). Comparative biochemistry of the flavonoids. London & New York. R. W. RIDING POLYGONUM MINUS HUDS. x P. PERSICARIA L. IN ANGLESEY In July 1975 a plant which appeared to have a combination of the characters of Polygonum minus Huds. and P. persicaria L. was found on the muddy shore of Llyn Coron, near Aberffraw, Anglesey, v.c. 52. This is the only known Anglesey locality for P. minus, but this species occurs here in good 256 SHORT NOTES numbers and often grows intermingled with P. persicaria, occasionally with plants of the two species actually touching. The features which made the plant conspicuous in the field were the decumbent habit and narrow, interrupted inflorescence as in P. minus, and the blotched leaves as in P. persicaria. Although a careful search of the site was made, no other plant of the same kind was seen. As the plant was branched from the base and rooting at the lower nodes, a portion was removed and potted in compost kept moist by standing in a pan of water, where it grew vigorously. Further observations on this piant showed that it was intermediate between the putative parents in a number of characters. A comparison of it with P. minus and P. persicaria from the same locality is shown in Table 1. TABLE 1. COMPARISON OF POLYGONUM MINUS HUDS., P. PERSICARIA L., AND THE PUTATIVE HYBRID P. minus Hybrid P. persicaria Habit Decumbent Decumbent Upright or ascending Stem Slender Intermediate Stout Inflorescence Very slender (c 3 mm Slender (c 5 mm wide); Stout (c 7-9 mm wide); wide); interrupted at interrupted at base scarcely interrupted at base base Leaf width Up to 7°5 mm Up to 9 mm Up to 12 mm Leaf length: width ratio 7-8 4-5-5-5 4-5-2 Leaf venation Inconspicuous beneath __ Fairly conspicuous Conspicuous beneath beneath Leaf marking None Blotched Usually blotched Perianth colour Pink Greenish, tinged with Greenish, tinged with pink pink Seed length and breadth 1:7x1mm 2:1 1-3 mm 2°6x 2:1 mm (mean of ten seeds) The pollen appeared to be normal, though the pollen grains were smaller (c 40 um diameter) than in either of the parents (c 50 um in P. minus; c 44 um in P. persicaria). Seed production also seemed to be normal, but the seeds were intermediate in size and shape between those of the parents. Herbarium specimens taken from the plant later in the season were sent to Dr J. Timson of the Department of Medical Genetics, University of Manchester, who (Timson in /itt. 1975) agreed that ‘it may well be the hybrid since all the characters are intermediate’. The specimens have been deposited in NMW. Timson (1965) has pointed out that interspecific hybrids in Polygonum Section Persicaria appear to be extremely rare in Britain; out of seven herbarium sheets of putative P. minus x P. persicaria examined by him, only one could be identified as the hybrid and that rather uncertainly. This specimen (BM) was collected by B. Welch in September 1955, on Ham Common, Richmond, Surrey, v.c. 17. There are unsubstantiated records of this hybrid from v.c. 11-14, 22 and 23 ({Timson 1975), but, apparently because of his uncertainty, Timson (1975) did not cite the Surrey record. The chromosome numbers of P. minus and P. persicaria are very close to one another: P. minus has 2” = 40, P. persicaria has 2n = 44. It is possible, as Timson (1965) has suggested, that their hybrid may be capable of some satisfactory meioses and thus account for the production of the normal-looking pollen and seeds. REFERENCES Timson, J. (1965). A study of hybridization in Polygonum Section Persicaria. J. Linn. Soc., Bot., 59: 155-161. Timson, J. (1975). Polygonum Section Persicaria, in STACE, C. A., ed. Hybridization and the flora of the British Isles, pp. 274-277. London. R. H. ROBERTS SHORT NOTES 257 VULPIA HYBRIDA (BROT.) PAU, NOMEN AMBIGUUM In an earlier note (Stace & Cotton 1976) we pointed out that the plants which occur in this country as wool aliens under the name Vulpia australis (Steudel) Blom are in fact identical with the southern European V. broteri Boiss. & Reut., for which the correct name is V. muralis (Kunth) Nees. Our reasons for using the name V. muralis were not, however, stated adequately, and the purpose of this note is to rectify the omission. The basionym of V. muralis (Festuca muralis Kunth) was published in 1822, and easily predates the publication of V. broteri Boiss. & Reut. in 1852. However, the name V. hybrida (Brot.) Pau, based on Festuca hybrida Brot., published in 1804, has also been used for the same plant and, of course, if it could be typified accordingly it would be the correct name. There is, unfortunately, uncertainty as to the application of the name Festuca hybrida, as indicated by Boissier & Reuter’s (1852) unwillingness to use it for the plant they named V. broteri. As pointed out by Hackel (1880) and by Kerguélen (1975, p. 285), it seems that Boissier & Reuter considered that Brotero confused two plants, or that his description and type did not match. It appears that Brotero’s herbarium has been lost or destroyed (Paunero 1964, p. 94; A. Fernandes in litt. 1973) and, in view of this and the uncertainty of the application of the name F. hybrida, we (like Paunero) prefer to treat it as a nomen ambiguum. Until such time as Festuca hybrida can be typified, the name used for the plant in question must be V. muralis (Kunth) Nees. REFERENCES Bolssier, E. & REUTER, G. F. (1852). Pugillus plantarum novarum africae borealis hispaniaeque australis, pp. 128-129. Geneva. HACKEL, E. (1880). Catalogue raisonné des Graminées du Portugal, p. 24. Coimbra. KERQUELEN, M. (1975). Les Gramineae de la flore francaise essai de mise au point taxonomique et nomen- claturale. Lejeunia, 75. PAUNERO, E. (1964). Notas sobre Gramineas, 2. Consideraciones acerca de las especies espafiolas del genero Vulpia Gmel. An. Inst. bot. A. J. Cavanillo, 22: 83-114. StTAcE, C. A. & CoTTON, R. (1976). Notes on two alien Vulpias in Britain. Watsonia, 11: 72-73. C. A. STACE & R. COTTON Watsonia, 11, 259-270 (1977). 259 Book Reviews The Naturalist in Britain. D. E. Allen. Pp. xii+292, with 13 black and white plates and 16 line illustrations. Allen Lane, London. 1976. Price £9-00. : The Naturalist in Britain is the unusual subject of this book, which traces the development of studies in natural history in the British Isles from 1600 to the present time. The 280 pages of concise and authoritative text are the results of 20 years’ research by the author, David Elliston Allen, who, _ with his special interest in social history and experience in geographical recording of man and _ other subjects, is well qualified to tackle the wide ramifications of his chosen topic. The author’s link with the B.S.B.I. is revealed on page 1, where he tells us that the idea of the book arose from his small exhibit on the Society’s membership (in 1952), showing how the alteration of its social composition had closely followed broad national patterns. The organization of studies in natural history is found to interrelate with the social fashions and economic conditions of past centuries, and Mr Allen describes the origins of societies for botany, geology, entomology, and taxidermy leading to ornithology and allied subjects. Also, the | history of the distinctive tools of each calling (e.g. hammer, vasculum and butterfly net), whose use was in many cases furthered by eccentrics who stamped their powerful personalities on the studies of their day, is traced. Groups of enthusiasts formed the early societies; the account shows how much of our present comprehensive knowledge of our native flora and fauna is based on the pains- taking observation and record-keeping of the early collectors and the continuity of their records through four centuries through these Societies, of which the first in Britain was the Temple Coffee House Botanic Club. Later the drawing-room hostesses originated ‘conversazione’—a word still used in this society’s calendar of events in most years. In turn the impact of natural theology, the turbulent introduction of Linnaeus’s new method, and the romantic movement of early Victorian times, followed by the links with professional science, are demonstrated. The influence of the _ fashionable collectors, peers of the realm and private patronage are then recounted with humour _ anda wealth of anecdote. Other topics woven into the narrative include transport, photography and publications, all with facets relevant to the theme. : From the 17th century, botanists were to the fore due largely to the practical application of : botanical studies through medicine, and there are evocative descriptions of days of simpling and herbarizing, also of expeditions in swaying horse-carriages (no travel pills in those days!), and tales of hardship and endurance associated with the botanical excursions. But one can envy the botanical discoveries of early expeditions and feel a link with the Cambridge undergraduate of 1755 who was ‘seized by the challenge of finding and adding to the localities for the rarer plants recorded by Ray in his early Flora of the area’. But we can claim a change of attitude from the botanist of whom it was said ‘he has charge of the physic garden of the apothecaries but treats it right scurvily, allowing no plants to multiply but either sticking them into his books or sending them away in exchange for others’. Five hundred individual naturalists are mentioned by name, but this book is by no means a cata- logue. The pages are packed with facts and information: one learns that ‘the accession of George III in 1760, had among its many effects, the minor and rather freakish one of placing at the nation’s head for the brief space of just two and a half years a trio of enthusiastic botanists’, or that English- women had a reputation for ‘strolling excessively in their garden—the French claimed it gave them big feet’. In addition to a careful index, particularly useful are ten pages of notes on sources which show the wide range of research, much of the material being hidden in little-known papers, the relevant significance of which this book brings to light. Some intriguing period illustrations add to the enjoyment. This book has been received with well-deserved wide interest, covering as it does a field of research 260 BOOK REVIEWS which is not only unique but one with many ramifications. It is one for both browsing and reference, and I would suggest that it will be a source of information and pleasure for many years ahead—by which time its present somewhat breath-taking cost will most probably be regarded as a mere trifle. M. BRIGGS The Seeds of Dicotyledons. A. J. H. Corner. Vol. 1, pp. xii+311; vol. 2, pp. viiit+ 552 (647 figs). Cambridge University Press, Cambridge. 1976. Price vol. 1, £15-00; vol. 2, £25-00. Professor Corner’s new topic is the structure of the seed-coat; his theme is again the phylogeny of the flowering plants. The taxonomic use of seed shapes and surfaces is commonplace, and in recent years such studies have often been supplemented by beautiful photographs from the electron scan- ning microscope. Corner’s subject-matter lies far from such superficial detail; it is the basic anatomical structure of the seed-coat and more especially that of seeds derived from ovules with two integuments, these being characteristic of most of the families we believe to be primitive. Each integument, of course, has an inner and an outer epidermis and between them is mesophyll. Thus there are six histological layers whose differentiation needs to be traced from ovule to seed. In addition there are the possible complications of arils, chalazal growth, vascularization and micro- pylar detail. One thing, then, is beyond dispute: seed structure is as much an integrated system as flower-structure; it is not to be dismissed as a unitary character. Argument and discussion occupy 60 concentrated and often provocative pages. The descriptive data (for the most part new) take up the remaining pages of volume 1. The whole 552 pages of volume 2 are devoted to the illustrative line drawings, many of them of great complexity—and often of fascinating beauty (e.g. Figs. 37 and 38 of Bixa, Fig. 94 of Sarawakodendron). These are, of course, Corner’s own work. The whole book is a tremendous achievement. £40 is a high price for two books, but is it so much for the results of many years’ devoted, original and stimulating research ? The data that are fundamental for Corner’s thesis relate to the seed-coats of Magnoliales. These are shown to be both complex and diverse in respect of which layer or layers show mechanical strengthening; in fact eight main types are recognized, but the basic structure is constant within a family. Corner’s conclusion is that the proto-angiospermous ancestor of these living families had a seed which developed a seed-coat of considerable complexity, and that the Magnolialean types represent different phyletic lines of simplification. These lines show further simplification in more advanced families, but nevertheless structure is often persistent enough to enable affinities to be traced. it is suggested that the nutmeg family, Myristicaceae, shows the closest approach among living dicotyledons to the seed structure of the early angiosperms. For clarity of description Corner reserves the terms inner and outer integument for the ovular structures; for the seed he changes to testa for the outer and tegmen for the inner. Thus a meso- testal seed has its predominant seed-coat layer developed from the mesophyll of the outer integu- ment of the ovule, an exotegmic one from the outer epidermis of the inner integument. When reviewing Takhtajan’s Flowering Plants, Origin and Dispersal a few years ago, I criticized that author for apparently relying solely on pollen structure to transfer Illiciaceae and Schisan- draceae from Magnoliales (monocolpate) to Ranunculales (tricolpate). Perhaps I owe Takhtajan an apology, for Corner makes the same move on the basis of seed-coat structure; however, Corner includes the monocolpate Winteraceae in this transfer, taking seed-coat in this case to outweigh both pollen and the presence of aromatic oil-cells. There are a number of other suggestions that will cause some surprise: Cruciferae are perhaps linked to Polygalaceae, rather than to Papa- veraceae or Capparaceae; Proteaceae may be linked to Papaveraceae. There are many more and it is clear the the digestion of this new information into a general system will require careful thought. Although Corner clearly feels that the families commonly grouped as Magnoliales (in the widest sense) need realignment into a number of distinct orders, he very wisely refrains from any name- making at this stage. He refers repeatedly to the limited material available to him and clearly there is much more to be done before the complete implications of this line of research are fully realized. Corner adduces from these studies of seed-coats much support for his views on angiosperm origins, which are commonly known as durian-theory. This he is well entitled to do. However the BOOK REVIEWS 261 issues must not be confused. This work on the structure of seeds stands as a major contribution to botanical knowledge in its own right. The taxonomic scope of the book will make it difficult for those who lack some knowledge of tropical and subtropical families; yet in this very fact it provides cogent argument for the view that the understanding of the angiosperms lies in tropical botany. But Corner does not neglect the temperate: Aquilegia, Althaea, Berberis, Chelidonium, Euonymus, Helleborus, Hypericum, Oeno- thera, Paeonia, Polygala, Ruta, Tilia, Vancouveria are all here in novel detail. I have a personal dislike for the larger page which demands a double-column setting. Good English deserves a better fate than burial in a narrow column. That apart, the Cambridge Uni- versity Press have done a very good job; misprints are trivial and the reproduction of the drawings is constantly excellent. This book is essential study for anyone interested in the evolution of the angiosperms; it is essential for any department teaching angiosperm classification; it is a splendid example for students of what can still be done by individual research. B. L. BurtTT Biological Identification with Computers. Edited by R. J. Pankhurst. Pp. x+333. Systematics Association Special Volume No. 7. Academic Press, London. 1975. Price £11-00. Those readers of Watsonia who are not at ease with the computing mathematical component in many recent taxonomic papers should read some (or parts of some) of the 16 contributions to this symposium volume (papers presented at a meeting of the Systematics Association in Cambridge in 1973). The remainder should read them all. I cannot summarize the detailed contents here; instead I shall try to encourage non-specialists to consult the volume, since the principles underlying so many of the papers must inevitably permeate the practise of taxonomy, to its everlasting benefit. Computer programs are not themselves mathematics. They are the mode of communicating the sequence of instructions to a computer to enable it to perform certain calculations or data re- arrangements which might well be too massive or complicated for an unaided or mathematically ignorant individual. But an initially less obvious benefit is that the methods of analysis require us to prepare our data in a strictly logical manner, indeed to compare like with like, and to define and standardize our terms. We need to have no fear of becoming automatons ourselves, since in spite of the hopes of the first numerical taxonomists, humans will almost always collect the initial data more or less subjectively, and must make equivalent interpretations at the end of the study. Chapters 1, 2 and especially 14 outline the general principles of biological identification, and distinguish between classification (placing individuals along with others in more or less homo- geneous groups) and identification (finding which group an as yet unclassified individual belongs to). An invaluable disciplinary benefit of incorporating computing into a taxonomic study is that one is forced into producing a full primary data matrix (i.e. all characters scored for all entities being studied). Deficiencies in the data matrices often mean that we have better means of analysis than data to analyse. A depressing conclusion is that many practising taxonomists are surprisingly illogical in their descriptive work and indeed frequently fail to eliminate avoidable errors. I would add from my own experience with bryophytes that a number of genera considered ‘critical’ owe this to poor quality keys rather than to any intrinsic taxonomic difficulties. Chapters 3 to 12 discuss (sometimes fairly technically) programs and techniques for key con- struction and automatic identification. The polyclave keys discussed in chapter 5 avoid the short- comings of traditional dichotomous keys, and allow characters to be employed in any order, some even permitting the comparison of a range of characters simultaneously. Many of these keys can be prepared by hand, and the majority can be used independently of any computer. They go a long way to dissociate keys for identification from systems of classification, with which they have been so confused in the past. Although medical diagnosis is otherwise excluded from the present volume, chapters 8, 9 and 11 make use of studies in bacteriological identification to illustrate some important contributions to general classification and identification theory. Microbiologists often need to search through great quantities of data, and their identifications really matter. Does it ‘matter’ so much that the Sali- cornia I have just eaten is S. fragilis rather than S. obscura? Maybe the precise strain of Escherichia coli (or worse) sitting on it matters more... 262 BOOK REVIEWS Summaries usually make easier reading than the material they summarize, so I recommend non-specialists to read chapter 16 and the four subsequent discussion papers. The volume concludes with a bibliography of papers on computing and identification, a list of relevant computer programs (directly concerned with identification or data capture) and a valuable glossary of terms. Incident- ally, these terms used by computer scientists are a study in themselves—are our data so wild as to need capturing? Perhaps they are. D. H. DALBYy The Natural History of Upper Teesdale, revised edition. Edited by M. E. Bradshaw. Pp. 75. Durham County Conservation Trust Limited. 1976. Obtainable from the Hon. Sales Officer, 1 Abbey Road, Darlington. Price 45p. This paperback booklet supplants the first edition of 1965 and within 61 pages of text (plus Notes for Visitors, Preface and Appendices of Latin names) covers, in seven chapters, a range of natural history subjects. The flora, including its history and vegetation (M. E. Bradshaw and W. A. Clark), has received very full treatment—logically, since Teesdale is internationally famous for this, above all else. Sadly, if typically, few concessions are made to the increasing number of amateur bryologists who will visit the area, and none at all to lichenologists. At least the bryophytes, in my experience, supplement many of the commenis made about the more spectacular flowering plants. Mankind (H. G. Proctor) is a chapter on the history of settlement and population movements which vividly captures the imagination, although the reader is left with rather vague impressions (chapter 7) about post-1905 and present-day activities of Teesdale residents. A supplementary short chapter on current land-use, and the influence on human resident populations of increased visitor- pressure, would have been welcome. The Invertebrates (J. C. Coulson) concentrates, understandably, on insects and should encourage entomologists, arachnologists and others to scurry up the Fells with nets and jars and add more interesting species to the English or British fauna. This is less likely among the vertebrates, the chapter on which (C. S. Waller and D. T. Crisp) will probably be of most use to ornithologists. The regional geology is fully and plainly laid out by G. A. L. Johnson, but I would have wel- comed a few illustrations of the more conspicuous fossils, and perhaps a selection of mineral crystals, which are often seen in the exposed limestones and numerous spoil-heaps of the area. The final chapter, on conservation (D. J. Bellamy), while dealing in a full and balanced way with the problems of Teesdale in particular, has much wider implications. One hopes that his cautiously optimistic note (p. 67) will be supported by future developments. There are fine illustrations by P. Nicholas, and the drawings by M. E. Bradshaw of Teesdale plant specialities, although of simpler line, adequately capture their habits. The general map (p. 4) is, however, somewhat diffuse, and would benefit from bolder treatment of the Cow Green Reser- voir, settlements, paths and boundaries, which are rather faint in this reproduction. The accent on birds, too, I find a little irritating, and their appearance at the head of chapters on invertebrates, flora and conservation in addition to their more logical one above Vertebrates, seems inappropriate (why not, for example, a Northern Eggar or selected pollen grains in the appropriate places ?). The booklet is packed with interesting information for the field and, to a lesser extent, armchair naturalist. Its overall tone seems to be aimed primarily at the educated visitor having some previous experience or training in a biological or geological field, but it has much to offer the casual layman. I believe it will prove to be an essential companion to the handbooks of the field naturalist visiting Teesdale and, moreover, at a price that is very favourable by any standards. A. EDDY Flora van Nederland. H. Heukels, 18th edition. Edited by S. J. Van Ooststroom. Pp. 913, with 1 map and 1,038 line-drawings. Wolters-Noordhoff, Groningen. 1975. Price Dfl. 43-25. The latest edition of this well-known Flora of Dutch vascular plants is again edited by Dr Oost- stroom. That it is only two years since the 17th edition is presumably testimony both to its popu- larity and the frequency of revision. ' BOOK REVIEWS 263 In his Foreword, Dr Ooststroom teils us that a number of new finds and distributional changes have been incorporated, together with sundry classificatory changes and references to recent literature. Some of these have been incorporated in the text, others are given in the Addenda (pp. 911-913). The long introductory section includes four keys: a traditional one to families and certain genera; one based on the ‘Linnaean system’ in which the emphasis is on genera rather than families; a third for species which seldom flower, have inconspicuous flowers or are otherwise difficult to identify; and a fourth to trees and shrubs. There are brief notes, and a map, of the major phyto- geographical divisions of the Netherlands and a detailed account of the classification of Dutch plant communities. The descriptions of individual specimens include references to distribution, the relevant plant communities and literature references, which are all the more valuable for being fuller than those we are accustomed to. Protected species are indicated and most species are illustrated by small line-drawings. The nomenclature is, in the main, conventional and none the worse for that. Many will welcome the traditional treatment afforded to Anthemis and Matricaria and the demotion of Arenaria lepto- clados to A. serpyllifolia subsp. tenuior (Mert. & Koch) Archang. Clearly the nomenclature does not follow Flora Europaea, but current usage (where different) or alternative treatments, are often indicated. Thus, in the Addenda we are told of the generic splits of Lycopodium and of the generic treatment of Hepatica and Ficaria. An exception is the Euphrasia account, which I found impossible to square with that in Flora Europaea. Surprises include the treatment of Polystichum setiferum (diploid) as a subspecies of P. aculeatum (tetraploid), and the generic status accorded to Lemna polyrhiza (Spirodela) and Mentha pulegium (Pulegium). The Addenda are important for descrip- tions of the two subspecies of Myosotis palustris and of Veronica hederifolia, and also for the inclusion of Carex cespitosa L., first discovered in 1973. Without wishing to be patronising, this is very much a Flora for the amateur. There is virtually no experimental taxonomic information. No chromosome numbers are given (this will, no doubt, please some people), nor is there any reference to polyploidy, even though, for example, the tetra- ploid and hexaploid Polypodium vulgare are given specific status (surprisingly so in view of the treatment of Polystichum). There is no reference to apomixis under Hieracium or Taraxacum, but this is presumably because of the lack of all generic descriptions other than the appropriate lead of the family key. Hybrids are summarily dealt with, usually only by a note on their distribution. This is a pity. Personally, I would far rather have had more information on hybrids and other critical taxa at the expense of the disproportionate amount of space devoted to introduced and cultivated plants. The wild Primula hybrids may get short shrift but there are descriptions of seven cultivated taxa. Eight species of Rhododendron are included (surely a very arbitrary selection), as are 14 species belonging to five introduced genera of the Polemoniaceae. No doubt this is a very personal view and there will be those who will value this Flora for precisely these reasons. Certainly it is a most useful Flora, attractively produced and not too large to be fairly described as pocket-sized. G. HALLIDAY Cytotaxonomical Atlas of the Arctic Flora. A. & D. Love. Pp. xxiii+ 598, with 1 map. J. Kramer, Vaduz. 1975. Price DM200. As the authors say in their Introduction, the purpose of this Atlas is twofold. It provides for the first time both an exhaustive check-list of the Arctic flora and a list of all the chromosome numbers recorded for these taxa up to the autumn of 1975. It is their hope that by so doing they will both serve and advance the appearance of a comprehensive Flora of the Arctic. Their southern limit of the Arctic in general follows the tree-line. Most of Fennoscandia is therefore excluded but Iceland is included. 404 genera and 1,629 species are listed and chromosome numbers are given for about 90% of these. This is nearly twice as many species as are included in Circumpolar Arctic Flora (Polunin 1959) and results both from their narrower species concept and from their rather wider interpretation of the Arctic. The basic chromosome number for each genus is followed by a list of species synonyms. Under the individual species are given synonyms, chromosome number(s), geographical distribution and 264 BOOK REVIEWS a full list of the various chromosome counts. The bibliography includes about 2,300 references and accounts for nearly a sixth of the book. There is an index to genera, including synonyms. Any taxonomic work by the Loves is guaranteed to raise some eyebrows, and this Atlas is no exception. As they say (p. xviii), they are convinced ‘that a correctly and exactly defined species is characterized by a single chromosome number only’. Operating on this basis it is inevitable that they will cross swords with those who regard taxonomy as an essentially pragmatic exercise in which morphological differentiation is still of prime importance. As one would expect, there are taxonomic surprises on nearly every page. Many genera have been split, although many of the unfamiliar names are in fact quite old: Sedum rosea goes into Rhodiola L., Potentilla anserina into Argentina Lam., Rumex acetosella and R. acetosa into Acetosa Fourr., Cerastium cerastoides into Dichodon (Bartl.) Rchb. and Trichophorum cespitosum into Baeothyron A. Dietr. For the reasons behind these changes and for the validation of their nomenclatural innovations we must consult a paper in Botaniska Notiser (Vol. 128: 497-523 (1975)), which was published after the Ailas. Some of their changes will meet with general approval. Luzula frigida and Juncus balticus are treated as subspecies of L. multiflora and J. arcticus respectively, and Tripleurospermum is included in Matricaria. However, unsatisfactory as are many grass genera, I for one flinch from the con- sequences of the massive migration of Puccinellia into the diminutive and obscure arctic genus Phippsia, of most of Agropyron and all Roegneria into Elymus and the transference of £. arenarius into Leymus Hochst. Unfamiliar specific epithets abound—Agrostis capillaris for A. tenuis, A. vinealis for A. canina subsp. montana and Empetrum eamesii for E. hermaphroditum. — Clearly most of their changes are based on chromosome number. Many botanists regard Minuartia as a tiresome split of Arenaria (and many of its species as no less tiresome). Not so the Loves. To them it is a cytotaxonomic challenge, without a basic number in sight. The solution is simple. The 11 arctic species are allocated to six genera all with different basic numbers. This involves the creation of two new genera, Porsildia and Lidia, and the disappearance of Minuartia. A similar fate has befallen Sagina, S. intermedia, S. cespitosa and S. nodosa having been removed to Spergella Rchb. Gentiana verna and G. nivalis appear in Calathiana Delarbe (x = 7,8), whilst Gentianella tenella is in Comastoma (Wettst.) Toyokuni. Species splits predictably include the elevation of the diploid Asplenium trichomanes to A. melanocaulon, of the diploid Anthoxanthum odoratum to A. nipponicum, and of the two cytotypes of Eleocharis palustris to specific rank. Fortunately they have left certain taxa well alone, such as Wilhelmsia physodes (2n = 50—110), Saxifraga cernua (2n = 36—72), and most of the obligate and facultative apomictic grasses. It would be possible to continue in this vein at length; a rough count suggests that at least 20° of the native species in The flora of Greenland (Bocher et al. 1968) appear under different names in this Atlas. It is unfortunate that the idiosyncratic taxonomic approach of the authors will inevitably alienate many of the users of what is an extraordinarily thorough and valuable book. Any future Flora of the Arctic will certainly be heavily indebted to it, but its interest and importance are far wider than the title might suggest. Unfortunately it is exorbitantly expensive at DM200, parti- cularly since it appears to have been produced by an off-set litho process. G. HALLIDAY British Water Plants. 8S. Haslam, C. Sinker & P. Wolseley. Reprinted from Field Studies, 4: 243-351 (1975). Obtainable from Field Studies Council, Preston Montford, Montford Bridge, Shrewsbury, Salop. Price £1-20. Despite the remarkable amateur and professional interest in our native flora and the vast amount of literature on British systematic botany there are very few books entirely devoted to the identifi- cation of fresh-water aquatic plants. There has long been a need for an ecological approach to the practical study of plants in the aquatic environment, and this most recent work makes a preliminary fulfilment of this need. Following a brief introduction on the ecological niches occupied by aquatic vascular plants, on the scope and use of the book, and how to set about collecting and examining water plants, are three principal sections. The first is a key to approximately 120 British vascular aquatics; the second is a systematic account of their distribution and ecology; and the third is a series of 92 line drawings, — BOOK REVIEWS 265 showing the natural habit of the majority of the species discussed. There is also a small glossary of terms employed in the key, a scantily written account on the importance, or otherwise, of phyto- sociological methods for the description of aquatic plant communities, and a useful index. The dichotomous bracket-key attempts to have as much all-year-round coverage as possible by being based entirely on vegetative characters. It is divided into two main parts. The first is a small 5-couplet key to six artificial groups (A—F) for assemblages of unrelated taxa with similar features such as submerged plants, floating plants, etc. The second part comprises keys to these main groups with a secondary series of generic keys (G-I) for taxonomically difficult groups such as Ranunculus subgen. Batrachium and Potamogeton. These will have great appeal to the beginner and younger users since they are written in simple terms, with admirable thumb-nail illustrations after each couplet for correct interpretation of the characters. The key is well organized and workable. When there are difficulties owing to confusion with cryptograms and younger stages, appropriate notes are given. At times the illustrations come into their own when attempts at simplification lead to ambiguity, e.g. leaves grass-like v. leaves strap- like, and leaves Jris-like v. leaves not Jris-like. The policy for omitting hybrids, however, may lead to confusion when identifying some problematic Potamogetons. The second section, in systematic order following Flora of the British Isles, 2nd ed. (Clapharn, Tutin & Warburg 1962), is a compila- tion of ecological and geographical data providing useful comparative information about the water type, flow rates, nutrient status, soil types and habitats of the plants. Geographical data are taken from Atlas of the British flora and Flora Europaea. The last section, to our minds, is undoubtedly one of the most useful parts of the book. Here some 92, mostly half-size line-drawings of habit and vegetative features are presented as complementary to the other two sections. The drawings by Mrs Wolseley are excellent and reaily convey the information they are intended to give. It is not an easy section to use directly, however, since it is only cross-referenced to the distribution list and not to the dichotomous key. This present volume, like others in the Field Studies Council series, fulfils an obvious role for the practical understanding of our otherwise overworked flora. The fact that it is modestly priced and aimed at the layman and amateur worker is commendable. The lack of critical content may be wished for by the professional worker, although it is worth purchasing for the illustrations alone. C. J. Humpurigs & A. C. JERMY A Treatise on Limnology, Limnological Botany. G. Evelyn Hutchinson. Pp. x+ 660 with 164 figures. John Wiley & Sons, New York, London, Sydney and Toronto. 1976. Price £16-75. The third volume in this series on limnology is a concise and very readable review of the literature on biological and ecological work carried out on both macrophytes and the algal benthos of fresh- water ecosystems. The first chapter (Chap. 27 in the series), ambiguously called “The Lower Rooted Vegetation’, discusses Bryophyta and Charophyta; that on the Charophytes is extensive but the Bryophyte papers are less so. Chapter 28 discusses the nature and diversity of aquatic tracheo- phytes and could perhaps have been fuller in its references to pteridophyte papers on taxonomy which have a modicum of biological observation in them, e.g. the papers on African Marsilea by E. Launert. Chapter 29 (130 pp.) discusses the biological characteristics of the tracheophytes of inland waters under such headings as: The change from air to water; heterophylly and environ- mentally determined variations; photosynthesis and respiratory adaptations; and life history (in which full reference is made to C. D. Sculthorpe’s excellent book The Biology of Vascular Aquatic Plants). Under pollination, Hutchinson misses the opportunity of discussing the implications of under- water pollination in relation to recognition-mechanisms between pollen-grain and stigma surfaces seen in terrestrial plants, although, in fairness to the author, the work on this aspect by Pettitt et al. (Micron, 5: 377-405 and 7: 21-32) is too recent for inclusion. Pollination of aquatic plants is in need of study, and simple observations of when and how pollen is released are useful data to record by anyone with the patience to make them. Much of our knowledge in this respect stems from observations made at the end of last century. For instance, is Zannichellia apomictic ?—there is virtually 100% fertility even when female flowers are covered by mud. Najas marina in Britain is 266 BOOK REVIEWS most certainly apomictic, as (to the knowledge of the reviewer) no male flowers have been found here; yet this annual plant produces abundant seed—or did before pollution of the Norfolk Broads occurred. It is in Chapter 30 (144 pp.) that Professor Hutchinson excels, and here chemical compounds and elements of all kinds are discussed in relation to the environment and the metabolism of water plants in general. This chapter is as lucid and readable as the previous ones, and there is a mass of references and ideas to lead the researcher into useful areas. The penultimate chapter reviews zonations of macrophytes, depth distribution and phytosociology, taking examples from tundra to tropical lakes, and the last chapter describes benthic algal communities from the top of the eulittoral through to the bottom of the euphotic zone. There is an index of lakes, of genera and species of organisms and a general index; and the very comprehensive bibliography contains c 740 references. This volume is a mine of information on aquatic biology; and, whilst many users finding the lucid style both readable and informative will want volumes 1 and 2 as well, it is a book that can be used with only passing reference to the earlier works and well worth owning for its own sake. A. C. JERMY Hayley Wood. Its History and Ecology. Oliver Rackham and others. Pp. xv+221, with 26 photo- graphic plates, 33 text figures and end-paper maps. Cambridgeshire and Isle of Ely Naturalists’ Trust Ltd, Cambridge. 1975. Obtainable only from the publishers, 1 Brookside, Cambridge. Price £3-00. 7 Hayley Wood, 122 acres in area, has long been known to Cambridge naturalists as a good example of a wood on chalky Boulder Clay; they used to go there, keepers permitting, to see oxlips. Since 1962 it has been the property of the Cambridgeshire and Isle of Ely Naturalists’ Trust, who are to be congratulated on producing this monograph, which is of very much more than local interest. Like many ancient woods in England, Hayley Wood is of an ill-defined rounded shape and lies on the boundary of its parish; it takes its name from a lost village which, judging from the name, must have originated as a clearing in the forest. Details such as this contribute to the picture of pre-13th-century history. There is more documentary evidence for its later history than exists for most woods, because it belonged to the Bishops of Ely and was involved in a legal dispute between tenants and landlord early in the 17th century. Combining all this evidence with that from neigh- bouring woods and from field work, Dr Rackham builds up a fascinating and valuable account of the history of the wood, its past treatment, the sizes and destinations of its produce, and its position in the rural economy of the district. Coppicing ceased in 1922/3, and 40 years of neglect that followed must have impoverished the herbaceous flora: though Dr Rackham says that Jris pseudacorus is the best example of a species which has been exterminated by the upgrowth of old coppice, his study of the flora of the disused railway which skirts the wood (see especially pp. 150-152) suggests that some of the species which are now restricted to the railway verges must once have grown in the woodland rides. The period of neglect also allowed fallow deer to establish themselves, thus greatly adding to the Naturalists’ problems. There is not only a conflict between deer and vegetation (the deer have already exter- minated Alisma plantago-aquatica and, having a passion for oxlip inflorescences, threaten to exterminate the oxlips also, and they will destroy any young coppice shoots to which they have access), but also, I suspect (Dr Rackham is very discreet in what he says about this), a conflict between naturalists who want to keep deer and those who don’t. In any case it would probably now be impracticable to exterminate the deer, and new coppice coupes, which are essential if the old flora is to be perpetuated, must be securely fenced—we are told the number of man-hours that the work takes, and given other practical hints. All these problems of management are lucidly discussed. There is a wealth of other good things: excellent chapters on the ecology of the herbaceous communities, of the woody plants—the clones of elm and their spread, the dying oaks, the ages of ash stools (which prove to be the oldest living inhabitants), the ancient hedegerows adjoining the wood, and a specially commendable chapter by M. H. Martin and C. D. Pigott on the soils. All these matters are expounded in a scholarly yet eminently readable way, which would make the BOOK REVIEWS 267 book a good introduction to woodland ecology equally for a university student or for an intelligent lay naturalist. A common short-coming of the amateur naturalist is that he limits himself to recording the mere presence of species; rather less than a quarter of ‘Hayley Wood?’ is taken up by the species lists that result from this kind of observation. The remainder of the book demonstrates how rewarding other kinds of observation, mostly well within the ability of amateur naturalists, can be. I have commented elsewhere (in Forestry, 48: 216-218) on some other aspects of ‘Hayley Wood’, and have only space to add that I find it a very stimulating book; there must be many naturalists interested in woodland ecology who would derive great pleasure and inspiration from it, even if they have never seen the Cambridgeshire Boulder Clay. E. W. JONES Flore de France. Fascicule 2. M. Guinochet & R. de Vilmorin. Pp. 818, with 68 plates. Centre National de la Recherche Scientifique, Paris. 1975. Price Fr 90-00. However many the objectives of a descriptive Flora, its main purpose is surely to provide the reader with a means of identifying the plants of a delimited area. The question as to how this can best be done depends to a very great extent on the size of the area concerned, and on the total number of taxa involved. With a fiora as relatively small as that of the British Isles, it is possible to provide a liberal amount of illustration and text within the compass of a single portable volume. It is also not wholly unreasonable to assume that a reader, defeated by the imperfections of keys and descriptions, may nonetheless reach a satisfactory conclusion by the simple expedient of thumbing through the illustrations until he sees something which answers to the plant before him, a surmise which can be further substantiated by reference to the relevant text. With a flora as large as that of France, the problems are of an order which rules out such hopeful groping. The reader must submit to the discipline of a system. Moreover if the guide is to be of any service in the field, there must be stringent economies both in text and illustration, and essential identificatory data must be sifted out from the mass of interesting but unnecessary fact. The present volume, the second of a projected set of four fascicles (the first of which has already been reviewed in this journal) is, insofar as one can judge without actual use in the field, a thought- fully planned attempt at such a condensation. It consists of short, simple keys to genera and rather more elaborate ones to species and subspecies, the latter often skilfully combining both discrimina- tory and descriptive characters. With regard to the generic keys, I am pleased to see that the con- tributors have tried to employ the obvious rather than the erudite. Too often, in their anxiety to attain infallibility, taxonomists ignore the needs of the less informed, and distinguish genera by minute technicalities when other, much more obvious characters would, in the particular circum- stances, serve equally well. The illustrations, though perhaps not as numerous as one might have wished, have evidently been selected with care to elucidate particular difficulties, or to explain unusual terms employed in the accompanying text. They are, like the text, admirably crisp and clear. Supplementary information relating to habitat, distribution and time of flowering, must of necessity be expressed in such abbreviated form that it is not always very easy to interpret. The habitat data make extensive use of the formal language of phytosociology, a language still very imperfectly understood in this country, and one which we are markedly reluctant to learn. If in fact this nomenclature of plant societies does lend precision to what must otherwise be expressed in very general, vague and unsatisfactory terms, then perhaps it is time we gave it more serious, and more critical, attention. I am glad to see that Gallic commonsense rules out the use of ‘popular’ names except where such names are widely and traditionally used; at the same time, it is difficult to see on what logical principle the present selection has been made. R. D. MEIKLE The flora of Lincolnshire. Joan Gibbons. Pp. 351, with 15 black & white photographs and 11 figures. Lincolnshire Natural History Brochure No. 6. Lincolnshire Naturalists’ Union, Lincoln. 1975. Price £4-20. 268 BOOK REVIEWS Until the publication of Miss Gibbons’ book, Lincolnshire was the only English county without a | County Flora of any sort. 1975 therefore marks the end of an era in local Flora writing, and | Miss Gibbons is the last of a long line of first-Flora writers that can be traced back to the 17th century. Like most first Floras, this one shows marked symptoms of condensation, and one is left with the impression that the author has a great deal of detailed information in the files or card indexes for which there was no room in the book. Lincolnshire is, however, the second largest English county, over three times the area of Leicestershire, whose first comprehensive Flora contained proportionally about as much information as that by Miss Gibbons, and whose second contained nearly 1,000 pages of text, much of it in very small type. Miss Gibbons’ introductory chapters, some of which are contributed by other authors, contain much information of value on topography, geology, meteorology, habitat types and botanical history. Inevitably, because of the size of the county, the maps are on a very small scale and the amount of topographical detail is limited. Some critics may regret the absence of a Gazetteer section; for a county as large as this it would inevitably have to be very incomplete, and it was probably wise not to attempt one. The section dealing with accounts of individual species occupies about 200 of the 350 pages of the book, and appendices concerned with aliens, extinctions, statistics and other related matter occupy a further 20. In this space the amount that can be said about each species is limited; in fact the strictest economy is observed over the quotation of individual records, distribution within the county, and habitat preferences. As regards distribution, Miss Gibbons prefers Woodruffe— Peacock’s Botanical Divisions to any kind of grid system, or to recording by parishes. It is worth noting, however, that The Atlas of the British flora already provides reasonably up-to-date maps of distribution in the county on a 10-kilometre basis, and to record in Lincolnshire on a ‘tetrad’ basis would be a monstrous task, requiring getting on for 2,000 tetrads and a total of 500,000 records. It is perhaps a pity that Woodruffe—Peacock chose to divide the county into only 18 divi- sions, since each of the presence or absence records in this Flora refers to presence or absence in an area almost as large as Middlesex. One senses at once the dilemma which faced the author in deciding how much of the available information to include. Compared with the pages of D. H. Kent’s The Historical Flora of Middlesex, the pages of this Flora are terribly empty, but to raise the amount of information on each species to the next practicable point on the scale would not only overfill the pages but also increase their number to a significant degree. It has been suggested that the chapter entitled The Lincolnshire Naturalists’ Union, consisting largely of a list of LNU Field Meetings from 1893 to 1973, is irrelevant and could well have been omitted, but the seven pages saved would not have provided the additional space needed to enlarge the accounts of species. My conclusion is that this is an excellent introduction to the study of the Lincolnshire flora, and that it in no way presents a discouragement to further study. Lincolnshire botanists are now left with the fascinating problem of whether to continue working along the lines so ably initiated by Miss Gibbons, perhaps with the parish as the basic recording unit, aiming ultimately at a Flora on the scale of Horwood & Gainsborough’s of Leicestershire and Rutland, or to go in for grid recording using a smaller grid than the 10-kilometre one already in use. K. G. MESSENGER Bedfordshire Plant Atlas. J. G. Dony. Pp. 132, with 800 plant distribution maps, 18 other maps and 2 perspex overlays. Borough of Luton Museum & Art Gallery, Luton, Beds. 1976. Price £3-00. Dr Dony has broken the records for county Flora writing; he has not only produced no less than three, but this Atlas is the second by the same author for the same area, and this must be the first such achievement. The new work is in many ways different from the older one (Flora of Bedford- shire, 1953). We now have 800 clearly printed maps of plant distributions, with notes on the remainder of the flora, including aliens; and an amount of information similar to that in the Flora is now presented much more compactly. The Atlas has a short but highly informative introduction, covering survey methods, geology, habitats and recent changes in the flora. Transparent overlays for the maps are included, giving an easy way to compare with factors such as river systems or built-up areas. On the other hand, the original Flora was expanded with a botanical history, habitat BOOK REVIEWS 269 surveys, records of non-vascular plants and a bibliography. Those who may wish to compare distributions of species of then as against now will not be able to do so very readily, on account of the very different methods of presentation. In making comparisons we in fact see the impact of Dony’s own thinking and its influence on county Flora writing, including the use of the tetrad mapping-unit (a square of 2 x 2 km) pioneered by him. The At/as compares very well for cost with the Flora (42s.), even without allowing for inflation. R. J. PANKHURST Plant and Animal Habitats in Town and Country. G. E. Simmons. Pp. 96. Blandford Press, Poole, Dorset. 1976. Price not stated. This book is one of a series with the overall title Approaches to Environmental Studies. Many aspects of the environment are studied; for example the weather, roads, railways, villages, churches and religions, police and fire services are but a few of the topics covered in 24 books. These are grouped in four sections, and to each section there is a teacher’s guide book as well as programmed texts and ancillary material. This particular book (No. 23) is one of the section that includes topics like pollution and parks and open spaces and is therefore more biological than some of the other sections. Plant and Animal Habitats is a concise little book defining briefly the meaning of the term ‘habitat’ and mentioning some of the more frequent plants and animals of habitats like gardens, streets, walls, hedgerows, commons, ponds and waste grounds. It is full of suggestions for observation and the keeping of records which, if carried out successfully, would be most rewarding. To be used really successfully, however, the pupil will require a fair measure of assistance. Thus instructions like ‘See how many woodlice you can find’ or ‘Make recordings of the insects that visit the umbels of Hogweed’ are difficult even if identification is carried only as far as the order, for many pupils find difficulty in distinguishing the main groups of insects. However, provided help is available, this is a stimulating little book. It is clearly set out and well illustrated, and there is a short biblio- graphy. C. T. PRIME Palaeobiology of Angiosperm Origins. N. F. Hughes. Pp. vii+ 242, with 73 half-tone and line illustra- tions. Cambridge University Press, Cambridge. 1976. Price £7-80. The voluminous literature on primitive flowering plants has been written largely by students of ‘recent’ angiosperms. An account of them from the point of view of a palaeobotanist is therefore all the more welcome. Alas! The point of view turns out to be a very individual one, and Dr Hughes makes few attempts to build bridges between the territories of palaeontology and neontology. Instead, he points out the dangers of ‘back-classification’, i.e. of extending the classification of recent organisms to accommodate fossils, and proposes that recent and fossil flowering plants be classified—and even named—in quite separate ways, e.g. by ‘development of a separate parallel data-handling code for palaeontology to decrease reliance on the inappropriate International Code of Botanical Nomenclature’. These generally negative conclusions are disappointing for the neontologist who takes up this book hoping to find concrete data that will either support one of the current theories of angio- sperm origin or suggest a new theory. Instead, after providing useful introductory sections on fossils and their interpretation, Dr Hughes examines the relevant evidence from Jurassic and Cre- taceous plant fossils only to conclude that ‘none of these pre-Cretaceous fossils represents an angiosperm’, and that one should ‘set aside all theoretical phylogeny until a genuine surplus of Cretaceous fossil information has been accumulated and has been stored under neutral descriptors’. He then occupies a mere 12 pages in examining current theories in the light of the fossil evidence as he sees it; but his conclusions are again depressingly negative. Even theories partly based on fossil evidence, such as the Gonophyll Theory, do not appear to excite his interest. I am glad to have read this book and would urge anyone interested in angiosperm origins and classification to read it. It will fili in the fossil background to these topics well; but it will also provide a cold douche to dampen evolutionary ardours. N. K. B. Rosson 270 BOOK REVIEWS The Biology and Chemistry of the Cruciferae. Edited by J. G. Vaughan, A. J. MacLeod & B. M.G. | Jones. Pp. xvi+ 355. Academic Press, London, New York and San Francisco. 1976. Price £7-80. This book presents the papers read at a Conference on the Biology of the Cruciferae arranged | jointly by the Linnean Society of London and the Phytochemical Society in London in January 1974. The addition of the word Chemistry to the book title reflects the predominance there of this aspect of biology; 5 of the 11 main chapters are devoted to chemical aspects—myrosinases (mustard- oil-splitting enzymes), glucosinolates (sulphur-containing compounds giving the Cruciferae their | characteristic pungent taste), lipids (mostly fatty acids), seed proteins and volatile flavour com- pounds. By a simple process of elimination, one can deduce that the non-chemical aspects of Biology in the present context include taxonomy, anatomy, cytogenetics, crop breeding and population genetics. The survey of systematic and phytogeography by I. C. Hedge makes interesting reading; but, as the author admits, it is still not possible to present a wholly satisfactory classification of this extremely natural family. D. J. Harberd’s survey of the cytotaxonomy of Brassica and related genera is a valuable summary of the accumulated information on the economically most important group of Crucifers, and it is nicely complemented by P. Crisp’s chapter on the breeding of Cruci- ferous crops, which naturally concentrates on the same group. ‘Seed Studies in the Cruciferae’ by J. G. Vaughan et al. covers both structural and chemical aspects and, contrary to what is often suggested by cytotaxonomists, indicates that Brassica and Sinapis are indeed separate genera. The remaining two chapters, on trichomes in Lesquerella (R. C. Rollins & E. C. Banerjee) and natural variation in Arabidopsis thaliana (M. J. Lawrence), are of a different nature, for the emphasis is on trichomes and natural variation respectively, and it is largely coincidental that members of the Cruciferae are the subject matter. Although there will be some who regret the use of unjustified photolitho type for a book of this — nature, the quality of presentation from both the editorial and publication points of view is high. However, the very unhelpful lack of titles of references, albeit now customary in many works of this sort, is to be much regretted, and in the present case is particularly frustrating because of the large amount of space wasted in the Bibliographies due to the format adopted. Thus the over-riding impression left by this book is that it contains several very useful chapters, such that a professional taxonomist will want to have it on his shelves, but that, like so many other conference reports, the contents do not hang together very well. This is largely because the organizers of conferences are able to include only those topics for which active research workers are available, and this problem is particularly acute when the conference theme is a single family. The book compares well with the earlier similar survey of the Umbelliferae (1971) and one awaits with interest the report of the conference on the Solanaceae held in 1976. C. A. STACE Flowering Tropical Climbers. G. Herklots. Pp. 194, with 260 figures and 16 colour plates. Dawson, Science History Publications, Folkestone. 1976. Price £17-50. This beautifully printed and boxed book is a delight to possess. I recognized at once all 11 climbers I grew in my garden in Malaya. The 270 line-drawings and 16 full-page colour plates are superb. All have been made by the author during his long life in the tropics. Date and locality are given so we can follow his peregrinations—Hong Kong, Trinidad and Nepal, where he resided, figure frequently. The slightly eclectic text is woven around the pictures; botanical detail (including precise notes on flower colour), habitat and range are included. Dr Herklots has not been well served by his editor and the text is marred by minor inconsistencies in citation and format, and the index, regrettably, is restricted to the scientific names. This should not deter one, for this is supremely an example of elegant book production, sublimely illustrated. T. C. WHITMORE Watsonia, 11, 271-279 (1977). 271 Reports ANNUAL GENERAL MEETING, 22nd MAY 1976 The Annual General Meeting of the Society was held in the Jodrell Laboratory, Royal Botanic Gardens, Kew, on Saturday, 22nd May 1976 at 12 noon, with 107 members present. Mr E. L. Swann (President), in the Chair, opened the meeting by expressing regret at the death in January of Mr J. E. Lousley, who had been a leading member of the Society and one of the foremost amateur British botanists of his time. The minutes of the last Annual General Meeting, as published in Watsonia, 11: 93-94 (1976), were passed as correct. REPORT OF COUNCIL The adoption of the Report for the year 1975, as circulated to members, was proposed by Dr C. P. Petch, seconded by Mr P. J. O. Trist, and carried unanimously. TREASURER’S REPORT AND ACCOUNTS The Treasurer pointed out that the Accounts showed the £2,000 deficit predicted by him in his 1974 Report. This was partly compensated for by assets in the form of posters and conference reports as yet unsold but, hopefully, soon to be converted to cash. Dr I. K. Ferguson commented on the 47% increase in the cost of printing and distribution of Watsonia. The Treasurer replied that this included heavy postal increases, and reported that the Editors and the Publications Committee had carried out a thorough investigation of possible economies in the publication costs of the Journals. Implementation of resulting economies should be apparent in the future but present figures were slightly inflated to allow for bills not yet received. The adoption of the Treasurer’s Report, together with the Accounts for the year 1975, was then proposed by Mrs B. H. S. Russell, seconded by Mr E. C. Wallace, and carried unanimously. The Treasurer asked that a vote of thanks to the Honorary Auditors should be recorded, and reported that he had presented them with a Raymond Piper Orchid print from the Society. The President had written thanking the Auditors and their reply expressing goodwill was read. On Dr W. T. Stearn’s proposal it was agreed to send to the Honorary Auditors the appreciation of this meeting. ELECTION OF OFFICERS The election of Mrs B. H. S. Russell as Vice-President was proposed by Mr E. Milne-Redhead, seconded by Dr J. G. Dony and carried unanimously. The Chairman thanked Mr J. C. Gardiner, retiring Vice-President, for his valuable work for the Society during his long term of office as Treasurer. The following officers were then proposed for re-election: Mrs M. Briggs Honorary General Secretary; Mr M. Walpole Honorary Treasurer; Dr G. Halliday, Dr N. K. B. Robson and Dr C. A. Stace Honorary Editors; Mrs R. Hamilton Honorary Membership Secretary; all proposed by Mrs J. G. Dony, seconded by Miss M. McC. Webster, carried unanimously and thanked from the Chair for the great amount of work achieved for the Society. The election of Mrs J. M. Mullin as Honorary Meetings Secretary, proposed by Mr E. J. Clement, seconded by Dr I. K. Ferguson; and the election of Miss L. Farrell as Honorary Field Secretary, proposed by Dr F. H. Perring, seconded by Mr R. J. Pankhurst, were carried unanimously. Welcoming these new officers, the President warmly thanked Mrs G. Beckett, retiring Field Secretary, who had been successively Programme and Meetings Secretary since 1967. A vote of thanks to Mr D. Langley, acting Meetings Secretary for part of the past year, was also recorded. ELECTION OF HONORARY MEMBERS Council had recommended the election of Dr S. M. Walters and Mr E. F. Greenwood. Dr Walters, an active member since 1944 and past President of the Society, had also served on many B.S.B.I. 272 REPORTS Committees including the Maps Committee, which had launched the survey for the Atlas of the British flora (1962). Mr Greenwood had been an Editor of the Society’s Journals for nine years (the Editor of Proceedings then an Editor of Watsonia) during which time considerable changes in format and of the business arrangements came under his supervision. The President from the Chair proposed election of these two worthy members and this was carried with prolonged applause. ELECTION OF HONORARY AUDITORS The election of Messrs Thornton Baker & Co. was carried unanimously. CHANGES OF RULE The following alterations to the Rules were passed without dissent: (a) Rule 9 Election of Vice-Presidents: *... to serve a term of four years and not then be eligible for immediate re-election’ (b) Rule 10 Length of service of Council Members: ‘A Council Member lost by death or resignation may be replaced for the remainder of the seniority by a co-opted member who shall then be eligible for election’. (c) Rule 30 Resignation: ‘. . . written notice to the Hon. General Secretary or the Hon. Treasurer before the last day of December... .’ (d) Rule 6 ‘The Officers of the Society shall be . . . the Honorary Editor or Editors of Watsonia’. ANY OTHER BUSINESS Mr K. H. Tough, commenting on the content of Watsonia, queried the possibility of a wider intake of articles. The Treasurer supported this request. Dr N. K. B. Robson, an Editor, replied that the content of the Journal depended on the subject material offered and that the Editors were willing to publish any relevant article offered if of appropriate quality. Mr E. Milne-Redhead suggested that some members might feel that they would not dare to write a paper attempting to meet the required standard. Mr S. A. Renvoize asked if the Editors could commission papers. The Secretary reminded members that the Editors had repeatedly appealed to members to submit Short Notes for Watsonia, and suggested that any members with a note or paper of scientific interest (and of interest to the Society) should offer this to the Hon. Receiving Editor, who would ensure that these would receive sympathetic consideration. Mr E. J. Clement asked if there was any intention to index B.S.B.I. News. The Secretary replied that this had not been considered to date. An index would add to the cost of B.S.B.J. News and with the recent stress on cutting costs of publications any decision on this would need careful thought. The Treasurer said that if B.S.B.J. News were to be indexed a volunteer for the work would be required. This point would be passed to the Publications Committee for consideration. The meeting closed at 12.36 and, at the end of the Programme for the day, Dr C. T. Prime, in the Chair, thanked the Director for his welcome and for his permission to hold the meeting at Kew in the very pleasant surroundings of the Jodrell Laboratory. Dr Prime also thanked Dr R. K. Brummitt for organizing local arrangements, and all the speakers, with special thanks to Mr C. Grey-Wilson who had come at short notice with excellent slides of Plants of Nepal. On the following day, Sunday 23rd May 1976, an excursion was made by 45 members to the Royal Botanic Gardens, Kew, at Wakehurst Place, Sussex, where a guided tour was given by Mr A. D. Schilling, Deputy Curator. M. BRIGGS FIELD MEETINGS 1975 ENGLAND STONE-—GREENHITHE, KENT. 27TH JULY The party of 17 met near Greenhithe Station and, under the leadership of Mr J. R. Palmer, spent some six hours under incredibly hot conditions exploring old lime-pits, the Thames reaches and derelict industrial land. En route to the first lime-pit near Greenhithe, we saw Petroselinum crispum and Silene italica REPORTS 273 growing in a hedgerow, while the densely wooded slopes surrounding the pit yielded gooseberry, a new tetrad record, and about thirty plants of Orobanche hederae in its only current Kent station. The pit was being filled with industrial waste and produced aliens such as Bromus madritensis, Nicotiana alata, Artemisia absinthium and Lycium barbatum. A large number of Ailanthus altissima seedlings were found as were also Lathyrus sylvestris, Sorghum halepense—in at least its second season judging from the impressive root system, and Hemerocallis fulva. A few bird-seed aliens were growing in the pit despite the early season for them—Setaria viridis, Linum usitatissimum and, of course, Phalaris canariensis. The second phase of the meeting involved a long hot walk starting from Greenhithe Station, where a hedge of Lonicera japonica caught the eye, and passing through an Industrial Estate where patches of dried-off Vicia dasycarpa, and later V. bithynica were pointed out. Along the track of a disused railway were Lotus tenuis and Carex divisa, while a nearby ditch contained Berula erecta. Lunch was eaten by a small beach near several areas of Parapholis strigosa, P. incurva and Agropyron pungens x A. repens. Walking towards Stone marshes, we inspected a good display of Verbascum phlomoides x V. thapsus. On the edge of the marshes were Calamagrostis epigejos and, to the delight of the grass enthusiasts, Puccinellia pseudodistans and P. fasciculata. The party then walked over the railway to a vast disused quarry which was about to befilled in with some of London’s refuse. We saw, probably for the last time in this pit, Nardurus maritimus in the open, while in a woodland area were scores of Epipactis phyllanthes, Listera ovata and several Hieracia. A clearing in the wood was carpeted with Oenothera parvifiora and, in the deepest part, our leader unerringly located numerous plants of Pyrola rotundifolia, all shortly to be buried by rubbish. After a discussion on conservation (the Pyrola site was man-made), the day ended with a short tour of Stone churchyard to view Salvia horminoides in the turf between the gravestones. C. G. HANSON ATTINGHAM PARK, SALOP. 21ST—24TH AUGUST This meeting gathered in the sad knowledge that it would be the last of a series begun in 1968, for in November 1975 Attingham Park would close as an Adult College and the wonderful hos- pitality botanists had enjoyed annually would no longer be available. This is particularly frustrating at a time when interest in the flora of Salop continues to grow and the demand for places on a meeting of this kind far outweighs the accommodation. The aim of the meeting, as in previous years, was to combine instruction in the identification of plants with work on a new Flora of Shropshire. To this end lectures were given each evening, whilst, during the day, visits were made to under-recorded parts of the county where particular 1 km squares were selected for detailed study. The morning of Friday 22nd was spent in the Hinkshay Field Studies Area, GR 33/69.07, on the edge of Telford New Town, a mixture of pools and woodland, the relics of an early nineteenth- century mining area. This was a good introduction for beginners since there was an abundance of common plants including a number of species pairs needing careful separation, e.g. Betula pendula and B. pubescens, and Rumex conglomeratus and R. sanguineus. During the afternoon we climbed the south slopes of the Wrekin near The Spout. Here a small stream cuts through damp alder/elm woodland which, though much affected by recent forestry, provided a rich flora of woodland ferns and grasses, notably Blechnum spicant, Dryopteris pseudo- mas, Festuca gigantea and Milium effusum, with, occasionally, such herbs as Lamiastrum galeobdolon and Valeriana dioica. One of the early decisions of the ‘Shropshire Flora Committee’ was that for map-making it would be best not to confine our work to the exact boundaries of v.c. 40 (Salop) but draw our limits instead at the edge of 10 km squares. So Saturday morning was spent in the Hereford (v.c. 36) part of 32/4.7 in the woods of Downton Castle Gorge, by kind permission of the owner. Here the River Teme has cut a deep gorge through Silurian rocks and the sides are clothed in mixed deci- duous woodland. Many new records were added to the 10 km square including a few specimens of Tilia platyphyllos, in full fruit, which could well be native in this locality. Scrophularia umbrosa was found in quantity by the river bank; it is a common plant in southern Salop and may well be widespread in v.c. 36 also. On the way back to Attingham, the whole party descended on the village of Shelderton. Splitting 974 REPORTS into about 5 groups we recorded about 170 species in an hour, 16 of which were new to 32/4.7, but this time well inside v.c. 40. Notable finds here included Agrostis gigantea, Pentaglottis semper- virens, Cicerbita macrophylla (a large patch established in the centre of the village), and Crepis capillaris var. glandulosa, always a puzzle for the uninitiated. For the last day of this course, and all the Attingham courses, we chose to return once again to the Llanymynech area on the Welsh border where Miss Doris Pugh led us to yet another un- explored corner—this time to Llyn Rhuddwyn (33/23.28), a small lake which lies in a hollow on a ridge at about 1,200 ft. With little water left after the long dry summer, the marshy margins and wooded surrounds were easily accessible and the following local Salop species were found: Carex vesicaria, Littorella unifiora, Myosotis secunda and Peplis portula. Lunch and the final farewells were taken in one of those innumerable Carboniferous limestone quarries which pepper this area. This one at Craig-llwyn (33/23.27) had the usual range of lime- stone grassland herbs and grasses but was more noteworthy for its aliens than its natives—notably a fine and apparently indestructible patch of Senecio tanguticus. A final roll-call disclosed what a mixed meeting this had been. Out of 20 participants 9 were members of the B.S.B.I., 9 of the British Naturalists’ Association and 5 of the W.F.S., yet we had worked happily and harmoniously together without being aware of these differences, which was just as well as some belonged to all three organizations! F. H. PERRING WALES WYE VALLEY, GWENT. 7TH JUNE 28 members and friends gathered together at the Wyndcliffe, Gwent from as far apart as Aberyst- wyth, Yorkshire, Scotland and Wiltshire. Five small areas of limestone woodland were visited as there are no large woodlands left in Gwent that have not been radically changed, since the war, by the Forestry Commission. The Wyndcliffe, with the panoramic viewpoint known as the ‘Eagles’ Nest’, lies in a Forestry Commission Reserve and it was here that members saw a remnant coppiced wood with ancient yews. Paris quadrifolia was not at its best and was half hidden by Mercurialis perennis. Near the top of the cliffs, shade had prevented Rudia peregrina from flowering. Arabis hirsuta was passed, clinging to a precipitous rock face, as the party descended the ‘365 steps’, at the bottom of which Pyrola minor was found among the beech leaves. The clearing of an unsafe rock face had prepared new ground for Atropa bella-donna and Euphorbia stricta to colonize, and Bromus lepidus was among the grasses on the roadside verge. Carex remota and C. divulsa were compared as they grew close to one another on the path to the car-park. The day was, by now, clear and hot and the cathedral-like clearing at Blackcliff must remain in the memory of those present for a long time. The atmosphere was cool and the light, beech-leaf- filtered green, fell on the fern-covered tumbled rocks, amid which were Chrysosplenium alterni- folium and Cardamine impatiens. Among other plants found were Epipactis helleborine, Cephal- anthera longifolia—the sole survivor of a colony wiped-out under stone dumped from road- widening, Neottia nidus-avis, Convallaria majalis, Carex digitata, and, at its most southern station in Britain, Melica nutans. In Cockshoot Wood a stop was made to see Dryopteris carthusiana and the leaves of Colchicum autumnale before crossing into Bishops’ Barnets Wood, where Carex pallescens, C. pilulifera, Melica nutans and Aquilegia vulgaris vied for space. A brief stop was made in the Coombe Valley to look at Symphytum x uplandicum. The final call was at Hardwick Plantation, where the Forestry Commission had set aside, at the request of Gwent Trust for Nature Conservation, a hectare or so for the preservation of some of the 360 plants recorded there and which might disappear if under conifers, like the rest of the wood. The party compared Platanthera bifolia and P. chlorantha and also saw Astragalus glycyphyllos, Aquilegia vulgaris, Trifolium medium, Picris hieracioides and P. echioides. T. G. EVANS REPORTS 275 SCOTLAND RIVER TAY, MID AND EAST PERTH. 25TH MAY This meeting was held jointly with the Andersonian Naturalists of Glasgow and the Perthshire Society of Natural Science, with the purpose of re-examining the flora of river-side shingle beds, previously visited in June 1969 (Watsonia, 8: 189-190). The party of 21 members and friends left Kinclaven Bridge at mid-day and travelled along the Murthly road to the farm of Kercock. In 1969 the shingle bed below the steading had been domi- nated by plants of broom and lupin; today, and now in full flower, only the broom was much in evidence. Selective nibbling by an increasing rabbit population was suggested as a possible cause of the lupin’s decline. The most noteworthy species in the ground flora were Aquilegia vulgaris, Thalictrum minus, Teesdalia nudicaulis, Silene maritima, Cerastium arvense, Geranium pratense, Ononis repens, Saxifraga granulata, Primula veris, Veronica filiformis, Galium boreale and Cirsium heterophyllum. At the eastern end, in silt overlying the shingle and under alder and willow, several colonies of Tolmiea menziesii were in full flower. Continuing westward, we crossed the river by the bridge near Caputh and the farm of Braecock was reached along the private road from the village. Beyond the farm lies The Inch, a shingle bank about half a mile in length, between the river and a backwater. Several of the species already noted at Kercock were seen again here and among those not previously listed were Lepidium heterophyllum, Arabis hirsuta, Geranium sylvaticum, Origanum vulgare and Carex vesicaria. Cattle now have access to this site, which may account for the decline of the lupin here; there was, however, a flourishing colony of Lupinus nootkatensis just across the river. This was reached by re-crossing Caputh Bridge and walking eastward along the wooded bank. In passing it was noted that the alien Salvia glutinosa, found here by Mr A. W. Robson in 1959, was still flourishing. Among the lupins, Trollius europaeus, Rubus saxatilis and Festuca arundinacea completed the total of noteworthy plants seen on that fine, sunny day. R. MACKECHNIE KIRKCUDBRIGHT. 1ST JUNE On the day following the Society’s Annual General Meeting at Dumfries, 37 members met at Dumfries Station and travelled into Kirkcudbright by chartered bus. The first stop on the road to Dalbeattie was by a steep road-bank near Lochanhead, where Dr Richards started by teaching the uninitiated how to identify various dandelions. Taraxacum cyanolepis, T. euryphyllum, T. raun- kiaerii and T. sellandii were examined. We also saw a Pyrola, identified by those present as P. minor. A specimen from this location had been identified as P. media by the Royal Botanic Garden, Edinburgh, so further investigation is needed to see if both are present. Lycopodium clavatum also occurred on this bank. The next stop was to examine ferns, by a ravine just east of Kirkcudbright Town. Phyllitis, Polystichum aculeatum, Asplenium trichomanes subsp. quadrivalens, Polypodium vulgare,P. interjectum and the hybrid P. x shivasii were all present, with a series of hybrids between Geum rivale and G. urbanum. Dandelions here included T. unguilobum, T. nordstedtii and T. croceiflorum. The bus then drove around Kirkcudbright Town, so that the Georgian architecture could be admired, and Erinus alpinus was noted on the walls of McLellan Castle. Our main destination was the shore near Borgue, to see an example of the coastal flora. Although Scilla verna was just going over, Astragalus danicus was in full flower, growing with Helianthemum nummularium, Armeriaand Anthyllis, making a very colourful spectacle. In the wetter places Schoenus nigricans was abundant with Eleocharis quinqueflora, and here Taraxacum palustre was noted. The nearest stations of this distinctive and rare dandelion are in Anglesey. The Borgue locality was discovered in 1973 and consists of over 100 plants, making this the largest colony known in the British Isles. Other dandelions included T. faeroense, T. naevosiforme, T. norstedtii, T. lacisto- phyllum, T. unguilobum and T. raunkiaerii. Viola canina was found on the cliffs, the first record for v.c. 73 since 1930. On the return journey, a stop was made on the merse (salt-marsh) near Southwick. The plant of most interest was a very large colony of Hierachloe odorata in full flower. A sedge was found that had the company guessing from C. diandra to C. divisa, but Mr Jermy decided it must be C. disticha, the first suggestion. Dandelions are rarely seen on salt-marshes, but here was a large 276 REPORTS colony of T. spectabile, with a few individuals of 7. maculigerum. In nearby woods grew Polypodium australe and a weil-naturalized group of Leycesteria formosa. T. unguilobum and T. sublaciniosum were here, together with a remarkable dandelion which appears to be undescribed. O. M. STEWART & A. J. RICHARDS DUMFRIES. 2ND-6TH JUNE On 2nd June the group of six and one guest set out to visit a wooded glen opposite Beattock. On the way we stopped at two places to show our visitors what we thought would interest them. Looking over St Ann’s Bridge into a deep gorge, we saw a fine show of Valeriana pyrenaica at the bottom. Both Beech and Oak ferns were visible. The next stop was at Lochwood, where there are peculiar old oaks. Spring flowers were still abundant, with Stellaria nemorum and Equisetum sylvaticum by the roadside. The Beattock glen had been chosen because herbarium specimens in the Dumfries Burgh Museum suggested that botanical forays used to be made to this beauty spot last century. We were rewarded by making the first post-1930 records of Cardamine amara, Carex laevigata, Thelypteris phegopteris, Gymnocarpium dryopteris, Dryopteris dilatata, Lycopodium clavatum, Myosotis secunda and Circaea intermedia. Anemone nemorosa and Epilobium brunnescens were added as first records for the 10 km square. On Tuesday, 3rd June we visited the eastern side of the county at Carronbridge, Nithside and Wanlockhead. As usual we were marking cards for the 10 km squares. By the riversides the more interesting additions were Carex hirta, Scirpus sylvaticus—not at that time in flower, Chryso- splenium alternifolium, and Geum x intermedium. The time-consuming run to the old lead-mining town of Wanlockhead left us with less than an hour for botanizing. Opening the car doors on to Botrychium lunaria and Viola tricolor raised our hopes, but the finds were few and, with one exception, ordinary. Miss E. Young returned from a look round the churchyard with a specimen of Draba muralis. This was the only new v.c. 72 record of the meeting. On Wednesday, 4th June we visited the part of the Hightae Local Reserve which is on old peat cuttings, where it was exciting to find Andromeda polifolia, Vaccinium oxycoccos and Drosera rotundifolia, enjoying an apparently safe existence in comparison with the other wetlands around Lochmaben. In the afternoon we went to the disused railway line, now grazed by cattle. Here we saw such characteristic plants as Chaenorhinum minus, Linum catharticum, Arenaria serpyllifolia, Anthyllis vulneraria and Aira caryophyllea. Cerastium arvense has spread on to the line from a dry bank nearby. Viola canina, determined by Professor D. H. Valentine, restores a record for the vice-county made in this same square before 1930. On the last morning a visit to the strand vegetation and salt-marsh at Powfoot was begun in the rain, but we added 11 records for the square, including Cochlearia anglica. On the salt-marsh, Blysmus rufus, Eleocharis uniglumis and Puccinellia maritima made up much of the sward. Ruppia maritima, determined by Mr J. E. Dandy and looking like green hair, filled a pool. In the afternoon we ended the meeting by looking at the invasive flora of a dried-up loop of the River Milk at Castlemilk, and adding 9 new records for the square. It was a worthwhile four days of good companionship enjoyed while adding these records to the squares visited. J. D.S. & M. E. R. MARTIN BALLANTRAE, AYR. 7TH—-8TH JUNE This poorly attended meeting, originally planned for two days, became in the event a one-day meeting, due to the early departure of those participating. However, in the short time available we were able to examine the stretch of cliff and shore between Portandea and Currarie Port lying some five miles south of Ballantrae. The wooded glen near Glendrisaig House was also explored. The most noteworthy plants observed on the cliffs were Asplenium marinum, Juniperus communis, Geranium sanguineum, Potentilla anglica, Sedum rosea, Carlina vulgaris, Antennaria dioica and Scilla verna; while in the glen we noted Polystichum setiferum, Equisetum telmateia, Hypericum androsaemum, Vicia sylvatica, Sanicula europaea and Allium ursinum-—the last covering the ground over considerable areas to the exclusion of all other vegetation. A brief visit was also paid REPORTS 277 to the shingle bank at the mouth of the Stinchar where Mertensia maritima was in fine condition, and an after-dinner visit to the coast near Lendalfoot produced Trifolium striatum. A total of 200 species was recorded for the 10 km square 25/0.7, of which 12 were additional to those recorded in the Aflas. A. McG. STIRLING KILLIN, MID PERTH. 22ND JUNE The Andersonian Naturalists of Glasgow and the Perthshire Society of Natural Science (Botanical Section) joined forces with the C.S.S.F. in a very successful ‘assault’ on the 2,900 ft mountain, which stands sentinel over Loch Tay and Glen Lochay where these two valleys meet. Messrs A. W. Robson and A. M. Stirling led 22 members and friends who met at the Ben Lawers car- park and, by kind permission of the landowners, were able to transport the party to a point c 2,200 ft up, almost at the base of the floristically rich cliff-faces. It was a day of brilliant sunshine and so the party concentrated on the arctic-alpine and associated flora of the higher parts. The party agreed to fan out over the whole hill and to assist in compiling a list of species on 10 km cards. Collating this effort produced a total of 166 species. The following rarities provide an idea of the interest this mountain holds and suggest that there are others awaiting discovery: Potentilla crantzii, Dryas octopetala, Salix lapponum, S. reticulata, Armeria maritima, Saussurea alpina, Juncus alpinoarticulatus, Eleocharis quinqueflora, Carex capillaris, C. saxatilis, C. vaginata, C. atrata, viviparous Poa alpina, and Sesleria caerulea. A. W. RoBson COWAL, MAIN ARGYLL. 12TH—19TH JULY The object of this meeting was to look at underworked areas in Cowal. Seven 10 km squares were visited and, although nothing unexpected was found, a good many species were added to the master cards. 12th July: We met at Loch Striven head and went by a forestry road up the glen behind to a fair height. Nothing unusual appeared to be threatened by the forestry operations. The long drought had not helped the flora and the most noticeable feature were the white patches of Eriophorum vaginatum. In the afternoon we visited the salt-marshes, disappointing because overgrazed, but we found Samolus valerandi, which is fairly uncommon here. 13th July: The party went up the west side of Loch Eck to Bernice on a damp, misty morning which was not helped by the drips from the natural oak woods; fortunately it cleared later. Lobelia dortmanna was growing in plenty and we found Veronica scutellata growing to about 24 ft in a bog. 14th July: We met at Achadunan farm in Glen Fyne and drove up to the Lairige hydro-electric dam at about 1,500 ft. It was a beautiful day, interrupted by some heavy thunder showers, and we found some pleasant but not unexpected sub-alpine plants before we went down to Glen Fyne proper, where a surprising number of common species were not on the master-card. A short visit to the house and policies of Ardkinglas House was marred by a thunderous downpour, and it was a damp party which enjoyed a welcome tea provided there by Mrs Noble. 15th July: The party went up Glen Masson and were joined, in pouring rain, by Dr Tayler. It was difficult to see anything and we could not climb the surrounding hills owing to mist. It did not appear a very rewarding area and we finally gave up after lunch. In the evening, however, it cleared and Miss Conacher took some of the party with Mrs Cassels to the salt-flats at the head of the Holy Loch, and added some species, several of garden origin, but Raphanus maritimus was interesting. 16th July: The party met at Kinlochruel and drove as far as the road goes past Colintraive up Loch Striven side. It was a beautiful day and the flushes down to the lochside were golden with Saxifraga aizoides. There was much Pinguicula lusitanica and some Parnassia palustris and Gymna- denia conopsea. On returning, we visited the policies of ruined ‘Southall’, very overgrown, and Polypodium interjectum and Listera ovata were seen. Nearer Colintraive, Polystichum setiferum grew on a damp rock, and a walk round Kinlochruel garden added several weeds to the list. 17th July: This was a most beautiful day. We met at the top of the pass into Hell’s Glen, where we were joined by Mr McClintock, and climbed Stob an Eas, 2,400 ft, where there was a real alpine flora. Asplenium viride, Cystopteris fragilis, Thalictrum alpinum, Rubus saxatilis, Alchemilla alpina and Juncus trifidus were surviving even the intense drought, and Salix herbacea looked very 278 REPORTS gay with scarlet, berry-like galls. We came down by the Ardnoe Burn, where there was a spectacular stand of Bromus ramosus, to Ardnoe, where we found Platanthera bifolia. There had been a dearth of orchids everywhere, probably due to the weather. A short visit to the shore of Loch Fyne ended the day. 18th July: This day was not really in an unworked area as we went to the serpentine outcrop in Glen Daruel to look for Asplenium cuneifolium, found by Mr McClintock, and to see the flora, which is unusual for this area. On the way up we saw a really magnificent display of Asplenium trichomanes in an old outhouse and a pretty trail of Mimulus moschata in a small burn. On the way down there were some of the largest patches of Lycopodium clavatum that we had ever seen. It was generally agreed that it had been an agreeable and rewarding week: though we had not added anything important to the underworked squares, we had found some overlooked plants and the weather had contributed to some very enjoyable days. R. MONTGOMERY CRAWFORD, LANARK. 21ST—26TH JULY Although the number of participants was low and the weather somewhat inclement, many of the 10 km squares in the south of the vice-county were visited and species added to the master cards, this being the main objective of the meeting. On Tuesday, the first interesting find was Erinus alpinus at Culter churchyard, with masses of white Cymbalaria muralis nearby on the walls. A roadside verge near Cormiston yielded the hybrid between Galium mollugo and G. verum—G. x pomeranicum. Pimpinella saxifraga was abundant here on the banks. In the afternoon Corehouse Nature Reserve was visited; Festuca altissima was found, but Poa chaixii proved elusive. Several Hieracia were collected for identification by Mr P. D. Sell. Snowhill was climbed on Wednesday, the gills providing the greater interest. Epilobium brun- nescens was abundant and Sedum villosum added a splash of colour to the bryophyte flushes. Gymnocarpium dryopteris and Thelypteris limbosperma were on the screes and Cochlearia alpina flowered in a restricted wet patch. A small area of open birch woodland near Earl’s Mill yielded ten sedges including Carex pallescens; Helianthemum chamaecistus and Cirsium heterophyllum were found on the roadside verge bordering the wood. : A brief visit to Carnworth Moss was made on the following day, but the more botanically — rewarding part were the railway banks nearby. Friday saw us climbing Tinto, which produced typical Calluna-heath with several rich flushes © near Greenshiels. Drosera rotundifolia and Narthecium ossifragum were found at the edge of a | fairly rich, upland meadow. Eguisetum sylvaticum formed a miniature forest on the broad roadside - | verge. Although there were few attenders, the time spent was enjoyable and I should particularly like to thank Mrs Allan for her fund of local knowledge. : L. FARRELL STORNOWAY, OUTER HEBRIDES. 2ND-9TH AUGUST | 18 members and guests attended the first official meeting to be held in Lewis since that led by. Miss U. K. Duncan in 1959. The aims were to study the grasses, sedges and rushes of the island | and to spend two days mapping the flora of under-recorded 10 km squares. | On Sunday, most of the party visited Lews Park, Stornoway, stopping first by a patch of sown grass to see Festuca pratensis, also found the previous day by Mr Copping beside the road at | Laxdale. On entering the park, a curious Agrostis, with short ligules, conspicuous paleas and lemmas long-awned from the base, was gathered and subsequently determined by Dr C. E. Hubbard as A. canina subsp. montana x A. tenuis. An awned form of Agrostis stolonifera var. palustris was found nearby, and in the woods surrounding Lews Castle grew Poa nemoralis, P. chaixii and Deschampsia cespitosa var. parviflora. An interesting discovery in stony ground by the castle was Festuca rubra subsp. megastachys. The Deschampsia, flowering later than the normal form of wet places, was particularly attractive. Miss E. R. T. Conacher and Mr Ribbons recorded the flora _ generally and found Trifolium hybridum, Epilobium adenocaulon, Salix purpurea x S. viminalis and Petasites fragrans. Rubus spectabilis and Cotoneaster microphyllus were noteworthy introduced species happily naturalized. | The next day was spent recording at Morsgail Lodge, Uig, by kind permission of Mr M. Mac- { REPORTS 279 donald. Mr Slack led most of the party in the ascent of Caultrashal Mor (749 ft), while four members concentrated on the area near the house. The former group discovered about 30 plants of Hammarbya paludosa, but in general found the area poor in species. The lower ground yielded Pinguicula lusitanica, flowering Utricularia minor, Carex limosa, C. pilulifera, C. pauciflora and Festuca pratensis again. It was, however, interesting to discover that we had added 12 of the Mapping Scheme’s 50 commonest species to a square where none had previously been found. Tuesday morning was devoted to visiting the Standing Stones at Callanish and the well-preserved broch at Dun Carloway, and the finding of Miss Conacher’s spectacles in uncharted moorland where she had mislaid them a month earlier on a private recording expedition with Miss Campbell. Encouraged by this good fortune we drove to the dunes, short turf and cliffs at Bosta on Great Bernera to see Catapodium marinum, and we also found splendid flowering spikes of the rosy- magenta-coloured Dactylorhiza fuchsii subsp. hebridensis, Carex hostiana, C. arenaria, Brachy- podium sylvaticum and Ammophila arenaria. A surprising discovery was Lamium moluccellifolium on the beach. ; Wednesday proved to be botanically one of the most rewarding days of the meeting. We first walked to an area of waste ground near Stornoway harbour to see Puccinellia capillaris, previously found there by Mr Copping. This species may well prove to be new to Lewis. On the walk back we passed Lolium multiflorum before setting off by car to the salt-marsh at Tong, which yielded Carex serotina and Puccinellia maritima. We continued to Coll, where Mrs Clark had previously seen Briza media, and ate our lunch among Agropyron junceiforme x A. repens. Then we explored the brackish stream, salt-marsh, pasture and pools to the west of the road. Within an hour we had seen Potamogeton pectinatus, Ruppia maritima, Zannichellia palustris, Juncus baiticus, Scirpus tabernaemontani, Eleocharis uniglumis, Blysmus rufus and a rich variety of commoner species. The final stop at Traigh Geiraha, north of North Tolsta, was remarkable for producing Eleocharis quinqueflora, E. multicaulis, E. palustris and E. uniglumis round the shores of a single loch, in addition to Catabrosa aquatica where fresh water was running over sand to the sea. Thursday was spent recording near Eishken Lodge by kind permission of Captain and Mrs A. M. Archdale, and Mr D. Kesting of Park Crofters Ltd. Three groups led by Mr Copping, Miss Conacher and Mr Ribbons explored Feirihisval (1,074 ft) and lochans, bogs and sea-cliffs to the east of the house. 192 species were found including Osmunda regalis, Juniperus communis, Utri- cularia neglecta, U. intermedia, Listera cordata, Hammarbya paludosa and Eriophorum vaginatum. _ Epilobium adenocaulon and planted Alnus incana were growing near the house. Friday took us to the Butt of Lewis, with rocky cliff tops dominated by Armeria maritima and Plantago maritima. However, we recorded Sagina maritima, Puccinellia maritima, P. capillaris (2 plants), and Catapodium marinum before moving to Eoropie beach with its sand-dunes and stream. There we were delighted to see Carex maritima and Avena strigosa growing in the sand. (Later, in Harris, four members of the party were surprised to find A. strigosa still being cultivated in the fields). The stream contained masses of Veronica anagallis—aquatica and Catabrosa aquatica. On our return to Stornoway we passed two fields at Barvas containing Phalaris arundinacea apparently being grown as a crop. The final botanizing of the week was an examination of the plants of the flower-beds outside our headquarters, the Acres Hotel, on Saturday morning. 24 wild species were found, none of which would have enhanced the reputation of the gardener. Suffice it to say that one was Agrostis gigantea, providing an addition to our list. So ended a meeting which had been blessed with fine, sunny weather almost throughout and had resulted in the discovery of 11 out of the 14 rushes, 30 of the 41 sedges, and 43 of the 49 grasses shown in the Atlas, and which, in addition, had added Festuca pratensis, Puccinellia capillaris, Briza media and Agrostis gigantea. A. CoPPING ot '# ¢ p* : “=e = eye | | re ae wot. olirtw:. ti enn) rah iad 2H HiNE, Jo manent +4 16. vd Xe sinsic 1 10 rely bsrsianounty wom BITIG Lot eon ahs.cu ron bebloiy batuorg aR! 7a od oo 1009.0710, ac ; ar ot bag axons 2 roll ie’ Seon) Zon) ptt ty sil . : oy, 10, £1. Bebb, bag ow ty 11, Tavopeh,.ad peribinyp sehen wid nsect dewoi ISI bel, scsi aigd ys 1) Ee fa yy evogs ; Dov s7919-1 wy. ori! bare cleins [in Sp OI? goi bass a 93 apiiein peta st) © poaioom ban sdogy th 206 gage z york RUG. 23 21M to. paibok ant, TPE wp “ey ini qaqa aati si neiiboges aolioces shavings keg Tihs plement denny: testi) OO eden is 21> Dos ‘hull ore. Lest mab =i 6 OL» ont, We ORO) beog, 4 . xii ta eplide.ernswor bibnsige bavol oaks, ow ‘bad eR man ibagpir choot .DOREAG...d.. DIAL. yi ARM. Ceaclas hess) ncitvelisnd | bewo Nolitkes: Muli Bow YIEVOID aniencqmue A sc wlidqorniunk, sap a 2 dlos-sirrogein Hall os fhoead ort) ste leu. aW .2nilsem.sd?>joevnb-soibamwer om ori To ene yileoy nod od 0} boven abe | feupivold 2D Nigns sibevigors 968 Of 1uodtad VeI/OrIOle J8e0, brugag)al em Ye RpRe, OT Dost lav dosed Alew ody gO. 2iwal olsyon.ad-ot sym, lie UPR 2790ga adt) .anigge> Mn tf bavecil hebisiy cot acot Js dean lice sdt.od io vd fo gaits: sidisd munca miod beeany ‘1 jauniveng. bed angio. oM .edw ile ol Douniicos.3 ORTON: plomaws, bee byioiqze ow codT made: box surcaligol gore vale a Has soaul y9.9)8 OMG ol ey ‘Nero avinate I Jo zozeam beniginos.t ait2 5 ; rf Wf ~wolat gniniainos asviwd ia ebb ows beebq ow Bg ‘ol 7 af Lat ; ehiziun siad-rawolk adsl: ‘SinBice ony. Yo ROUSCI AISI tn-sn0e aval saw roger” Phin’ bs, AeROry ¥8p ’ Ae PRM 900,180 Be OL BL salle *rrobisg ent, 10 gaits . lee La Puth Sti RS bid juaayo, ‘noma. ‘watiaove MOTI, «at icky by gozqinie, 26 oft to. th If, “eogtiaa | Atgds, 10,08 26 b Pe aka IDA, alana ANY pow be f chee iy ie ieee pone a fone fs fot ore {IID ah ¥, a? 9 ube Reods sande ever she ia emaplivas yur. parethira aes Fet) isbers, Sty. ® pasties, bape! i } ppg : oO ee ruveri 'y tative i += aa Seearmnity oe Tush Rape ae ) Ta rit Gon Srugpans Ring & 4] oka a 4 es Bioopammertsns el. 7 Te day ‘wih aper ring ah re ee Ae ts) i o y . aa is 7 14. 15. Botanical Society of the British Isles Conference Reports . BRITISH FLOWERING PLANTS AND MODERN SYSTEMATIC METHODS Ed. A. J. Wilmott. Pp. 104, 18 plates, wrappers. 1948. . THE STUDY OF THE DISTRIBUTION OF BRITISH PLANTS Ed. J. E. Lousley. Pp. 128, illustr., wrappers. 1951. . THE CHANGING FLORA OF BRITAIN Ed. J. E. Lousley. Pp. 204, 6 plates. 1953. . SPECIES STUDIES IN THE BRITISH FLORA Ed. J. E. Lousley. Pp. 189, 2 plates. 1955. . PROGRESS IN THE STUDY OF THE BRITISH FLORA Ed. J. E. Lousley. Pp. 128, 4 plates. 1957. . A DARWIN CENTENARY Ed. P. J. Wanstall. Pp. 140, 7 plates. 1961. . LOCAL FLORAS Ed. P. J. Wanstall. Pp. 120, 2 plates. 1963. . THE CONSERVATION OF THE BRITISH FLORA Ed. E. Milne-Redhead. Pp. 90. 1963. . REPRODUCTIVE BIOLOGY AND TAXONOMY OF VASCULAR PLANTS Ed. J. G. Hawkes. Pp. 183, 1 plate. 1966. . MODERN METHODS IN PLANT TAXONOMY Ed. V. H. Heywood. Pp. 312, illustr. 1968. . FLORA OF A CHANGING BRITAIN Ed. F. H. Perring. Pp. 158, illustr. 1970 (Paperback reprint: 1973). . TAXONOMY AND PHYTOGEOGRAPHY OF HIGHER PLANTS IN RELATION TO EVOLUTION Ed. D. H. Valentine. Pp. xii+ 432, illustr. 1972. . PLANTS WILD AND CULTIVATED Ed. P. S. Green. Pp. 232, 8 plates. 1973. THE OAK: ITS HISTORY AND NATURAL HISTORY Ed. M. G. Morris and F. H. Perring. Pp. 376, 8 plates. 1974. EUROPEAN FLORISTIC AND TAXONOMIC STUDIES Ed. S. M. Walters, with the assistance of C. J. King. Pp. 144, 4 plates.1975. Obtainable from: DR F. H. PERRING, OUNDLE LODGE, OUNDLE, PETERBOROUGH, PE8 5TN Hybridization and the flora of the British Isles Edited by C. A. STACE A comprehensive account of each of the 975 hybrids that has been recorded from the British Isles based on accounts prepared by over 80 specialists and skilfully edited and brought together in a single volume by Dr Stace. Up-to- date data are provided on their appearance, identification, fertility and distri- bution, and on the results of any experimental work which has been carried out on them. Many of the so-called hybrids are mere fanciful identifications; the evidence in such cases is assessed. A literature list is given for each hybrid, and an introductory section provides a general background to the whole subject of hybridization. In addition, 464 hybrids between British species which have been found abroad but not yet in the British Isles are listed. This reference work is an authoritative source of information for field botanists who wish to discover hybrids in the wild, and for professional botanists who wish to use hybrids for both research and teaching purposes. It also provides a stimulus for further research, as for the first time the gaps and deficiencies in our knowledge are precisely defined. The introductory section is a more complete synthesis of information on hybridization than has hitherto been available, and for the most part uses British and Continental examples to illustrate the principles discussed. Published in collaboration with the Botanical Society of the British Isles by Academic Press, London, New York, San Francisco. Pp. xiv+640. 1975. Price £14-80/$39.25. Obtainable from Academic Press, London, etc., 24-28 Oval Road, London, NW1 7DX. Natural History Book Reviews A new thrice-yearly journal (January, May, September) offering authoritative reviews of books on all aspects of ornithology, botany, zoology, entomology, conservation and the environment. Books reviewed range from the most advanced specialist monograph to elementary texts and children’s titles. While the emphasis is on titles in English, works in other languages of exceptional interest are also reviewed. Each issue contains one or more extended review articles surveying available literature in a particular subject-field. Natural History Book Reviews offers all those concerned with purchasing, or recommending the purchase of, natural history books at all levels an invaluable guide and enables the teacher and the interested amateur to remain abreast of current publishing developments in this field. Annual subscription (3 issues): £6.50 (U.S. $15.60) The Richmond Publishing Co. 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