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Kent, 75 Adelaide Road, West Ealing, London, W.13 9ED Watsonia, 11, 1-5 (1976). 1 Epipactis leptochila (Godfery) Godfery and E. phyllanthes G. E. Sm. occurring in South Northumberland on lead and zinc soils A. J. RICHARDS and G. A. SWAN University of Newcastle-upon-Tyne ABSTRACT Epipactis leptochila (Godfery) Godfery has been found in 11 stations in S. Northumberland (v.c. 67), 180 miles north of the nearest previously known site. Plants appear to be typical except for short perianth segments. All sites are under birch and associated with apparently toxic heavy metal soils; one site in particular has a very high level of zinc. Most sites are on apparently natural gravels beside the Rivers Tyne and South Tyne, but one is by a disused lead mine. At another lead mine E. phyllanthes G. E. Sm. var. pendula D. P. Young is reported. This species is new to South Northumberland, the nearest previously reported station being in south-west Furness (v.c. 69b). On lead spoils at another site E. helleborine was found; depauperate forms of this species resemble FE. leptochila vegetatively. The significance of Epipactis on heavy metal sites in Northumberland is discussed. INTRODUCTION Epipactis leptochila (Godfery) Godfery was said by Young (1962a) to be confined in England to calcareous areas in the south, where it is frequent very locally on the escarpments of the Chilterns and Cotswolds. Elsewhere it is uncommon, in scattered localities along the Chalk range from Wiltshire to Kent, on the Carboniferous Limestone of the Wye Valley and Mendips, and in two localities in Devon (Perring & Walters 1962). It also occurs in widely separated localities in France, Germany, Denmark, Sweden and Switzerland. The plant is said to be strongly calcicolous — much more so than any other European species of Epipactis. Every station for which geological data are available is on calcareous rock, and these comprise the great bulk of records. Conversely no station had been recorded on neutral or acid rocks. The usual habitat is in beech-woods under heavy shade, and with only a sparse ground flora; it often occurs with E. helleborine (L.) Crantz and E. purpurata Sm. From these, and from other related Epipactis species, it is readily distinguished by rather narrow, yellowish, two-ranked leaves, relatively few, yellowish flowers, which are held in an inclined position and which lack a rostellum, with rather long, narrow perianth segments, and a patent epichile which is longer than broad, and which equals or exceeds the hypochile. The anthers are borne on a stalk (clinandrium) in such a way that when viewed sideways a distinc- tive hole is visible between the column and anther (Young 1962a). As such it is usually a rather distinct species, and it has not normally been considered as being a critical member of this difficult genus. It was therefore surprising to receive from M. Rawes and B. M. Cunliffe, towards the end of 1973, a record of E. leptochila from birch scrub on the east bank of the River South Tyne near Williamston, S. Northumberland (v.c. 67), 4 miles north-north-east of Alston (grid references of all sites are withheld, but have been deposited with the Biological Records Centre, Monks Wood). This identification had been confirmed by Dr J. T. Knight. It later transpired that the same colony had been observed and photographed a month earlier on the 21st July 1973 by Dr A. W. Williams, who had known it for some years. HABITAT AND DISTRIBUTION The colony was located on the 14th July 1974 by G. A. and M. Swan, although on that date it was only in bud, and a further search by A. J. Richards and G. A. Swan on 31st July revealed that rN 2 A. J. RICHARDS AND G. A. SWAN it was part of a larger colony of Epipactis with about 290 flowering spikes within a limited area. A voucher specimen was taken (herb. Swan) and some mature flowers were pickled in 70% ethanol. Reference in the field to Clapham (1962) and Young (1962a) established that the colony consisted of typical E. leptochila, differing only in short outer (mean 7-4 mm) and inner (mean 5-9 mm) perianth segments. All mature flowers examined in this and other populations lacked a rostellum and possessed a clinandrium. In addition, all had a narrow, patent, triangular epichile. In view of the apparent lack of floral measurements in the literature (except Godfery 1933), those from one population of E. leptochila and two of E. helleborine are presented in Table 1, together with measurements made from Ross-Craig’s (1971) excellent scale drawings. At this locality the River South Tyne forms a broad valley at an altitude of 700 ft (220 m) with level gravels in the immediate vicinity of the river. To the east of the gravels, away from but scarcely raised above the river, is a thinly grassed bank on which birch scrub grows. E. leptochila occurs in scattered colonies both within and at the margin of this scrub. We noted the following species growing within 1m of E. leptochila: Angelica sylvestris Conopodium majus Leontodon hispidus Anthoxanthum odoratum Dactylorhiza fuchsii Ranunculus acris Armeria maritima Epipactis helleborine Salix nigricans Arrhenatherum elatius Equisetum arvense Silene dioica Betula pubescens Geum rivale Succisa pratensis Carex ovalis Holcus lanatus Trifolium repens Centaurea nigra Open gravel and pathside within 5m of E. leptochila colonies, to the south and east, also supported : Acer pseudoplatanus G. verum Trisetum flavescens Aegopodium podagraria Hieracium sp. Ulex europaeus Agrostis canina Linum catharticum Vicia cracca A. tenuis Lotus corniculatus V. sepium Campanula rotundifolia Luzula sylvatica Viola lutea Cerastium fontanum Minuartia verna Cruciata laevipes Parnassia palustris Bryum caespiticium Deschampsia cespitosa Plantago lanceolata B. capillare Euphrasia nemorosa Rubus idaeus Pleurozium schreberi Festuca rubra Rumex acetosa Polytrichum juniperinum Filipendula ulmaria Sorbus aucuparia Rhytidiadelphus squarrosus Fraxinus excelsior Thlaspi alpestre Tetraplodon mnioides Galium saxatile Thymus drucei Weissia controversa vat. densifolia The open nature of these gravels, which are almost bare in places, and the occurrence of species such as Armeria maritima, Minuartia verna, Thlaspi alpestre and Weissia controversa vat. densifolia led us to suspect that the mineral content of the soil might be unusual. In the vicinity of Alston, lead, zinc and barium ores have been mined in the past. Soil samples were therefore taken from 2-10 cm below the surface in the immediate vicinity of E. leptochila, and from the bare patches in the adjacent open gravel. These were analysed by Dr K. Shaw (Agricultural Development and Advisory Service, Kenton Bar, Newcastle-upon-Tyne) (Table 2). The Epipactis sites are remarkable, with high levels of extractable zinc, of an order likely to result in toxicity in many plants (Bradshaw et al. 1965), although if the calcium level (unknown) is high relative to magnesium, there would be a less severe effect. However, the zinc levels are nearly twice the ‘critical’ level suggested by Halliday (1960). Levels of extractable lead are also rather high. The levels of extractable phosphorus and potassium are very low although not untypical of many upland soils, particularly those with a high pH. The pH of these soils is about 7 (neutral), which is unusual in this predominantly acidic area, and may well be comparable with the southern chalk soils on which this species has been previously reported. The bare gravel areas are not dissimilar, although the figures are less extreme, particularly with regard to zinc, which is only a tenth of the Epipactis soils. This, coupled with the relatively high levels of lead and the low levels of phosphorus, are sufficient to account for the sparse but florist- ically interesting vegetation found in these areas. Nevertheless, it is noteworthy that the more EPIPACTIS LEPTOCHILA AND E. PHYLLANTHES IN S. NORTHUMBERLAND 3 extreme soils, under birch, support a more vigorous ground vegetation. This may be due to such ameliorating factors as the shade, moisture and higher humus of these areas, and the higher pH. It may be that the open nature of the gravel communities owes much to free drainage, high leaching and low humus content, as Halliday (1960) has suggested that little direct heavy metal toxicity is associated with lead/zinc spoil soils in the Pennines. Similar possibly toxic gravels are found beside the River South Tyne from Williamston down to the junction with the River North Tyne at Warden, and species such as Armeria maritima, Thlaspi alpestre and Minuartia verna occur beside the combined River Tyne even below Wylam, only 7 miles west of Newcastle. Since we thought that birch scrub growing on apparently toxic Tyne gravel might constitute a habitat for E. leptochila, we went downstream on the same day in search of other examples of this type of habitat, and indeed we never failed to find the Epipactis in suitable areas. South-east of Harper Town, the South Tyne forms a bend, on the western (and southern) bank of which a fairly level area is covered with birch scrub at an altitude of 500 ft (160 m). Here, growing in the scrub and also on an open grassy bank just above the river, we saw about 1,000 TABLE 1. FLORAL CHARACTERS OF EPIPACTIS LEPTOCHILA AND E. HELLEBORINE Outer perianth Inner perianth Epichile Epichile L Origin Sample segment (mean) segment (mean) (mean) Hypochile L size (mean) Length L/B Length L/B Length L/B Ros- = Clin- tellum andrium E. leptochila Williamston 9 T4 24 5-9 I-75 3-5 1:9 1-10 —_ ae Tip patent Clapham (1962) _- 12-15 ca. 10 Longer _ + than broad Tip patent Ross-Craig (1971) ~ 10 1-9 9-6 1-8 4-5 a2 0-90 _ a= Tip patent E. helleborine Haydon Bridge 10 9-6 1:6 7:9 ies 2°4 0-8 0-44 + — Tip recurved Langley 9 8°38) .91:5 7:0 1-4 3-0 0-7 0-64 + —_ Tip recurved Clapham (1962) — ca. 10 Broader + — than long Tip recurved Ross-Craig (1971) — 11:0 =1-55 9-6 1-5 5:0 0:8 1-0 + ~ Tip recurved All measurements in mm spikes of E. leptochila. These were very similar to those at Williamston, although some were much bigger plants. There was also one plant of E. helleborine. The associated species were very similar to those at Williamston, but the following additional species were also noted: Anthyllis vulneraria Polygala vulgaris Taraxacum unguilobum Geranium sylvaticum Rhinanthus minor Two further sites were visited the same day. On the southern bank, west of Beltingham, 13 spikes were found; and on the northern bank, near Crow Hall, 6 spikes were discovered after a long search. In both these sites, birch occurred on riverside gravels, but there was little evidence of soil toxicity and none of the characteristic gravel plants were present. The vegetation under the birch was much more rank and species-rich, and the Epipactis were poorly developed and apparently suffering from competition. This was also true of the other six riverside sites sub- sequently discovered downstream to Wylam during August. All were on riverside gravels or banks associated with birch, and in some cases (as at Wylam and Haltwhistle) open gravel or banks with 4 A. J. RICHARDS AND G. A. SWAN Armeria, Minuartia and Thlaspi occurred nearby. Nevertheless, colonies were invariably small both in number of spikes (2-20) and height, occurring in areas with vigorous ground cover. It is suggested that these birch woods might have colonized toxic gravels, and have developed a deep humus above the gravels, allowing more vigorous ground cover to develop. In view of the association of E. leptochila with apparently toxic sites under birch, spoil heaps from disused lead mines were examined. Some of these proved negative, but a thriving colony of more than 50 spikes, accompanied by E. helleborine, was discovered in birch scrub on spoil heaps (altitude 450 ft, 140 m) some distance north of the River Tyne at Hexham. TABLE 2. ANALYSES OF SOIL SAMPLES FROM WILLIAMSTON Site pH! p2 K3 Meg? Cu5 Zn6 Pb® E. leptochila 72 3 56 99 26 2,370 692 E. leptochila 7:0 4 54 113 18 2,300 300 Open gravel 5:2 6 91 61 12 192 540 Open gravel Dy 6 91 48 15 222 720 All element concentrations as p.p.m. of air-dry soil. 1 Measured by glass electrode on 1:2 soil: water mixture. 2 Extracted in sodium bicarbonate and determined by the phosphomolybdate procedure. 3 Extracted in N ammonium nitrate and determined by flame-photometer. 4 Extracted in N ammonium nitrate and determined by atomic absorption spectrophotometer. 5 Extracted in EDTA and determined by atomic absorption spectrophotometer. 6 Extracted in 0-5N acetic acid and determined by atomic absorption spectrophotometer. On 11th August 1974, G. A. and M. Swan visited the old heaps of a lead mine (altitude 400 ft, 125 m) north of Haydon Bridge. Although birch was abundant they failed to find E. leptochila. However, this disappointment was more than compensated for by the discovery of a flourishing colony of E. phyllanthes G. E. Sm. var. pendula D. P. Young. About 50 spikes were present, growing particularly under Corylus avellana, along with Salix spp., Alnus glutinosa, and Betula sp. Although the plant occurred to within 5 m of the edge of a lead spoil tip 20 m high, and the ground vegetation was open, it was under heavy shade, and there was no external indication that the spoil was in any way affecting the soil or the vegetation. However, patches of birch scrub adjacent to and on the heaps held magnificent colonies of E. helleborine, of which there were several hundred spikes. The nearest records for E. phyllanthes to Haydon Bridge appear to be in W. Lancs. (v.c. 60) and S.E. Yorks. (v.c. 61) (Young 1962b), and south-west Furness (v.c. 69b) (Young 1952). The Lancashire record is known to be of var. pendula. This locality, therefore, is not only the first record of E. phyllanthes for S. Northumberland but it is also the northernmost known in the British Isles. The record from Holy Island (Cheviot, v.c. 68) cited by Swinton (1967) must surely be an error for E. dunensis (T. & T. A. Stephenson) Godfery, which is well known there (Young 1962a). Comparison of the Haydon Bridge plants (voucher in herb. Swan) with the account in Young (1952) shows these attractive and distinctive little plants to be in agreement with his var. pendula in every characteristic. We also inspected the heaps at the site of the disused lead smelting works at Langley (altitude 700 ft, 220 m) where very many magnificent plants of E. helleborine were growing, particularly under conifers. Among birch scrub on these heaps were also some smaller, yellower plants with leaves in two ranks, which had the superficial appearance of E. leptochila. However, the flowers of both possessed a rostellum, lacked a clinandrium, and had an epichile and perianth segments indistinguishable from the Haydon Bridge population of E. helleborine (Table 1). Doubtless, they were merely odd and rather depauperate forms of this species. DISCUSSION The origin of E. leptochila in the Tyne Valley is quite unknown, although it can give rise to some interesting speculation. In view of the constancy of its habitat elsewhere it may be supposed that EPIPACTIS LEPTOCHILA AND E. PH YLLANTHES IN 8S. NORTHUMBERLAND 5 it originally arose in woodland sites on chalk and limestone where it is widespread in this country and abroad. The majority of its close relatives are also most usually found in this type of habitat. In common with most members of the Orchidaceae it possesses a very light seed which is very readily windborne, perhaps over large distances on occasion. Thus the natural colonization of suitable habitats in Northumberland from the south of England is not beyond question. The Tyne Valley sites are not apparently of very recent origin. Helleborine latifolia Druce is recorded from open sites by the banks of the Tyne near Haltwhistle by Blackburn (1927), and her report is accompanied by an editorial note from J. E. Hull that he knew it in similar situations at Feather- stone in 1897. Since E. leptochila had not been distinguished at that time, it has probably been present in the Tyne Valley for at least 80 years. The Tyne Valley habitats share with southern sites features of shade, rather sparse ground flora, and, in one case at least, a neutral pH. Furthermore, they seem to be attractive to other species of Epipactis, E. helleborine being larger and more frequent here than elsewhere in Northumberland, and the very local £. phyllanthes occurring in one site. Nevertheless, the relatively high altitude (up to nearly 800 ft) must render these sites much colder than those in the south, and the unusual mineral content of the soil would be thought to pose many problems to these plants. However, it apparently has the effect of creating open ground suitable for these species; the poor showing of Epipactis in downstream sites with good ground cover was notable, and it may be that previously vigorous colonies have suffered from the progressive colonization of heavy metal gravels by birch, with subsequent increase of ground cover. REFERENCES BLACKBURN, D. B. (1927). Observations in the neighbourhood of Haltwhistle (vice-county 67). Vasculum, 10: 62-63. BRADSHAW, A. D., MCNEILLY, T. S. & GREGORY, R. P. (1965). Industrialisation, evolution, and the develop- ment of heavy metal! tolerance in plants, in GOODMAN, G. T. et al., eds. Ecology in the industrial society, British Ecological Society Symposium, 5: 327-343. CLAPHAM, A. R. (1962). Epipactis, in CLAPHAM, A. R., TuTIN, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed., pp. 1016-1021. Cambridge. GoprerY, M. J. (1933). Monograph and iconograph of the native British Orchidaceae. Cambridge. HALLIDAY, G. (1960). Studies on the Arenaria ciliata and Minuartia verna complexes. Ph.D. thesis, University of Cambridge. PERRING, F. H. & WALTERS, S. M., eds (1962). Atlas of the British flora, p. 333. London. Ross-Cralia, S. (1971). Drawings of British plants, 28: Plate 38. London. SWINTON, Mrs (1967). Lindisfarne (Holy Island) Botanical meeting. Hist. Berwicksh. Nat. Club, 37: 204. Youna, D. P. (1952). Studies in the British Epipactis, 3. Epipactis phyilanthes G. E. Sm., an overlooked species. Watsonia, 2: 253-276. Youna, D. P. (1962a). Studies in the British Epipactis, 5. Epipactis leptochila; with some notes on E. dunensis and E. muelleri. Watsonia, 5: 127-135. Youn, D. P. (1962b). Studies in the British Epipactis, 6. Some further notes on E. phyllanthes. Watsonia, 5: 136-139. (Accepted May 1975) bis NAS ae Brest st oils en tt TR ee ey, ie bint Ota halle ella en Oakey ze ridobery ienmrivbe! e Pt) a! einai 6 weadone ae, bed - ahs on ef 7; rior ae yy i ik acai, pn ci 8 ine eng wy: 20: “i inate a0 ‘ ‘oy ; ‘hue aN 10 +t Sokal Hibs seme | on bed ‘ abihosasy, a gone, ; % proud ca holt Vath Ga T Tha igite eistid. Hs, meet di) ba oe Re etn pee ; Ab latin ee ues my ee hy Sy) fr3 a sel wy Nea Be Los i ANS hd ie oa ile * seine gait, TION TLE Ha RTL mad 8 “a ne Ineuport mony brs wgial gaied “agin ie gota i: 30 oe sae _ ) nudwol § nk eye yout an pee lrg i A UE Nl py ae ent : @ f ' Bt a. hee hare, PRAT gel ty iri wos f eaten ecw oO WO HI Re. PEPE PEAR, Bake yor bermtive ovett ‘ ARE ue ; nb baad 1a, la ‘ { oy Wax 1 " Re eo manta » Lapa. 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As eA a PAs fe : i : wa tonlth ee selthiy a i} Choe | } he aa y Revive ‘aif Voth ian e Neb pete hae Ts hairs said ‘vibe 3 ag et een y AUT Eto oe ott is De era \e BVM: ere ay ae tele 2 eth Be epic HARTA et wagnvornnnt . ‘ feauin pad L.tbrstel wha) urate ba at nA RE 4 Te Bee Gh eLabier : i heads rage J RNG Ged (gh eine 6h Gi, eayhul sf Ae. tal ssi ane a ne - hy " ‘bY PD acer (em eae ae nt y il ie Dae ey A, eo TOE BY Oe SW TRS OIORS ETOO CIA) oe AVVO beta te yh * of 2 ¥ AY “ Os Pes LX ORION Kee f orl Ph Gan ae Ff ‘ \ ae i j yrs i 1 } * i hs ae i iN 4 " \ ee ee tr eee TAN eR hum GCC cw AMP f f ; CARL A crf ha is he : : a ‘ ai q Yai 5 f Rpts ‘ f ) 5 Pet i ! i. \ q a 5 pi Vi 4 j i ‘ " Ai! y 4 Dae at Ny Wy } | Y aye ’ i ; y i} ye Fi Watsonia, 11, 7-31 (1976). 7 Studies on variation and evolution in Centaurium erythraea Ratn and C. littorale (D. Turner) Gilmour in the British Isles 1. Taxonomy and biometrical studies R. A. E. UBSDELL Department of Botany, University of Reading* ABSTRACT Analysis of populations of Centaurium erythraea Rafn subsp. erythraea and C. littorale (D. Turner) Gilmour subsp. littorale has shown that both show a wide range of morphological variation. Much of this variation is retained in cultivation and five varieties are recognized of subsp. erythraea and four of subsp. littorale. Despite this variation the two subspecies are quite distinct. Analysis of mixed populations of the two species from the coasts of Anglesey and Lancashire has shown that some plants are, to varying degrees, intermediate and probably of hybrid origin. In the majority of populations from the Lancashire coast, backcrossing to C. littorale appears to have taken place, although two suggest backcrossing to C. erythraea. This is in contrast to the situation in the population from Anglesey in which only F,-like hybrids were present. Analysis of two populations from northern Germany show all plants to be intermediate and F,-like. INTRODUCTION The genus Centaurium Hill (Erythraea Borkh. nom. illegit.) of the Gentianaceae is widespread in Europe, especially around the Mediterranean. Taxonomic investigations of the genus have pre- sented great difficulties since the species are extremely variable. Parallel variation in several characters is common in the groups of related species, and there have also been reports of natural hybrids. Species and infra-specific taxa are, therefore, often difficult to define and the literature is full of nomenclatural confusion. However, Zeltner’s work (1970) has led to a better understanding of relationships within the genus. He concentrated primarily on cytological studies, but correlated this with evidence from morphology, ecology and geographical distribution. He showed that in Europe the genus consists of certain ancient, morphologically distinct species, together with a number of taxonomically difficult complexes. It is in the subsections Parviflorae (Ronniger) Melderis, Vulgaria Melderis and Centaurium of the section Centaurium that the nomenclature is particularly confusing since the principal species of each, C. pulchellum (Sw.) Druce, C. littorale (D. Turner) Gilmour and C. erythraea Rafn, show parallel variation in many of the characters and numerous subspecies and varieties of each have been described without adequate study. Subsection Parviflorae consists of two annual species, C. pulchellum and C. tenuiflorum (Hoff- manns. & Link) Fritsch, both of which grow in the British Isles. The former is hypotetraploid (2n = 36), while the latter has both diploid (2n = 20) and tetraploid (2n = 40) races, of which only the diploid is found in the British Isles. Subsection Vulgaria consists of the narrow-leaved, biennial species. Three are diploid (2n = 20) and confined to Spain, Portugal and southern France, while another two are tetraploid (2n = 40). Of the latter, one is found only in Spain and Portugal, while C. /ittorale is present in central and northern Europe and is the only species of this subsection found in the British Isles. There is a sixth species, confined to Spain and Portugal, for which no chromosome count is available. Subsection Centaurium consists of the broad-leaved, biennial species. One of these is diploid and confined to the Mediterranean region, while C. erythraea is the most widespread species of the * Present address: Botany School, University of Oxford. 8 R. A. E. UBSDELL genus and is the only one of this subsection present in the British Isles. It has both diploid and tetraploid races, the diploids being confined to the Mediterranean region while the tetraploids have spread into central and northern Europe including Russia and the British Isles. The tetra- ploids also appear to have been introduced into North America and Australia. Zeltner’s work and Melderis’s recent account of the genus for Flora Europaea (Melderis 1972a & b) have cleared up much of the nomenclatural confusion surrounding the European species of these three subsections. However, little experimental work has been carried out on the taxa of northern Europe, especially on C. erythraea subsp. erythraea and C. littorale subsp. littorale. Both are extremely polymorphic and numerous varieties of each have been described, particularly in the British Isles (Wheldon & Salmon 1925, Gilmour 1937). There is still much confusion over the status of many of these varieties as they are based upon poorly defined characters and given only vague descriptions. Four varieties of C. erythraea subsp. erythraea were recognised by Gilmour and named under C. umbellatum Gilib. as follows: i) Var. centaurium (L.) Gilmour is the typical, tall variety of the species. ii) Var. fasciculare (Duby) Gilmour is distinguished by its numerous axillary branches (the plant is often as broad as it is tall). It occurs in exposed places on the coasts of Scotland and Ireland. iii) Var. subcapitatum (Corb.) Gilmour is strictly maritime and distinguished by its solitary stem, crowded cauline nodes, compact inflorescence and dwarfness. It has been confused with C. capi- tatum (Willd.) Borbas, a dwarf, maritime plant with the stamens inserted at the base of the corolla- tube. C. capitatum was originally treated as a distinct species but is now included in C. erythraea subsp. erythraea as var. capitatum (Willd.) Melderis (Melderis 1972b). iv) Var. sublitorale (Wheldon & Salmon) Druce was thought (Wheldon & Salmon 1925) to be a hybrid between C. erythraea and C. littorale as it has certain characteristics of both species, but without definite evidence they treated it as a variety of the former. These varietal names will be used throughout this account since the combinations under C. erythraea do not exist, although Melderis (1972b) has shown that the name C. umbellatum Gilib. is an invalidly published, nomenclatural synonym of C. erythraea Rafn. Gilmour also recognized four varieties of C. littorale subsp. littorale. Two from the coasts of north-eastern England and eastern Scotland are glabrous, while the other two from the coasts of North Wales, north-western England and south-western Scotland are scabrid with a dense covering of small papillae on the stem, leaves, bracts and calyx: i) Var. occidentale (Wheldon & Salmon) Gilmour is the common scabrid variety found on the coasts of North Wales, north-western England and south-western Scotland. ii) Var. bayleyi (Wheldon & Salmon) Gilmour is distinguished from the former by its dwarf, much branched habit (the plant is as broad as it is tall), compact inflorescence, longer leaves and larger flowers. It is reported from the coasts of Anglesey and Lancashire. iii) Var. littorale is a dwarf, glabrous variety distinguished by its habit, short calyx and long capsules. It is reported from the coasts of north-eastern England and eastern Scotland. iv) Var. minor (Hartm.) Gilmour is reported from a few places on the coasts of north-eastern England and eastern Scotland, but it is difficult to see how it differs from var. littorale except in the lengths of the calyx and capsule. There have also been reports of hybrids, plants morphologically intermediate between the two species, from mixed populations on the coasts of Lancashire and North Wales, and these add to the confusion. Wheldon (1897) described in detail the characteristics of some plants which he collected from Hightown, S. Lancashire, v.c. 59 in 1894 (specimens in BM). He thought them to be hybrids between C. littorale and C. erythraea since they had some of the characteristics of both, but named them Erythraea littoralis var. intermedia Wheldon as.they resembled C. littorale more closely and were also highly fertile. Salmon & Thompson (1902) referred to a series of plants collected by them from Ansdell, W. Lancs., v.c. 60 (in BM). Some of them were clearly C. erythraea, others C. littorale, while the rest were intermediate and sterile. They thought that the latter might be hybrids. CENTAURIUM ERYTHRAEA AND C. LITTORALE: MORPHOLOGY 9 Wheldon & Salmon (1925) later described a variety of C. erythraea under the name of Erythraea centaurium var. sublitoralis Wheldon & Salmon from western England and Wales, which they considered to be a hybrid between C. erythraea and C. littorale. However, without definite evidence of this they treated it as a variety of the former, and the name C. umbellatum var. sublitorale (Wheldon & Salmon) Druce was later given to it (Druce 1926). O’Connor (1955) described the results of an investigation of Centaurium from the dunes at Freshfield, S. Lancs., v.c. 59 carried out by her in 1954. She found distinct populations of each species, and other populations which contained both species together with plants that could not be referred to either. Vohra (1970) aiso looked in detail at populations from Freshfield and Ainsdale, S. Lancs., and his results were the same as those of O’Connor. O’Connor also remarked that plants possessing characters similar to those classed as inter- mediates in mixed populations from the west coast have been reported from East Anglia, although C. littorale is not known there. A detailed biosystematic study was therefore carried out by the present author to investigate and describe the variation patterns shown by populations of C. erythraea subsp. erythraea and C. littorale subsp. littorale in the British Isles and northern Europe. It was hoped that an experi- mental approach to this problem wouid both help to reduce some of the taxonomic confusion surrounding the numerous varieties of these two subspecies and explain the nature of the morpho- logically intermediate plants in mixed populations. This work formed the contents of a thesis accepted for the degree of Ph.D. by the University of Reading (Ubsdeil 1973) and will be presented in three papers. This first paper is concerned with a biometric analysis of morphological characters. The purpose of this study was to assess the extent of the variation in certain morphological characters, and by so doing to delimit the range of variation shown by these two species. This information was then used as a basis for assessing the morphological evidence for hybridization. An account of the cytology will be given in a second paper, and a final paper will deal with the isolating mechanisms which normally keep these two species distinct in the wild. TABLE 1. LOCALITIES OF POPULATIONS SAMPLED Centaurium erythraea Centauriuin littorale 1. Kevington, W. Kent, v.c. 16 A. Aijinsdale, S. Lancashire, v.c. 59 2. Headley, Surrey, v.c. 17 D. Ajnsdale, S. Lancashire, v.c. 59 3. Box Hill, Surrey, v.c. 17 F. Ainsdale, S. Lancashire, v.c. 59 4. Orpington, W. Kent, v.c. 16 G. Ajinsdale, S. Lancashire, v.c. 59 5. Folkestone, E. Kent, v.c. 15 H. Ajnsdale, S. Lancashire, v.c. 59 6. Luccombe, Isle of Wight, v.c. 10 BA. Newborough, Anglesey, v.c. 52 7. Sandown, Isle of Wight, v.c. 10 N. Newborough, Anglesey, v.c. 52 8. Swanage, Dorset, v.c. 9 24. Holy Island, Cheviot v.c. 68 10. Bonchurch, Isle of Wight, v.c. 10 R. Ross, Cheviot v.c. 68 11. Freshwater, Isle of Wight, v.c. 10 22. Nairn, v.c. 96b 12. Steyning, W. Sussex, v.c. 13 EL. near Elgin, Moray, v.c. 95 13. Bude, E. Cornwall, v.c. 2 T. Tain, E. Ross, v.c. 106 23. Aberlady, Haddington, v.c. 82 DBO. Dornoch, E. Sutherland, v.c. 107 25. Newbiggin, S. Northumberland, v.c. 67 26. Cardiff, Glamorgan, v.c. 41 Mixed populations of the two species 27. Fanore, Clare, v.c. H9 14. Hightown, S. Lancashire, v.c. 59 28. Funshin, S.E. Galway, v.c. H15 15. Freshfield, S. Lancashire, v.c. 59 I. Minsmere, E. Suffolk, v.c. 25 16. Freshfield, S. Lancashire, v.c. 59 J. |Minsmere, E. Suffolk, v.c. 25 17. Freshfield, S. Lancashire, v.c. 59 K. Wells, W. Norfolk, v.c. 28 18. Freshfield, S. Lancashire, v.c. 59 L. Cley, E. Norfolk, v.c. 27 19. Freshfield, S. Lancashire, v.c. 59 20. St Annes, W. Lancashire, v.c. 60 Populations of C. erythraea with stamens 21. Ainsdale, S. Lancashire, v.c. 59 inserted at the base of the corolla-tube AN. Newborough, Anglesey, v.c. 52 29. Freshwater, Isle of Wight, v.c. 10 B, C, E, Ainsdale, S. Lancashire, v.c. 59 30. Freshwater, Isle of Wight, v.c. 10 FAL. Falshdft, Schleswig-Holstein, Germany Sae. St Peter, Schleswig-Holstein, Germany 10 R. A. E. UBSDELL MATERIALS During the summer months of 1970-72, populations of C. erythraea subsp. erythraea and C. littorale subsp. littorale were sampled from many parts of the British Isles and from two localities on the coast of north-western Germany. These consisted of either a single species or mixed populations of both, and the localities are listed in Table 1. METHODS The populations of single species were subjected to detailed biometric analyses, and the results compared with those from similar analyses of mixed populations, to determine whether the overlap in variation in the mixed populations was due to variation within the species themselves or to hybridity. Samples of seed from many of the wild populations were sown in the greenhouse and raised to maturity under uniform conditions. Mature flowering plants were pressed and measured for comparison with the wild plants. Plants of C. erythraea subsp. erythraea and C. littorale subsp. littorale from the Continent (northern Europe) were obtained as seeds from Botanic Gardens and raised in the greenhouse, as only two populations (both mixed) were sampled in the wild. Sets of voucher specimens have been deposited in BM and OXF. CHARACTERS CHOSEN AND METHOD OF ASSESSMENT After careful consideration of the characters used by Wheldon & Salmon (1925), Melderis (1932) and Warburg (1962) to distinguish between C. erythraea subsp. erythraea and C. littorale subsp. littorale including the numerous varieties of each, the following characters were chosen: 1. Height of stem This was measured vertically from the base of the basal rosette of leaves to the tip of the highest flower. 2. Habit The number of main flowering stems and cauline nodes were noted. The lengths of the internodes were also measured to see if the cauline nodes were well spaced or crowded together, and the following three categories were used for scoring: a) L = long, internodes >2-5 cm long, b) M = medium, internodes 1-0—2-°5 cm long, c) S = short, internodes < 1-0 cm long. 3. Inflorescence The total number of cymes present on each flowering stem was scored as follows: a) M = many, >5 cymes, b) I = intermediate, 3-5 cymes, c) F = few, <3 cymes: The length of the internodes between each cyme in the inflorescence was scored to give an estimate of the relative laxity or compactness of the inflorescence and was scored as follows: a) L = lax, >2-0 cm long, b) I intermediate, 1-0—-1-9 cm long, c) D_ = dense, 0:5-0-9 cm long, d) VD = very dense, <0-5 cm long. 4. Length|breadth ratio of cauline leaves This ratio was obtained by dividing the total length of the leaf (in mm) by the greatest breadth. For each plant the first three cauline leaves above the basal rosette of leaves were measured and a mean value obtained. CENTAURIUM ERYTHRAEA AND C. LITTORALE: MORPHOLOGY 11 5. Shape of cauline leaves All cauline leaves could be classified without much difficulty into one of the following categories: a) Elliptical = leaves short and broad with sides never parallel,an acute apex and 5 prominent veins, b) Linear = leaves long and narrow with parallel sides, an obtuse apex and 1 prominent vein, c) Linear-elliptical = leaves intermediate in dimensions but with sides never parallel, an acute apex and 3 prominent veins. 6. Indumentum Plants were scored as follows: a) Scabrid = margins and midribs of cauline leaves, upper parts of stem, bracts and calyx densely covered in small papillae, b) Semi-scabrid = all parts lightly covered in papillae, c) Glabrous = papillae absent. 7. Calyx/corolla-tube ratio This ratio was obtained by dividing the total length of the calyx from the base to the tip of the teeth (in mm), by the length of the corolla-tube. Ten flowers per plant were scored and a mean value obtained. 8. Length of corolla lobes This was measured from the top of the corolla-tube to the tip of the corolla lobes on live material only. Ten flowers per plant were scored and a mean value obtained. 9. Diameter of the pollen grains The pollen grains were mounted in cotton blue/lactophenol and a mean diameter calculated for each plant from 25 grains. 10. Shape of stigma This could be classified into one of three categories: (a) (b) (c) 11. Length of filaments This was measured from their point of attachment to the corolla-tube to the base of the anther. 12. Length of capsule RESULTS The data derived from the morphological studies can best be considered in three sections: a) The variation shown by individual characters, b) An assessment of the overall variation shown by different populations, c) The correlation of the different characters within individual plants. VARIATION OF INDIVIDUAL CHARACTERS For each of the characters studied, the mean, standard deviation and coefficient of variation were calculated for twenty plants in each population. The results are given for populations of C. erythraea and C. littorale in Tables 2 and 3 respectively, and for mixed populations of the two in Table 4. A comparison of C. umbellatum var. subcapitatum and C. erythraea subsp. erythraea var. capitatum is given in Table 5. R. A. E. 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LITTORALE: MORPHOLOGY rs TABLE 5. MEAN AND STANDARD DEVIATION FOR CHARACTERS OF PLANTS FROM POPULATIONS OF C. ERYTHRAEA VAR. CAPITATUM, C. UMBELLATUM VAR. SUBCAPITATUM AND C. UMBELLATUM VAR. CENTAURIUM (for abbreviations see Table 2) C. umbellatum C.umbellatum C.erythraea C.umbellatum C. erythraea var. var. var. var. var. Character centaurium subcapitatum capitatum subcapitatum capitatum pop. 5 pop. 11 pop. 30 pop. 29 pop. 29 X Ss x Ss x Ss X Ss X Ss Height (cm) 20°38 “35-6 4-4 1:0 1-4 0-2 oo) 275 ie 0-2 Habit (a) t(b) Tt (c)T | Oat aes © it 4°5 3 **S 1 4 § por? S Inflorescence M D M VD M VD M VD M VD Leaf L/B ratio 28 7:4 24 0:3 25 0-4 2:9 0-3 fs jee OP Leaf-shape ELL BLL ELL ELL ELL Indumentum G G G G G Calyx/corolla-tube ratio 0-55 0-05 0-61 0-06 1:0 =0-09 0-79 O-11 Le” 0-87, Corolla-tube length (mm) 8-7 0-5 8:6 0:6 4:6 0-4 7-3 1:0 4:8 0-7 Corolla-lobe length (mm) 49 0-9 5°7 0:4 4-7 0-6 4:8 0-7 4-5 0-4 Pollen grain diameter (um) 25-0 25:0 26:0 25-0 26:0 Stigma-shape a a a a a Capsule length (mm) 9-3 0:8 — — —- -—— 89 1-1 6:6 08 Filament length(mm) ~~ 3:3 0-2 FD POW SOs Ms 3-4 0-2 Sl 0-6 Point of insertion of stamens Mouth of tube Mouth of tube Base of Tube Mouth of tube Base of tube 1-3. Height, Habit, Inflorescence All three characters are extremely variable and this is to be expected since all are strongly affected by the environment. a) C. erythraea All plants of this species have 4—6 cauline nodes, and most have a compound inflorescence consisting of many branches each terminating in a compound cyme (more than 5 cymes), although some plants have only 1 or 2 cymes. Populations vary in the length of the cauline internodes, and in the relative laxity/compactness of the inflorescence, and can be divided into three major groups: i) Populations 1-7, 10, 12, 25, 26, 28, L and K are medium to tall (10-30 cm) with a single main flowering stem, 4-6 cauline nodes, long internodes with well-spaced leaves, and a dense inflorescence of more than 5 cymes, or sometimes only 1 or 2. Plants from populations 1-4, 10, 12, 26 and 28 from inland habitats, 5-7 and 25 from the bases of cliffs by the sea and L and K from pinewoods near sand-dunes, became taller but otherwise retained these characteristics when grown from seed under uniform conditions. Continental plants grown in the greenhouse resembled the British plants placed in this group. ii) Populations 8, 11, 13, 27, I and J, together with plants from 29 with normal stamen insertion, are all dwarf (4-8 cm) with a single main flowering stem, 4 or more cauline nodes, short inter- nodes with crowded cauline leaves, and a dense or very dense inflorescence of many (>5) or few (<3) cymes. Plants from populations 27, I and J from sand-dunes became taller and identical to plants in group (i) when cultivated under uniform conditions, whereas plants from populations 8, 11, 13 and 29 from exposed, maritime clifftops retained their original characteristics. iii) All plants from population 30 and those from population 29 with stamens inserted at the base of the corolla-tube, are even dwarfer (1-1—1-4 cm) with a single main flowering stem, 2-3 cauline nodes, very short internodes with crowded cauline leaves, and a very dense inflorescence of many cymes. These characteristics were retained in cultivation. iv) Population 23 from sand-dunes is dwarf (4-8 cm) with several main flowering stems (the plant is as broad as it is tall), 4 cauline nodes, short internodes with crowded, cauline leaves, and a dense inflorescence of many cymes. All characteristics were retained in cultivation. b) C. littorale 16 R. A. E. UBSDELL Plants of this species can be divided into three main groups which, apart from becoming taller, remained constant in cultivation: i) Populations A, D, F, G and H from sand-dunes on the Lancashire coast and N from Anglesey have one main fiowering stem, 3 or 4 cauline nodes (the leaves are longer than the internodes) and a lax inflorescence of few (<3) cymes. Continental plants grown in the greenhouse resembled the British plants belonging to this group. ii) Populations 22, 24, R, EL, DO and T from sand-dunes on the coasts of Northumberland and north-eastern Scotland have one main flowering stem, ft or 2 cauline nodes with long leaves exceeding the internodes in length, and a lax inflorescence of few (<3) cymes. iii) Population BA from sand-dunes on the coast of Anglesey has several main stems (the plant is as broad as it is tall), several crowded, cauline nodes, and a dense inflorescence of few (<3) cymes. It thus resembles group (iti) of C. erythraea. c) Intermediate plants These are plants from mixed populations on the coasts of Anglesey and Lancashire which are intermediate between C. erythraea and C. littorale in a number of characters and cannot be referred to either. These plants are tall (over 10 cm) with 3-6 well-spaced cauline nodes and an inter- mediate inflorescence of 3-5 cymes. Many of these characters seem to be adaptations to local environmental conditions and are of little use for separating the two species. 4,5. Length|breadth ratio and shape of cauline leaves These leaf characters have proved easy to score and are of great use for separating the two species. a) C. erythraea Plants of this species can be divided into two groups which remained constant in cultivation: i) The majority of plants have a mean cauline leaf length/breadth ratio falling within the range 2:1-3-3 and elliptical leaves. ii) Populations J and L have a higher mean leaf ratio with values of 3-5 and 4-7 respectively. These are the narrow-leaved variety with linear-elliptical leaves reported by previous authors from Norfolk and Suffolk. b) C. littorale Populations of this species show more variation, both within and between populations, although the leaf-shape is linear in all plants. Three main groups can be recognized and all remain constant in cultivation: i) Populations A, D, F, G and H from the Lancashire coast, 24 and R from Northumberland, 22 and EL from the coast of north-eastern Scotland and plants from the Continent grown in the greenhouse have a mean leaf ratio within the range 5-0-6°3. ii) Populations BA and N from the coast of Anglesey have a higher mean leaf ratio within the range 7:1-7-6. iii) Populations DO and T from the coast of north-eastern Scotland have a lower mean leaf ratio within the range 4-1-4°3. c) Intermediate plants As all mixed populations in the British Isles occur on the coasts of Lancashire and Anglesey, the value of C. littorale is taken to be above 5:0 and that for C. erythraea below 3:3. Plants of these two species from the Continent also fall within these values. The leaf ratios of the inter- mediate plants from the British Isles and Germany fall within the range 3-5—4:8 and the leaves are linear-elliptical. The mean leaf ratios of the mixed populations fall within the range 3-1-5-3. 6. Indumentum- a) C. erythraea All populations are glabrous, and this character is retained in cultivation. b) C. littorale Populations vary in this character and fall into three main groups which remain constant in cultivation. i) Populations A, D, F, G, H from the Lancashire coast and N from the coast of Anglesey corresponding to var. occidentale and population BA from Anglesey corresponding to var. bayleyi are densely scabrid. ii) Populations 22, 24, EL, DO, R and T from the east coast and corresponding to var. littorale and var. minor are either glabrous or semi-scabrid. CENTAURIUM ERYTHRAEA AND C. LITTORALE: MORPHOLOGY 17 iii) Plants from the Continent are completely glabrous. c) Intermediate plants As all plants of C. erythraea from single species populations are glabrous and all plants of C. littorale from single species populations on the coasts of Lancashire and Anglesey are scabrid, this character can be used to separate them in mixed populations from the British Isles. The intermediate plants vary in this character, as in some mixed populations all are glabrous while in the rest all intermediates are scabrid. Intermediate plants from mixed populations in Germany were glabrous as are all Continental plants of C. erythraea and C. littorale subsp. littorale. 7. Calyx/corolla-tube ratio This character proved easy to calculate and is one of the best for separating the two species. a) C. erythraea Populations of this species fall into three main groups: i) Most have a mean value within the range 0-44—0-59, but this increases in cultivation to 0-64— 0-71. ii) Populations 26, 27, 28 from Wales and western Ireland, plants of this species from mixed populations on the coasts of Lancashire and Anglesey, and population L from eastern Norfolk, have a slightly higher mean value, 0:63-0-64, which increases in cultivation to 0:74-0:76. iii) Population 30 and plants from 29 with stamens inserted at the base of the corolla-tube have a mean value of 1-0—-1-2, which is much higher than that of all other plants of this species. The rest of the plants from population 29 with normal stamen insertion also have a higher mean value of 0-79. Examination of the length of the calyx and corolla-tube shows that the calyx of these plants is of the same length as the calyx of typical C. erythraea, and the difference in ratio is the result of the very short corolla-tube of the plants from populations 29 and 30. b) C. littorale Populations of this species fall into three main groups: i) Most have a mean value within the range 0:80-0:87, but this increases in cultivation to 0-88-0-97. ii) Populations N and BA from Anglesey have an even higher value of 0:97-0:98. iii) Populations DO and T from Sutherland and Ross have a lower mean value of 0-50-0-65. This is atypical of the species and in fact falls within the range of C. erythraea. c) Intermediate plants Plants of C. littorale from the coasts of Lancashire and Anglesey can be clearly separated from plants of C. erythraea using this character. Intermediate plants from mixed populations in the British Isles vary with some falling within the range of C. littorale from the west coast, others falling within the range of C. erythraea, while the rest are intermediate and fall within the range 0:72-0:78. Mixed populations have mean values of 0:66-0:82. Intermediate plants from Germany have mean values of 0:66-0:73, which is intermediate between the values of Continental plants of C. erythraea and C. littorale. 8. Length of corolla lobes This character can only be accurately measured on living material. a) C. erythraea All populations of this species have mean values of 4:4 to 5-5 mm and remain constant in cultivation. b) C. littorale Populations of this species are more variable and fall into two main groups: i) Populations from the coasts of Lancashire and Anglesey, and plants from the Continent have mean values within the range 5-8-6:°6 mm. ii) Populations 24, R, 22, EL, DO and T from the east coast have lower mean values within the range 4-5-5-4 mm. c) Intermediate plants This character can be used to distinguish plants of C. littorale from the coasts of Lancashire and Anglesey from plants of C. erythraea. Intermediate plants from mixed populations have mean values within the range 5-4—6-1 mm. 9. Diameter of the pollen grains No mention of this character has been made by previous authors, yet it has proved easy to measure accurately and is of great value in separating C. erythraea from C. littorale. B 18 R. A. E. UBSDELL a) C. erythraea All plants of this species were found to have pollen grains ranging from 23 to 26 um with mean values for populations within the range 24-26 pm. b) C. littorale | All plants of this species were found to have larger pollen grains ranging from 28 to 32 pm with mean values for populations within the range 29-32 yum. c) Intermediate plants Although some of the intermediate plants from mixed populations have mostly sterile and dis- torted pollen, most have fertile, spherical pollen grains, like those of C. erythraea and C. littorale, ranging in diameter from 26 to 30 um and with population mean values from 27 to 28 um, which is intermediate between the two species. 10. Shape of the stigma No mention of this character has been made by other authors, yet it has proved easy to score and is of great value in separating living material of the two species. Stigmas of intermediate plants from mixed populations are clearly intermediate in shape between those of the two species. 11. Length of filaments Most plants of C. erythraea, and all of C. littorale and the intermediates have the filaments ranging in length from 3-0 to 3-8 mm (mean values for populations of 3-3—3-5 mm) and they are inserted at the mouth of the corolla-tube. All plants of C. erythraea from population 30 and some from 29 have the filaments ranging in length from 4-8 to 6-0 mm (mean values for populations of 5-0-5-1 mm) and they are inserted at the base of the corolla-tube. 12. Length of capsule Most plants of C. erythraea, and all of C. littorale and the intermediates have a similar range {mean values for populations of 8-9-9-3 mm) and the capsules are equal in length to the corolla- tubes. Plants of C. erythraea from populations 29 and 30 with the filaments inserted at the base of the | corolla-tube differ. These have shorter capsules (mean value 6-6 mm) which are longer than the stunted corolla-tubes. SUMMARY OF INDIVIDUAL CHARACTERS As mixed populations of the two species are found only on the coasts of Lancashire and Anglesey, only the variation shown by plants of C. /littorale from these areas needs to be considered when selecting characters to distinguish this species from C. erythraea in mixed populations from the British Isles. The above analysis of individual characters has shown that the best ones are those listed in Table 6. TABLE 6. CHARACTERS USED TO DISTINGUISH C. ERYTHRAEA FROM C. LITTORALE VAR. OCCIDENTALE AND VAR. BAYLEYI Character C. erythraea Intermediate C. littorale 1. Leaf length:breadth 2-1-3-3, short and broad 3-4—4:9, intermediate 5-0-7:6, long and narrow ratio 2. Leaf-shape elliptic, sides never linear-elliptic, sides linear with sides parallel, parallel, 5-veined, apex never parallel, 3-veined, 1-veined, apex obtuse acute apex acute 3. Indumentum glabrous semi-scabrid scabrid 4. Calyx:corolla-tube 0-40-0-64, calyx about 0:65-0:75, intermediate 0-76-0-98, calyx nearly ratio half as long as as long as corolla-tube corolla-tube 5. Corolla-lobe length 4-5-5-4 mm 5:5-5-6 mm 5:7-6:2 mm 6. Diameter of pollen grains 24-26 wm 27-28 um 29-32 «m 7. Stigma-shape! a b Cc Score for each 0 5 10 Total hybrid-index score 0 Intermediate 70 1 See page 11. CENTAURIUM ERYTHRAEA AND C. LITTORALE: MORPHOLOGY 19 AN ASSESSMENT OF THE TOTAL VARIATION SHOWN BY POPULATIONS To arrive at an assessment of the total variation shown by each plant, a total hybrid-index score was calculated using the characters and numerical scores given in Table 6. Thus, the total hybrid- index score for a plant with all C. erythraea characters would be 0 and for a plant with all C. littorale characters would be 70. Hybrid-index histograms were constructed to give an indication of the proportion of C. erythraea-like, C. littorale-like, and intermediate plants present in each population (Figs 1 & 2). SINGLE SPECIES POPULATIONS OF C. ERYTHRAEA Even in single species populations it can be seen that some plants will have a hybrid-index slightly higher than the theoretical value of 0 (Fig. 1: a—h; Fig. 2: i-m, v). Thus plants with a score of 0, 5, 10 and 15 can be considered to be characteristic of C. erythraea. | Plants from the East Anglian populations J (Suffolk) and L (Norfolk) have noticeably higher scores than all other plants of this species (Fig. 2: n & 0). In cultivation, plants of this species range from 0 to 15 (Fig. 2: s) except those from East Anglia, which fall within the range 15-235. SINGLE SPECIES POPULATIONS OF C. LITTORALE Populations of this species are far more variable in their hybrid-index scores than those of C. erythraea. a) Plants from the coasts of Lancashire and Anglesey have scores of 65—70 (Fig. 1: 1-5; Fig. 2: 7 & 8). b) Plants from the coasts of Northumberland, Easterness, Moray and those from the Continent grown in the greenhouse, have scores of 50-65, which are slightly lower than the theoretical value (Fig. 1: 6; Fig. 2:9 & 10, w). c) Plants from the coasts of E. Ross and E. Sutherland are very different from all other popula- tions of this species since, with values of 30-35 (Fig. 2: 12) and 40-45 (Fig. 2: 11), they fall within the range considered to be intermediate between all other plants of C. littorale and C. erythraea. All plants of this species except those from E. Ross and E. Sutherland, which unfortunately were not cultivated, showed scores of 55-70 in cultivation (Fig. 2: t). MIXED POPULATIONS The normal range of variation shown by the single species populations does not cause any overlap in the hybrid-index values of the two species, except for plants from E. Ross and E. Sutherland. This is in contrast to the situation shown by mixed populations from the coasts of Lancashire, Anglesey and Germany. a) Histograms i-vii (Fig. 1) for populations from Hightown, St Annes, Ainsdale, Freshfield and Anglesey all show a broken and uneven transition from C. erythraea to C. littorale. Obviously there are some plants that are referable to C. erythraea with scores of 0-15, and others with scores of 55-70 that are referable to C. littorale. The rest are intermediate in a number of characters, and with scores of 20-50 cannot be referred to either. b) Histograms viii (Fig. 1) and ix (Fig. 2) for other populations from Freshfield show that there are some plants referable to C. erythraea and the rest are intermediate. c) Histograms x and xi (Fig. 2), for other populations from Ainsdale and Freshfield respectively, show some plants referable to C. littorale while the rest are intermediate. d) Histogram xii (Fig. 2) for another population from Freshfield, and x and y (Fig. 2) for German populations show that only intermediate plants are present. The presence of plants in mixed populations with scores intermediate between those of C. erythraea and C. littorale is not unreasonable evidence of hybridization. It is possible that of these intermediate plants those with a score of 25-40 might belong to the F, generation, those with scores of 45-50 might be backcrosses to C. littorale, and those with scores of 15-20 might be backcrosses to C. erythraea. If this is so, it would seem that backcrossing and introgression to C. littorale has taken place in population 14 at Hightown (Fig. 1: i), popu- lations 16-19 at Freshfield (Fig. 1: viii; Fig. 2: ix, xi & xii), and in populations 21 and E at Ainsdale (Fig. 1: iti & vi). By contrast, populations 15 from Freshfield and 20 from St Annes (Fig. 1: ii & v) appear to consist of backcrosses to C. erythraea. 20 R. A. E. UBSDELL 1 ¢ ry > w w Percentage frequency LL TLL = f | < Vili oe ° w °o “ e © 5S 10 15 20 25 30 35 40 45 50 55 6065 76 () Hybrid index 0-70 Ficure 1. Hybrid index histograms. In populations B and C from Ainsdale (Fig. 1: vii; Fig. 2: x), and those from Anglesey (Fig. 1: iv) and Germany (Fig. 2: x & y) individual plants with scores of 20, 25, 35 and 40 are present Progeny of intermediate plants grown in cultivation In cultivation, the progeny of plants from single species populations together with plants of C. erythraea and C. littorale from mixed populations show hybrid-index values identical to those shown by them in the wild. However, the progeny of intermediate plants from Ainsdale, Freshfielp and Hightown show a variety of intermediate scores (Fig. 2: u). A few had values of 25-30 but the majority ranged from 40 to 50, thus resembling C. littorale more closely than C. erythraea and strongly suggesting backcrossing to the former species. THE CORRELATION OF DIFFERENT CHARACTERS WITHIN INDIVIDUAL PLANTS A more detailed picture of the characters of the individual plants, particularly those of the hybrids, can be seen from pictorialized scatter-diagrams. As space does not allow presentation of all popu- lations examined by this method, only a few representative examples are given. CENTAURIUM ERYTHRAEA AND C. LITTORALE: MORPHOLOGY 21 Percentage frequency Hybrid index 0-70 FiGure 2. Hybrid index histograms. (The unlabelled histogram is u.) SINGLE SPECIES POPULATIONS OF C. ERYTHRAEA Figs. 3 and 4 show the range of characters found in typical individuals of C. erythraea from popu- lations in Surrey (v.c. 17), E. Kent (v.c. 15), Dorset (v.c. 9), Haddington (v.c. 82) and Clare (v.c. H 9). All the characters fall within the range characteristic of C. erythraea subsp. erythraea. These studies show that all plants from single species populations of C. erythraea subsp. erythraea, except those from populations J (E. Suffolk), L CE. Norfolk), 29 and 30 (Isle of Wight), are very similar in most characters, and the only variation is shown by plants from exposed, coastal situa- tions. The three varieties described by Gilmour (1937) under C. umbellatum as var. centaurium, var. fasciculare and var. subcapitatum can be recognized, but, as Melderis (1972b) has shown that the correct name for this species is C. erythraea Rafn, these three varieties should be placed under C. erythraea subsp. erythraea as follows: i) C. erythraea subsp. erythraea var. erythraea C. umbellatum Gilib. var. centaurium sensu Gilmour, Kew Bull., 19: 497 (1937) Plants of this variety, the type variety of the species (Melderis 1972b), are tall (9-30 cm high) with a single main stem, 4—6 cauline nodes, long internodes with well-spaced leaves and a dense 22 R. A. E. UBSDELL 8.0, 7.0 © ra bi ¥ B 6 * hs < + 4 by fa o. ny ay 9 7 g Se ey allie! Fe Wrietew tit oe ie > i 7 hat phate eel + corte done ae | Teen Ant. /iterale Tram ie eee 7 ’ ; i prery by ’ , . pr , “ ye Y. : mils ae. Oe): aie cen’ eaferahie ta 0 ee ; $ Vee at Uh LORE a Pa eres La a ee ire es ae ae oe ie» hy ) 7 a od s > as > ‘ Sy aay pair sate thee UR See bre oar ‘aie Bi hit yt t¢ Lon Sid onialy hackerophce th 2. cryeeaee ae PRN CRE EPG: 54 qt f ear) ; sf =f is re reysey a Fy any aie pes ee mi ” 7 Sea % . sa } ; 4 fo PLETE ; ® f . % et EP a ee Fee ih i. b diay bh Ns ty me Ged ai asl y j Aub Pye" * (ey mie Vere ef katie i) Pooissy yy ap ine) wood fe tyra th tee AON aS Se er Ae ite oe Resting. me er eres eas | fede hid Wee ' raves ted by sian 9 4 eA CL NS OLE OF Se ees haeniie rae ie Bae RF CS OLS OD ida: HES TREN. a Ne | bets ‘ "ee y ; phy vg bute: Sg Teh is oh ‘abe Naletuliel 1the chia, aa ‘ un : « 5 ; em gets are } ee Ae - Cia eet, Sa High . ; hy ay oc ae Ty ame NEL, ATs shi SA aah? rs earner oe. WSuisskr. en wie eee bigs ror t aisha: » . im MAR aT ; : ; ay ‘ een se Ae He) Cpveiilwn, Sia ene eRe Ne whereas: ..P ETP) oe ae Osea ati ae fae ER alee, sex ARN RES | | | ae hy ‘ y ; = of i ! 7 ¥ i } nl i 7 ; ay Mars bah Watsonia, 11, 33-43 (1976) 33 Studies on variation and evolution in Centaurium erythraea Rain and C. littorale (D. Turner) Gilmour in the British Isles 2. Cytology R. A. E. UBSDELL Department of Botany, University of Reading* ABSTRACT Chromosome counts of Centaurium erythraea Rafn and C. littorale (D. Turner) Gilmour from northern Europe are given. They confirm those made by Zeltner (1970) and show both species to be tetraploid (2n = 40) with regular meiosis. Morphologically intermediate (F,-like) plants from Ainsdale, S. Lancs. (v.c. 59), were found to be tetraploid with irregular meiosis. They were almost sterile, as were F,-like plants from Newborough, Anglesey (v.c. 52), and were almost identical in their morphology and cytology to artificial F, hybrids. Morphologically intermediate plants from northern Germany were found to be tetraploid, but with regular meiosis and a high fertility. It is suggested that they are stabilized derivatives of the F, hybrid. Hybrid plants from St Annes, W. Lancs. (v.c. 60), which resemble C. erythraea in their morphology and are probably backcrosses to this species, were found to be tetraploid with regular meiosis and a high fertility. Hybrid plants from Ainsdale, Freshfield and Hightown, S. Lancs. (v.c. 59), which closely resemble C. littorale in their morphology, were found to be hexaploid (27 = 60) with regular meiosis and a high fertility. They are shown to breed true, to be isolated from their tetraploid parents and other (tetraploid) hybrids by a difference in chromosome number, and to be able to compete successfully with their parents. They are, therefore, considered to constitute a new species, and a possible origin involving hybridization and polyploidy is discussed. INTRODUCTION Prior to Zeltner’s work (1961, 1962, 1963, 1966, 1967) only four species of the genus Centaurium Hill had been examined cytologically, and the results obtained by different authors contradicted each other so much that it was impossible even to establish a basic number for the genus (Wulff 1937, Warburg 1939, Rork 1949, Mesquita Rodrigues 1953, Khoshoo & Khushu 1966, Brink 1967). Zeltner made numerous, reliable chromosome counts of all the European species and finally produced a comprehensive treatment summarizing his work on the genus (Zeltner 1970). He found all the karyotypes to be very similar and decided that they were of no taxonomic value. He also established two basic chromosome numbers for the European species, of x = 11 for C. spicatum (L.) Fritsch and x = 10 for all the other species. Six of the latter were found to be diploid (2n = 20), two tetraploid (2n = 40) and the other three species were found to have both diploid and tetraploid races, the diploids occurring in the Mediterranean region and the tetraploids in central and northern Europe. His work showed the importance of polyploidy, and of the resulting genetical isolation, in the evolution of the genus. CENTAURIUM ERYTHRAEA RAFN The first chromosome count (2n = 42) for this species was determined by Rork (1949) on a plant from New York. Zeltner (1962) carried out chromosome counts on 113 populations from Europe. * Present address: Botany School, University of Oxford. Cc 34 R. A. E. UBSDELL Of these, 16 from southern Europe were found to be diploid (2n = 20) with regular meiosis, while the other 97, all from northern Europe, were found to be tetraploid (2n = 40) with regular meiosis. It is not known how the tetraploids arose, but Zeltner suggested that it could have been by auto- polyploidy. C. capitatum (Willd.) Borbas, which is now considered to be only a variety of C. erythraea (Melderis 1972, Ubsdell 1976), was also found by Zeltner to be tetraploid with regular meiosis. CENTAURIUM LITTORALE (D. TURNER) GILMOUR The first chromosome count (n = 19) for this species was determined by Wulff (1937) on material from northern Germany. Warburg (1939) recorded 2” = 38 and 2” = 56 on material from the British Isles, while Brink (1967) reported that most plants of this species from the Netherlands had 2n = 40 although she had found a few plants with 2n = 38 or 2n = 42. Zeltner (1962) ex- amined 15 populations from Europe and similarly found all to be tetraploid with regular meiosis. The diploid relatives of this species are confined to the Iberian peninsula and southern France, and Zeltner suggested that C. /ittorale could have been derived from them either by autopolyploidy or by allopolyploidy. NATURALLY OCCURRING HYBRIDS Zeltner (1970) examined some hybrid plants (C. erythraea x C. littorale) from Ainsdale, S. Lancs. (v.c. 59). He found 40 chromosomes at mitosis, but observations of meiosis showed some of the cells dividing regularly to give two equal groups of 20 chromosomes, and others showing irregular division with groups of 20/24 and 16/24. 80% of the pollen was distorted and sterile. K. Jakobsen (in litt. 1972) reported that tetraploid hybrids between these two species occur on the coasts of Denmark. Some of the hybrids were almost sterile with irregular meiosis, while others appeared to be cytologically stable. This paper is concerned with the cytology and pollen fertility of plants of C. erythraea and C. littorale from northern Europe (chiefly from the British Isles), their naturally occurring hybrids (Ubsdell 1976) and artificial hybrids. METHODS CHROMOSOME COUNTS Somatic chromosomes were examined at mitosis in the root-tips. Initially they were difficult to count as all the chromosomes tended to conglomerate on the metaphase plate. Eventually, however, pre-treatment with saturated, aqueous paradichlorobenzene for twelve hours before fixation in 3:1 ethanol:acetic acid was found to give satisfactory preparations with the chromosomes well spread. After fixation the root-tips were softened in N HCl for three minutes before washing and staining in propionic-orcein. Buds were fixed in 3:1 ethanol:acetic acid and stored in 70% ethanol. After softening in 1:1 ethanol:concentrated HCl, good results were obtained for the meiotic chromosomes by staining in propionic-orcein; acetic-orcein and acetocarmine gave poor results. All preparations were made permanent by freezing with Arcton 12, dehydrating in ethanol and mounting in Euparal. POLLEN FERTILITY Fresh pollen grains were mounted in cotton blue/lactophenol. Spherical pollen grains took up the stain and were considered to be viable, while unstained, distorted, elliptical grains were considered to be infertile. 100 pollen grains were scored for each plant. RESULTS Details of the chromosome number, pairing at meiosis and pollen fertility are given for various collections of C. erythraea (Table 1), C. littorale (Table 2), and their natural (Table 3) and artificial (Table 4) hybrids. CENTAURIUM ERYTHRAEA AND C. LITTORALE: CYTOLOGY TABLE 1. CHROMOSOME NUMBER, MEIOTIC PAIRING AND POLLEN FERTILITY OF C. ERYTHRAEA Single species populations 4. Orpington, W. Kent, v.c. 16 5. Folkestone, E. Kent, v.c. 15 6. Luccombe, Isle of Wight, v.c. 10 7. Sandown, Isle of Wight, v.c. 10 8. Swanage, Dorset, v.c. 9 10. Bonchurch, Isle of Wight, v.c. 10 11. Freshwater, Isle of Wight, v.c. 10 12. Steyning, W. Sussex, v.c. 13 23. Aberlady, Haddington, v.c. 82 25. Newbiggin, S. Northumberland, v.c. 67 27. Fanore, Clare, v.c. H9 28. Funshin, S.E. Galway, v.c. H1I5 J. Minsmere, E. Suffolk, v.c. 25 L. Minsmere, E. Suffolk, v.c. 25 Mixed populations 14. Hightown, S. Lancs., v.c. 59 15. Freshfield, S. Lancs., v.c. 59 20. St Annes, W. Lancs., v.c. 60 21. Ainsdale, S. Lancs., v.c. 59 AN. Newborough, Anglesey, v.c. 52 Continental plants raised in greenhouse (all seed obtained from Botanic Gardens) Rigen, Germany Wageningen, Netherlands Copenhagen, Denmark TABLE 2. CHROMOSOME NUMBER, MEIOTIC PAIRING AND POLLEN Meiosis metaphase I (No. of bivalents) anaphase I 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 ales FERTILITY OF C. LITTORALE Single species populations D. _ Aiunsdale, S. Lancs., v.c. 59 G. Aijinsdale, S. Lancs., v.c. 59 H. Aijnsdale, S. Lancs., v.c. 59 BA. Newborough, Anglesey, v.c. 52 N. Newborough, Anglesey, v.c. 52 24. Holy Island, Cheviot, v.c. 68 22. Nairn, v.c. 96b Mixed populations E. Ainsdale, S. Lancs., v.c. 59 21. Ainsdale, S. Lancs., v.c. 59 14. Hightown, S. Lancs., v.c. 59 16. Freshfield, S. Lancs., v.c. 59 20. St Annes, W. Lancs., v.c. 60 AN. Newborough, Anglesey, WiCw 2 Continental plants raised in the greenhouse. (all seed obtained from Botanic Gardens) Rigen, Germany Blid6, Sweden Helsinki, Finland Copenhagen, Denmark Meiosis metaphase I (No. of bivalents) anaphase I 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 Root-tip mitosis 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 Root-tip mitosis 2n = 40 2n = 40 2n = 40 2n = 40 2 A0 2n = 40 2n = 40 2n = 40 2n = 40 2n = 40 Pollen fertility Pollen fertility 35 36 R. A. E. UBSDELL TABLE 3. CHROMOSOME NUMBER, MEIOTIC PAIRING AND POLLEN FERTILITY OF NATURAL HYBRIDS Melosis 14. 16. 20. 21, AN. Newborough, Anglesey Cultivation code and status Hightown, S. Lancs. Progeny in cultivation R60 Freshfield, S. Lancs. Progeny in cultivation R61 St Annes, W. Lancs. Ainsdale, S. Lancs. Ainsdale, S. Lancs. Progeny in cultivation R52 Selfed progeny of R10, R11 Ainsdale, S. Lancs. Progeny in cultivation R23 R50 Ainsdale, S. Lancs. Progeny in cultivation R28 FAL. Falshoft, Germany S.P. St Peter, Germany metaphase I (No. of bivalents, univalents and quadrivalents) 30 Il 30 II 30 IL 30 I 20 IL 30 I 30 I 29/30 I 30 I 30 I 30 Il 30 Il 28 I1+11V 26 If+4 1 18 +41 18 Ii+11V 1ISTI+11V+61 141I+11V+81I IZ 1+1IV+61 and as for R23 20 II 20 II anaphase I (pattern of Segregation and diploid no.) 30-30 (2n = 60) 31-26 (2n = 57) 29-28 (2n = 57) 31-25 (2n = 56) 26-25 (2n = 51) 26-24 (2n = 50) 30-30 (2n = 60) 28-28°(2n = 56) 27-27 (2n = 54) 28-27 (2n = 55) 29-29 (2n = 58) 27-25 (2n = 52) 30-29 (2n = 59) and those given for 14 30-30 (2n = 60) 20-20 (2n = 40) an ='40 2n = 60 30-30 (2n = 60) 29-29 (2n = 58) 30-30 (2n = 60) 30-30 (2n = 60) 30-30 (2n = 60) 30-30 (2n = 60) 30-30 (2n = 60) 2n:= 49 on == AO 31-29 (2n = 60) 31-27 (2n = 58) 30-25 (2n = 55) 30-24 (2n = 54) 2n = 40 2n = 40 20-20 (2n = 40) 20-20 (2n = 40) Root-tip mitosis 2n = 60 2n = 60 2n = 60 2n = 60 2n = 60 2n = 60 2n = 60 2n = 60 2n = 40 2n = 60 2n = 40 Pollen fertility 45, 58, 63, 69, 70, 80, 81, 84% 81-90% 76, 82, 84, 88, 92, 94, 96% 85-90% 67-87% 0-10% 0-10% 0-10% 77-85% 78-82% CENTAURIUM ERYTHRAEA AND C. LITTORALE: CYTOLOGY 37 TABLE 4. CHROMOSOME NUMBER, MEIOTIC PAIRING, POLLEN FERTILITY AND SEED SET OF ARTIFICIAL HYBRIDS x seed Cultivation set per code and Meiosis Root-tip Pollen capsule status metaphase I anaphase I mitosis fertility by open pollination Fi 1 20 I 24-20 (27 = 44) 2n=40 0-10% 0-6 2 1914+21 23-20 (2n = 43) 18 +41 22-19 (2n = 41) 3 17 11+61 20-20 (27 = 40) 18 U+1I1V 20-18 (2n = 38) 4 17 11+11V+21 19-19 (2n = 38) 1611+11V+41 19-17 (2n = 36) 15 11+21V+2I 20-15 (2n = 35) 1411+21V+41 15 Il+ch. of 5+51 Tetraploid hybrid R23 x C. erythraea 5 20 IL 20-20 (2n = 40) 2n=40 67% 110 18 1+11V 15 11+21V+21 Tetraploid hybrid R23 x C. littorale 6 Not clear but IIs, ls and 20—-1-1-17 (2n = 40) 2n=40 6-9% 2 IVs 18—2-2-18 Hexaploid hybrid x C. erythraea 7 24 11+2 1 28-22 (2n = 50) 2n = 50 26-54% 50 8 22 Ti+2 Ill 26-24 (2n = 50) minute (reciprocal crosses) 19 II+chs. of 4, 6 25—5-20 (2n = 50) malformed 26-22+ 1 If (Qn = 50) 21-21+4 II (2n = 50) Hexaploid hybrid x C. littorale 9 20 Il+chs. of 3, 3, 4 25-25 (2n = 50) 2 = 00) “3-25 7, 10 10 20 II+chs. of 4,4+2 I 25—1-24 (2n = 50) minute (reciprocal 19 II+chs. of 4,6+2 1 25-2-23 (2n = 50) malformed crosses) 19 Il+chs. of 6+6 I 24—8-18 (2n = 50) 18 IL+chs. of 4,6+2 1 20-15~—15 (2n = 50) 18 Il+rgs. of 4,44+4 1 20-3-27 (2n = 50) 17 If+chs. of 3, 3, 4, 6 20-30 (2n = 50) 17 I1+chs. of 3, 3, 4,4+21 16 If+chs. of 4, 4, 4, 6 16 I1+chs. of 3, 4,6+4 1 16 Il+rgs. of 4,4+41 16 I1+chs. of 3, 3, 5, 5 Hexaploid hybrid x Hexaploid hybrid 11 30 IL 30-30 (2n = 60) 2n = 60 79-89% 200 CENTAURIUM ERYTHRAEA All plants of this species, both from single species populations and from mixed ones, were found to be tetraploid (2n = 40) with regular meiosis. All plants were highly fertile, with a pollen fertility of over 80% in single species and mixed populations (Table 1). 38 R. A. E. UBSDELL CENTAURIUM LITTORALE All plants of this species, both from single species populations and from mixed ones, were also found to be tetraploid (2n = 40) with regular meiosis. All plants were highly fertile, with a pollen fertility of over 79% in single species and mixed populations (Table 2). NATURALLY OCCURRING HYBRIDS These hybrids showed a variety of chromosome numbers and meiotic irregularities, and seemed to fall into three main cytological types. 1. This is represented by the morphologically intermediate, F;-like plants from Ainsdale, S. Lancs. (Table 3: populations C, E & 21). These plants were tetraploid (2n = 40) with irregular meiosis. No clear metaphase counts were made, but preparations of first anaphase showed lagging chromosomes, fragments and apparent chromatid bridges. All plants had a very low pollen fertility of 0-10%. | é Seed collected from these hybrids in the field was sown in the greenhouse and the progeny raised to maturity. These cultivated plants, which were at least second generation hybrids (Table 3: R23 & R28), were also found to be tetraploid (2” = 40) and to have irregular meiosis. The number of bivalents varied from 13 to 20, the other chromosomes forming univalents or occasionally a quadrivalent (Fig. 1: a & b). There were lagging chromosomes and other irregularities at first anaphase, and despite the large number of bivalents all plants had a low pollen fertility (0-10 %). Morphologically intermediate, F,-like plants from Newborough, Anglesey (v.c. 52) (Table 3: population AN), also had a low pollen fertility, but unfortunately no chromosome counts were made. , age he Me a U4 Fup. rs FicureE 1. Tetraploid hybrids. Progeny of morphologically intermediate, F,-like hybrids from Lancashire (a) R28 metaphase I of meiosis (18 IL[+1 IV), (b) R23 metaphase I of meiosis (17 I+ 6 I). Artificial F, hybrid (c) metaphase I of meiosis (15 I[+5 I and a chain of 5). 2. This is represented by plants from St Annes, W. Lancs. (v.c. 60). Table 3, population 20), which closely resemble C. erythraea in their morphology and may be backcrosses to this species, and also by the morphologically intermediate plants from the northern German populations (Table 3: populations FAL. & S.P.). These plants were found to be tetraploid (2n = 40) with regular meiosis, and all were highly fertile with a pollen fertility comparable to that of the parents. Unfortunately, no progeny were raised in the greenhouse. 3. This is represented by the majority of hybrid plants from Ainsdale, Freshfield and Hightown, S. Lancs. (Table 3: populations 14, 16 & 21), which closely resemble C. littorale in their morpho- logy and may possibly be backcrosses to this species. These plants were found to be basically CENTAURIUM ERYTHRAEA AND C. LITTORALE: CYTOLOGY 39 hexaploid (2n = 60), but meiotic counts of 50, 51, 52, 54, 55, 56, 57, 58 and 59 were also made, more than one number often being counted from the same individual. Three preparations of first metaphase showed complete pairing with 30 bivalents, but many other preparations were not clear enough for a count to be made. Twenty preparations of first anaphase showed cells dividing regularly into two equal groups of from 27 to 30 chromosomes, while a further ten preparations showed cells dividing irregularly into groups of 24/26, 25/26, 25/31, 27/28, 26/31, 28/29 chromosomes (Fig. 2). These plants showed a wide range of pollen fertility (45-96%), but the majority were quite fertile (over 80%). (a) (2n= 50) (b) (2n=31) ee @ ® ® a2 ue fsa @ e ° PY vie a © D “of 9eree? >. * yA 7 ‘et ate e, * on M0, ee ow % Fy a’ ace @@? (c) (30 — 30) FiGureE 2. Hexaploid hybrids (a) anaphase I of meiosis (2n = 50), (b) anaphase I of meiosis (31-26, 2n = 57), (c) anaphase I of meiosis (30-30, 2” = 60). Wiild-collected seed from these hybrids was sown in the greenhouse and the progeny raised to maturity. All plants were found to be basically hexaploid (counts of 2n = 60 for root-tip chromo- somes). A few plants (Table 3: R50) showed irregular meiosis, the number of bivalents varying from 26 to 29 with the other chromosomes forming univalents or occasionally a quadrivalent. Anaphase segregation was unequal, and the pollen fertility of these plants fell within the range 67-87 %. However, the majority of plants (Table 3: R10, R11, R31, R51, R52, R60, R61) showed regular meiosis with complete pairing (30 bivalents) and equal anaphase segregation into two groups of 30 chromosomes. The pollen fertility of these cytologically stable, hexaploid plants was high (74-90%) and comparable to that of the parents. : Seed set by selfing these hexaploid hybrids in the greenhouse produced progeny identical to the parents in their morphology, cytology and fertility (Fig. 3, R10). ARTIFICIAL Fj HYBRIDS AND THEIR PROGENY These hybrids resulted from crosses made in the greenhouse between C. erythraea and C. littorale- All were found to be morphologically intermediate between the parents (Fig. 3: 1-4), and tetraploid (2n = 40) with irregular meiosis (Table 4: 1-4). The number of bivalents varied from 14 to 20, the other chromosomes forming univalents and multivalents (Fig. 1c). There were also lagging chromosomes and other irregularities at first anaphase. Despite the large number of bivalents, pollen fertility was low (0-10%) and all plants set very little seed by artificial self-pollination. 40 R. A. E. UBSDELL LEAF LENGTH : BREADTH RATIO 0:5 0-6 0-7 0-38 0-9 CALYX : COROLLA TUBE RATIO KEY Leaf shape Indumentum Pollen (diameter) e elliptical e # « glabrous @ua 24-26Uu. w linear-elliptical e@ # « semi-scabrid @e 4 27-28U. a linear @-s- « SCabrid @8#4 29-327u. @ C.erythraea FF, * C. littorale | . (1, 2, 3, 4) © R 23 (F3) ® R 10 (Hexaploid) ® 5 (Backcross) F 6 (Backcross) oO 7,8 (Backcross) 9,10 (Backcross) @ 1i (Hexaploid x hexaploid) FiGuRE 3. Scatter diagram comparing all artificial hybrids with plants of C. erythraea, C. littorale and their natural hybrids raised in the same greenhouse in the same year. OTHER ARTIFICIAL HYBRIDS Crosses between natural, almost sterile, tetraploid hybrids (R23) and natural, hexaploid hybrids produced pentaploid (2n = 50) progeny with irregular meiosis and a very low pollen fertility and seed set. No such plants were found in the wild. Using the same natural, tetraploid hybrid (R23), backcrosses were made to C. erythraea and C. littorale. Surprisingly the two backcrosses had very different fertility. That to C. littorale was morphologically intermediate between the tetraploid and hexaploid hybrids, had irregular meiosis, a low pollen fertility and seed-set (Table 4: 6), and was unlike any plant found in the wild. By contrast, the backcrosses to C. erythraea closely resembled that species and had a more regular meiosis, higher pollen fertility and better seed set than both the natural and artificial F, hybrids CENTAURIUM ERYTHRAEA AND C. LITTORALE: CYTOLOGY 41 (Table 4: 5). These plants strongly resembled the fertile, tetraploid hybrids (Table 3: population 20) from St Annes, W. Lancs. in their morphology, cytology and fertility. Backcrossing the natural, hexaploid hybrids (R10, R11) to C. erythraea produced progeny intermediate in morphology between the F, hybrid and C. erythraea (Fig. 3: 7 & 8), but the back- crosses to C. littorale closely resembled the hexaploid parents (Fig. 3: 9 & 10). Both backcrosses were pentaploid (2” = 50) with irregular meiosis, a low pollen fertility and poor seed-set (Table 4: 7-10). No plants with these characteristics were found in the wild. DISCUSSION Chromosome counts of C. erythraea and C. littorale from northern Europe show both species to be tetraploid (2n = 40) with regular meiosis. These results confirm those of Zeltner (1970) and Brink (1967), but contradict those of Wulff (1937), Warburg (1939) and Rork (1949). It is possible that faulty observation of this cytologically difficult material might explain the counts of 2n = 42 for C. erythraea made by Rork (1949), and 2” = 38 for C. littorale made by Wulff (1937) and Warburg (1939). It also seems probable that the count of 2” = 56 for C. littorale by Warburg (1939) was made from the hexaploid hybrid between this species and C. erythraea, since the hexa- ploid hybrids closely resemble C. littorale and are easily mistaken for this species. The artificial F; hybrids are shown to be tetraploid (2n = 40) with fairly good pairing at first metaphase, but irregularities at first anaphase and a low pollen fertility. It therefore seems likely that the morphologically intermediate (Hybrid-index 25-40 (Ubsdell 1976)), tetraploid plants from Ainsdale, S. Lancs., and Newborough, Anglesey, with irregular meiosis and low pollen fertility also belong to the F; generation, as do those examined by Zeltner (1970). The morphologically intermediate, tetraploid plants (Hybrid-index 25-40) from northern Germany with regular meiosis and a high pollen fertility seem unlikely to belong to the F; genera- tion, since artificial Fy hybrids produced by crossing plants of C. erythraea and C. littorale from northern Germany were found to have irregular meiosis and a low pollen fertility similar to those produced by crossing British plants. It is possible that they are segregates of the F; hybrid which have become cytologically stable, and a similar suggestion was made by Jakobsen (in litt. 1972) to explain the origin of cytologically stable, fertile, tetraploid hybrids found in populations on the coast of Denmark. The highly fertile, hexaploid, hybrid plants from Ainsdale, Freshfield and Hightown, S. Lancs. (Hybrid-index 45-50) could have been formed by one of several methods. While it is possible that it may have been derived from an octoploid parent and a normal, tetraploid parent, this seems very unlikely since the number 2” = 80 has never been recorded in Centaurium. It seems more likely that the hexaploid was produced from two tetraploid parents, C. littorale and C. erythraea, and this must almost certainly have involved an unreduced gamete, possibly as a result of non-disjunction and restitution during first meiotic division. It is possible that the unreduced gamete was from C. littorale since the hexaploids most closely resemble this species. With three sets of chromosomes, two of which would be from C. littorale, they would also have at least 20 bivalents and it is possible that some of the C. erythraea chromo- somes might also pair. Those that do not pair might get lost during anaphase segregation and this would explain the numbers of 2n = 50-59 observed at anaphase in some of the natural, hexaploid hybrids. This theory, however, assumes that non-reduction has only occurred in C. littorale since no hexaploid hybrids resembling C. erythraea have been found, although it is possible that they might be formed but do not survive in the wild. It seems more likely that non-reduction occurred in a hybrid plant, since natural and artificial, tetraploid F; hybrids are almost sterile owing to meiotic irregularities. Furthermore, it has been shown that neither selfing the artificial F; hybrids nor crossing between them increases their fertility so there would be a strong selective advantage for genotypes that could overcome their sterility by some other means. It has been shown that artificially backcrossing the natural, almost sterile, tetraploid hybrids from Ainsdale to C. erythraea produces tetraploid plants closely resembling this species in their morphology, and with regular meiosis and high fertility. Such plants closely resemble the fertile, 42 R. A. E. UBSDELL tetraploid hybrids from St Annes, W. Lancs. (Hybrid-index 15-20), and so these plants may have originated in this way. By this method, the fertility of the hybrids has been increased at the tetra- ploid level. However, it has also been shown that artificially backcrossing the same Ainsdale hybrids to C. littorale does not increase the fertility at the tetraploid level, all such backcrosses being almost sterile. If, therefore, fertilization occurred between a non-reduced, tetraploid hybrid gamete and a normal, reduced, diploid gamete of C. littorale, the more fertile hexaploid genotypes might be at a selective advantage. By this method, the hexaploids would resemble C. /ittorale more closely in their morphology and pairing at metaphase and anaphase segregation would also be the same as it would have been if non-reduction had occurred in C. Jittorale. Experiments have also shown that crossing the hexaploid hybrids with either of the tetraploid parents, or with the tetraploid hybrids, produces a few almost sterile, pentaploid plants of varying morphology, none of which has been found in the wild. It seems, therefore, that the hexaploids are isolated from their tetraploid parents and all tetraploid hybrids by a difference in chromosome number, and so in order to persist must either self or cross with each other. As has already been shown, the hexaploids are characterized by a more or less regular meiosis and good seed-set; they also breed true. Since these hexaploids are as abundant as their parents in certain populations at Ainsdale, Freshfield and Hightown, S. Lancs., and since they breed true and are also isolated from their parents by a difference in chromosome number, they should be considered to constitute a new species. Their nomenclature will be discussed in the third and final paper in this series. This study has consequently shown that, in addition to introgression, which might have been expected, hybridization between C. erythraea and C. littorale has resulted, in certain areas, in the evolution of a new species. ACKNOWLEDGMENTS I wish to express my sincere thanks to Dr D. M. Moore under whose supervision this work was initiated and completed. I am also very grateful to Professor V. H. Heywood for providing facilities in the Department of Botany at the University of Reading. This work forms part of a thesis accepted for the degree of Ph.D. at the University of Reading, and was carried out during the tenure of a grant awarded by the Science Research Council. REFERENCES BRINK, A. (1967). In FREUSEN, A. H. J. A field study on the ecology of Centaurium vulgare Rafn. Thesis, University of Utrecht. KHosHoo, T. N. & KHUSHU, C. L. (1966). Biosystematics of Indian plants, 2. The problem of Centaurium pulchellum. Proc. Ind. Acad. Sci. (B), 63: 152-160. MELpeRIs, A. (1972). Taxonomic studies on the European species of the genus Centaurium Hill. Bot. J. Linn. Soc., 65: 224-250. RODRIGUES, J. E. DE (1953). Contribuicao para o conhecimento cariologico das halofitas e psamOfitas litorais. Diss. Dout. Coimbra Univ. RorkK, C. (1949). Cytological studies in the Gentianaceae. Amer. J. Bot., 36: 687-701. UBSDELL, R. A. E. (1976). Studies on variation and evolution in Centaurium erythraea Rafn and C. littorale (D. Turner) Gilmour in the British Isles, 1. Taxonomy and biometrical studies, Watsonia, 11: 7-31. WarsurG, E. F. (1939). In MAuDE, P. F. The Merton Catalogue: A list of the chromosome numerals of species of British flowering plants. New Phytol., 38: 14. Wutrr, H. D. (1937). Karyologische Untersuchungen an der Halophyten-Flora Schleswig-Holstein. Jahrb. Wiss. Bot., 84: 820. ZELTNER, L. (1961). Contribution a'l’étude cytologique des genres Blackstonia Huds. et Centaurium Hill (Gentianacées). Bull. Soc. bot. Suisse, 71: 17-24. ZELTNER, L. (1962). Deuxiéme contribution a l’étude cytologique des genres Blackstonia Huds. et Centaurium Hill (Gentianacées). Bull. Soc. neuchdtel. Sci. nat., 85: 83-95. ZELTNER, L. (1963). Recherches sur quelques taxa méditerranéens du genre Centaurium Hill (Gentianacées). Bull. Soc. neuchdtel. Sci. nat., 86: 93-100. CENTAURIUM ERYTHRAEA AND C. LITTORALE: CYTOLOGY 43 ZELTNER, L. (1966). Sur quelques taxa nouveaux ou méconnus des genres Blackstonia Huds. et Centaurium Hill (Gentianacées). Bull. Soc. neuchdtel. Sci. nat., 89: 61-73. ZELTNER, L. (1967). Etude cytologique de deux espéces relativement rares du genre Centaurium Hill (Gen- tianacées). Bull. Soc. neuchdtel. Sci. nat., 90: 241-246. ZELTNER, L. (1970). Recherches de biosystématique sur les genres Blackstonia Huds. et Centaurium Hill (Gentianacées). Bull. Soc. neuchdatel. Sci. nat., 93: 1-164. (Accepted April 1975) é Rises lov 9%: os job ab aupigolo} 3 ee Pe. cs engl | ue ‘t Watsonia, 11, 45-52 (1976) 45 Notes on British Rubi, 4 Ei S. EDEES 23 Dartmouth Avenue, Newcastle, Staffs. ABSTRACT The names of seven brambles on the British list are discussed and the following new names are proposed: Rubus painteri for R. eifeliensis sensu W. C. R. Wats. ex parte, non Wirtg., R. turritus for R. apricus sensu W.C. R. Wats. ex parte, non Wimm., R. infestior for R. infestus sensu W. C. R. Wats., non Weihe ex Boenn., and R. anglohirtus for R. hirtus sensu W.C.R. Wats. ex parte, non Waldst. & Kit. R. pampinosus Lees is shown to be an earlier name for R. favonii W. C. R. Wats. and R. infestus Weihe ex Boenn. for R. taeniarum Lindeb. R. condensatus P. J. Muell. is confirmed as a British species. 1. Rubus pampinosus Lees, Bot. Malv. Hills, ed. 2, 55 (1852) R. favonii W. C. R. Wats., J. Ecol., 33: 338 (1946) R. buttii Barton & Riddelsd. in Riddelsd., Hedley & Price, F/. Gloucester, 148 (1948) There is a specimen in Babington’s herbarium (no. 233) in CGE named R. pampinosus Lees, which is undoubtedly the same taxon as R. favonii W. C. R. Wats. It is a good specimen mounted on two sheets, one bearing a panicle in advanced fruit and the other a piece of the first-year stem with two leaves. It was collected by W. Mathews from ‘open hedge banks in the Forest of Wyre’, Worcester, v.c. 37, 25/9/1851, and named by E. Lees. Babington had determined it R. villicaulis Weihe & Nees and maintained that opinion in later years (1869, p. 144). In his original description Lees (1852) described the stem as ‘polished’ and referred to the long, leafy panicle with wide- spreading branches. He called it ‘a very remarkable bramble, with leaves so large and numerous as almost to conceal the stem’ and said it grew in dense thickets in Cowleigh Park, v.c. 37. Later on (1867) he gave the Wyre Forest as a second locality. I have seen this bramble both in Cowleigh Park and in the Wyre Forest and have several specimens, including one from the Wyre Forest, Salop, v.c. 40, collected by W. C. R. Watson, 15/7/1953. This specimen is unnamed but is no doubt the source of Watson’s record (1958) of R. favonii for the Wyre Forest. R. buttii Barton & Riddelsd. is the same taxon. R. favonii must now be known as R. pampinosus. It is locally common in woods in the west of England from Staffordshire through Gloucestershire to Dorset. Exsiccata from the following vice- counties have been examined by A. Newton or myself: 6, 9, 11, 33, 36-40, 46. 2. Rubus condensatus P. J. Muell., Flora (Regensb.), 41: 167 (1858) Among the specimens which Sudre (1911) cited to illustrate this taxon is a series (Sudre’s Rubi rari no. 68) collected by P. J. Mueller from Reissbach, Alsace, the locus classicus, 20/7/1857. There is a good example in MANCH which tallies well with the original description. Watson (1958) thought that a bramble he found on Harrow Weald Common, Middlesex, v.c. 21, was R. con- densatus. But his specimen in my herbarium does not match the MANCH specimen closely. However a bramble of central England, which has puzzled batologists for many years, clearly does. There are specimens of this bramble in MANCH and SLBI collected by A. Ley, 25/8/1904, from the Wyre Forest, Salop, v.c. 40, which Ley named R. serpens Weihe with the approval of W.M. Rogers. But they differ from Rogers’ (1900) description of R. serpens, which, however, may not be reliable. The true R. serpens Weihe ex Lejeune & Court. has often been misinterpreted, but it was evidently a hairy, very glandular bramble with many quinate as well as some ternate leaves. There is another specimen in BIRM collected by Ley, 2/9/1909, from Wenlock Edge, Presthope, Salop, which Ley called R. serpens forma. 46 E. S. EDEES Watson, to whom I sent a Staffordshire specimen collected in 1953, associated it with R. scaber Weihe & Nees, and B. A. Miles told me (in /itt. 1968) that he had seen Staffordshire specimens, collected by J. Fraser in 1877 and 1878, which had also been named R. scaber. R. condensatus resembles R. scaber in having ternate leaves, small white petals and erect sepals. But there are many differences, particularly in the armature and glandular development of the stem and rachis and in the shape, texture and serration of the leaflets. The following description of the Wyre Forest bramble was prepared before I had seen the MANCH specimen or had read Mueller’s original account. It is offered for comparison with Mueller’s description and as a substitute for Watson’s (1958): Stems bluntly angled, becoming reddish purple in exposure, thinly pilose, with a few scattered acicles and glandular hairs. Prickles small, often shorter than the diameter of the stem, declining or sometimes slightly curved, unevenly arranged. Leaves ternate. Leaflets plicate, thinly strigose and shining green above, paler below and almost glabrous apart from a few hairs on the veins. Margins shallowly crenate. Terminal leaflets obovate- oblong, acuminate, emarginate or cordate, three to four times as long as their petiolules. Petioles with a few weak, declining or curved prickles. Flowering branches with adpressed and short, spreading hairs, frequent short acicles and a few glandular hairs. Prickles slender, declining or curved. Lower leaves ternate, upper leaves simple, broad, cordate, nearly glabrous on both sides. Panicles dense with interlacing pedicels and sub- sidiary clusters on long, suberect peduncles in the axils of the leaves. Sepals sparingly aculeolate and glandular, long-pointed, white-margined, reflexed in flower, erect in young fruit. Petals white, spaced, obovate, 10 x 5 mm. Stamens white, equalling the green styles. Anthers pilose or subpilose. Carpels densely hairy at first, bearded later. R. condensatus is frequent in the Wyre Forest, GR 32/7.7, and in south-western Staffs., v.c. 39. I have a series of specimens in my herbarium from Kinver, GR 32/8.8, Codsall, GR 33/8.0, and Wrottesley, GR 33/8.0. 3. Rubus painteri E. S. Edees, sp. nov. R. eifeliensis sensu W. C. R. Wats. ex parte, non Wirtg. Turiones pilis patentibus, albis, simplicibus dense vestiti. Glanduli stipitati pauci vel nulli. Aculei pilosi, numerosi (15—20 per 5 cm), ad angulos dispositi, aeque longi ac diametrum caulis, patentes vel paulo declinati. Folia ternata vel quinata pedata. Foliola utrinque griseoviridia, superne strigosa, inferne pilis multis ad nervos dispositis villosa, aequaliter duplicato-serrata. Foliolum terminale late obovatum vel paene rotundatum, cuspidatum, basi emarginatum vel subcordatum, petiolulo proprio triplo longius. Foliola infima breviter petiolulata (2-3 mm). Petioli pilis albis patentibus dense vestiti aculeisque parum curvatis muniti. Rami floriferi aculeis pilisque illis caulis similibus instructi. Inflorescentiae late elongatae, usque ad apicem foliiferae, infra foliis ternatis, supra bracteis foliaceis et longe petiolatis praeditae. Foliola utrinque cinerascentia, subtus ad nervos molliter pilosa. Ramuli inferiores (usque ad 5 cm longae) adscendentes, superiores (2-4 cm) patuli, omnes floribus breviter pedicillatis ornati. Sepala pilosa, aculeolata, laxe reflexa. Petala alba vel dilute rosea, obovato-cuneata, 12 x 7 mm. Stamina alba stylos pallidos superantia. Receptacula hirsuta. Stems densely hairy with patent, white, simple hairs. Glandular hairs rare or absent. Prickles hairy, numerous (15-20 per 5 cm length of stem), situated on the angles, equal in length to stem diameter, patent or slightly declining. Leaves ternate or quinate-pedate. Leaflets greyish green on both surfaces, strigose above and soft with many hairs on the veins beneath, evenly but coarsely serrate. Terminal leaflets broadly obovate or nearly round, with a cuspidate point and emarginate or subcordate base, three times as long as their petiolules. Basal leaflets with short petiolules (2-3 mm). Petioles with dense, spreading, white hairs and nearly patent, slightly curved prickles. NOTES ON BRITISH RUBI, 4 AT Flowering branches armed and clothed like the stems. Panicles, when well developed, broadly cylindrical and leafy to the summit, the lower ternate leaves giving place to leafy bracts on long petioles. Leaflets ashy grey on both sides and soft with many hairs on the veins beneath. Lower panicle branches ascending, about 5 cm long, upper branches patent and decreasing in length from 4 to 2 cm, all bearing clusters of flowers on short pedicels. Sepals hairy, aculeolate, loosely reflexed. Petals white or pale pink, obovate-cuneate, 12 x 7 mm. Stamens white, longer than the pale styles. Receptacles hirsute. HOLOTYPUS: Raven’s Clough, Rushton, GR 33/91.63, Staffs., v.c. 39, 22/7/1944, W. D. Graddon (herb. E. S. Edees 3824) This is a common bramble on the foothills of the Pennines in Staffordshire and Cheshire. When Watson visited Staffordshire in 1950 he examined several bushes in the field and later on identified them with R. eifeliensis Wirtg., a bramble which he claimed to have found already at Boar’s Hill, near Oxford, Berks., v.c. 22. But the Staffordshire and Cheshire bramble, of which I have several specimens and which I know well in the field, is not identical with Watson’s specimen from Boar’s Hill, 7/7/1936, now in my herbarium, nor with authentic specimens of R. eifeliensis Wirtg. Indeed the Boar’s Hill bramble is not a close match for R. eifeliensis either, nor is another specimen in my herbarium from Fancy, near Plymouth, S. Devon, v.c. 3, 20/7/1939, collected by Watson and also named R eifeliensis. No British exsiccata which can be unequivocally named R. eifeliensis have yet been seen. R. eifeliensis was first described by Wirtgen in 1858 on the labels of specimens sent out as Herbarium Ruborum Rhenanorum, ed. 1, no. 94. It was a very brief description, stating simply that the bramble resembled R. silesiacus Weihe, but differed in having a shaggy inflorescence, spreading and leafy below, and large, reddish petals. I have not seen a specimen of this number, but I have one of Herb. Rub. Rhen., ed. 2, no. 69, in my herbarium, gathered by E. Ley from the locus classicus in 1860 and sent out by Wirtgen under the same name. There are certainly resemblances between the Staffordshire and Eifel plants, but the Eifel bramble has more finely toothed leaflets, more strongly curved prickles on the stem, felted leaflets and is said to have red petals. A. Newton, who saw the German plant in situ in 1974, considers it quite different from the Cheshire and Staffordshire bramble we both know well. The German plant has a much stronger development of glands and acicles, is dark green rather than grey-green and has pink to deep pink petals. R. painteri, on the other hand, is notable for its white petals, nearly eglandular stem, nearly patent prickles, soft but not felted leaflets and strongly hirsute receptacles. Like R. eifeliensis it is a very shaggy bramble. In addition to the holytype I have Staffordshire paratypes collected by myself (herb. E.S.E.) from Biddulph, GR 33/89.59, Black Bank, near Newcastle, GR 33/81.47, Highshutt, near Cheadle, GR 43/03.43, and Meerbrook, GR 33/98.59, and one from Congleton, GR 33/89.62, Cheshire, v.c. 58. The name commemorates the work of W. H. Painter (1835-1910), who took a special interest in Rubi. There are specimens of R. painteri in MANCH, collected by Painter from Biddulph in 1889 and 1890, under the name R. pyramidalis Kalt. 4. Rubus turritus E. S. Edees, sp. nov. R. apricus sensu W. C. R. Wats. ex parte, non Wimm. Turiones obtuse angulati, faciebus planis, glabrescentes, purpurascentes. Aculei maiores tenues, declinati, subaequales, diametro caulis paulo longiores, plerumque ad angulos dispositi. Aculei minores inaequales sed non in ceteros abeuntes. Aculeoli numerosi, glanduli stipitati aliquantum sparsiores. Folia quinata pedata vel rarius ternata. Foliola utrinque viridia, supra glabra, infra pilosa sed non tomentosa, plus minusve aequaliter serrata. Foliolum terminale obovatum, longe acumina- tum, in basin emarginatum saepe angustatum, petiolulo proprio triplo longius. Petioli glandulosi et leviter villosi aculeis tenuibus aculeolisque armati. Stipulae lineares, glandulosae. Rami floriferi inferne leviter pilosi aculeolis tuberculisque muniti, superne dense villoso-tomen- tosi, glandulosi, aculeolati. Aculei tenues, debiles, declinati, aequales, diametro caulis vix longiores. 48 E. S. EDEES Folia ternata. Foliola obovata, utrinque viridia. Inflorescentiae apicem versus late cylindratae pedunculis longis, subaequalibus, 1—3-floris. Flores 2.5 cm diametro. Sepala dense hirsuta, aculeolata, glandulosa, attenuata, erecta. Petala alba vel primo dilute rosea, remota, lanceolato-obovata, 15 x 6 mm, glabra. Stamina alba stylos virides vix superantia. Carpella glabra. Receptacula pilosa. Stems bluntly angled with flat sides, green, becoming purple on the angles, glabrescent. Prickles slender, declining, somewhat variable in length, the longest slightly exceeding the diameter of the stem, mostly confined to the angles. Pricklets and acicles numerous and varying in size but distinct from the main prickles. Glandular hairs frequent, but less numerous than the pricklets and acicles. Leaves sometimes ternate but usually quinate-pedate. Leaflets green on both surfaces, glabrous above and hairy, though not felted, beneath; regularly toothed except sometimes towards the tip. Terminal leaflets obovate, acuminate, often with straight sides converging to a narrow, emarginate base, three times as long as their petiolules. Petioles with slender declining prickles, acicles and glandular hairs, and thinly clothed with patent white hairs. Stipules narrow and fringed with glandular hairs. Flowering branches with sparse, white spreading hairs and numerous pricklets and tubercles below, felted above with dense, spreading simple hairs, glandular hairs and acicles. Prickles slender, weak, declining and of even length, scarcely longer than the stem-width. Leaves ternate with obovate leaflets, green on both sides. Panicles broadly cylindrical, with long, subequal peduncles, — each bearing 1-3 flowers. Flowers 2.5 cm in diameter. Sepals densely hairy, aculeolate, glandular, long-pointed and erect. Petals white, though sometimes pinkish in bud, spaced, obovate-lanceolate, 15 x 6 mm, glabrous. Stamens white, slightly longer than the green styles. Carpels glabrous. Receptacles hairy. HOLOTYPUS: Hanchurch Hills, on the south side of the water-tower enclosure, GR 33/840.397, Staffs., v.c. 39, 1/8/1972, E. S. Edees 20823 (herb. E.S.E.) This is another local bramble which is abundant on Hanchurch Hills and distributed over a wide area of north Staffordshire. I have paratypes in my herbarium from Barlaston, GR 33/92.39, Fulford, GR 33/93.38, Leigh, GR 43/00.33, Mucklestone, GR 33/74.37, Sandon, GR 33/93.33, Stone, GR 33/90.37, Swynnerton, GR 33/84.41, and Whitmore, GR 33/82.41, in addition to several specimens from Hanchurch Hills. Watson, who saw the bramble in situ, identified it with R. apricus Wimm. But it does not match closely specimens of R. apricus from the original station, of which I have seen several. I have two in my own herbarium and there is one in K, collected by F. Schwarzer in 1869, ‘Am Streitberg in Schlesien’, with a note, ‘Specimen typicum in statione originali autoris lectum’, pasted on to the sheet. Prof. H. E. Weber, to whom I sent a specimen of the Staffordshire bramble with Watson’s suggested name, replied ‘Very far away from the true plant which I know from original specimens’. Nor is R. turritus closely related to R. bercheriensis (Druce ex Rogers) Rogers, which Watson (1935) described as R. apricus var. sparsipilus W. C. R. Wats. 5. Rubus infestus Weihe ex Boenn., Prodr. Fl. Monast., 153 (1824) R. taeniarum Lindeb., Nov. Fl. Scand., 5 (1858) There is a widely distributed bramble in northern England and southern Scotland which Rogers (1900) identified with R. infestus Weihe ex Boenn. and Watson (1958) with R. taeniarum Lindeb. Apart from Heslop-Harrison (1968), most living batologists agree that these are two names for the same taxon. Many Continental batologists have for long held this view. Even Lindeberg admitted a close relationship between R. taeniarum and R. infestus. A specimen of his Herbarium Ruborum Scandinaviae no. 19 in MANCH from the locus classicus for R. taeniarum, ‘In insula Oroust Bahusiae, August 1880’, has a Latin description which begins: ‘R. infesto Whe proximus eiusque quasi forma borealis.” But Lindeberg thought that the grooved, more lightly-armed stem, with fewer glandular hairs, the felted leaflets and compound panicles sufficiently distinguished it. Focke (1877, 1914) cited R. taeniarum as a synonym of R. infestus and Sudre (1911) described it NOTES ON BRITISH RUBI, 4 49 as a variety of R. infestus having a rhomboid terminal leaflet, with a nearly entire base, and a lax inflorescence. Of the two numbers which Sudre (1911) cited to illustrate R. infestus, one at least (Sudre’s Batotheca Europaea no. 37) is identical with R. taeniarum. There is a good example in BM which I have carefully examined. The petals are rounded and notched and said to be rose-coloured, the stamens are scarcely longer than the styles and the carpels are strongly pilose. I quickly con- cluded that this was R. taeniarum and then I discovered a note with one of Watson’s specimens in my herbarium which stated that in Watson’s opinion too Sudre’s no. 37 was R. taeniarum. The other number which Sudre cited for R. infestus was no. 122 of the Set of British Rubi. Unfor- tunately pieces for this number were cut from bushes from two widely separated areas, viz. Thursley Common, Surrey, v.c. 17, and Bethesda, Caernarvon, v.c. 49, and represent two different taxa. The Bethesda bramble is R. taeniarum, but the bramble of Thursley Common is R. infestus sensu Watson, non Weihe ex Boenn., which (see below) it is proposed to call R. infestior. Both were distributed as R. infestus and Sudre (1904) said of the specimen or specimens he saw: ‘Espéce bien caractérisée’. Watson (1931, 1949) followed Neuman (1915) in thinking R. infestus and R. taeniarum to be distinct. At first he thought that the bramble recorded for northern England and Scotland should be called R. spurius Neuman, a name which according to Weber (1972) was given to a weak specimen of R. infestus from its origina! German station. When Watson visited Staffordshire in 1950 he called several bushes, which we examined together in the field, R. spurius, maintaining that the true R. infestus was a quite different bramble. In his Handbook (1958) he substituted the name R. taeniarum for R. spurius. There is a note on one of his specimens in my herbarium, dated 1951 and followed by his initials, with the equation R. taeniarum = R. spurius. Watson (1931) claimed that the true R. infestus had white fiowers and imbricate leaflets reminiscent of the Sub- erecti and was more fiercely armed, and he applied this name to a bramble which he had seen in several places in south-eastern England, as, for example, on Farnborough Common, Hayes Common and in Barnet Wood, W. Kent, v.c. 16, and on Netley Heath and in the Roughs near Guildford, Surrey, v.c. 17. This is a well marked taxon which resembles the plate of R. infestus in Rubi Germanici (Weihe & Nees 1825) to some extent, but not the lectotype at Kiel. The lectotype, designated by Professor Weber in 1975, was labelled by Weihe ‘Rubus infestus Weihe, Minden’, and is a good specimen of the taxon which Lindeberg later called R. taeniarum. R. infestus was first recorded for Minden, West Germany, where Weihe & Nees (1825) said it was far from rare and easily recognized. Focke (1877) said it was plentiful in that province in the neighbourhood of Menninghiiffen. There is a specimen in K of Focke’s Rubi Selecti no. 66 from Minden, ‘In Hecken und Gebiischen zu Heddingsen bei Menninghiiffen’, collected by H. Banning in 1871. Weber sent me a recent specimen from the same place, collected by himself in 1968, and told me (in litt. 1975) that it was still very common. These specimens match one another and also ' specimens of R. taeniarum collected by Lindeberg and others from Sweden and Denmark, of which there is a good series in MANCH, leaving no doubt that R. infestus Weihe and R. taeniarum Lindeb. are conspecific. Nor is there any doubt that the bramble of northern England and southern Scotland, which Watson called R. taeniarum and his predecessors R. infestus, was also correctly identified. British exsiccata have been compared with authentic specimens of R. taeniarum and the Minden specimens of R. infestus and match them both. But R. infestus is the earlier name. Accord- ingly specimens which have been determined R. taeniarum for British collectors should now be re-named R. infestus. The bramble from south-eastern England requires a new name. 6. Rubus infestior E. S. Edees, sp. nov. R. infestus sensu W. C. R. Wats., non Weihe ex Boenn. Turiones angulati, glabrescentes. Aculei crebri, e basi lata declinati vel curvati, inter se valde diversi, maiores diametro caulis parum longiores, minores multo breviores, haud ad angulos omnino dispositi. Aciculi glandulique stipitati vulgo sparsi. Folia quinata subdigitata. Foliola nonnunquam imbricata, utrinque viridia, supra glabrescentia, subtus pallide pubescentia, inaequaliter serrata. Foliolum terminale ovatum, cordatum, acu- D 30 E. S. EDEES minatum, circa 8x6 cm, petiolulo proprio triplo longius. Petioli aculeis falcatis aculeolisque inaequalibus confertim obsiti. Rami floriferi aculeis declinatis falcatis et aculeolis multis dense armati. Aciculi glandulique stipitati, quorum plurimi minus quam 1 mm longi, praesertim in pedunculis pedicellisque numerosi. Inflorescentiae angustae, ramulis brevibus, adscendentibus, paucifloris instructae. Sepala aculeolata, laxe reflexa vel patentia. Petala alba. Stamina alba stylos pallidos supendncd: Carpella glabra. Stems angled, glabrescent. Prickles crowded, broad-based, declining and curved, varying greatly in size from pricklets much shorter than the stem-width to prickles which are a little longer than it, occurring on the faces as well as the angles. Acicles and glandular hairs usually few. Leaves quinate-subdigitate. Leaflets frequently imbricate, green on both sides, nearly glabrous above, soft with short hairs and paler below, rather coarsely serrate. Terminal leaflet ovate, cordate, acuminate, about 8 x 6 cm, three times as long as its petiolule. Petioles strongly armed with falcate prickles and many prickiets of variable length. Flowering branches densely armed with crowded falcate and declining prickles sid pricklets of various sizes. Acicles and glandular hairs, which are usually less than 1 mm long, mostly on the peduncles and pedicels. Panicles narrow, with short, ascending, few-flowered branches. | Sepals aculeolate, loosely reflexed or patent. Petals white. Stamens white, longer than the green styles. Carpels glabrous. HOLOTYPUS: Witley Common, near Milford, GR 41/9.4, Surrey, v.c. 17, 17/7/1935, C. Avery, “oe W.C. R. Watson as R. infestus Weihe (herb. E. S. Edees) The strong, unequal prickles, which are sometimes so crowded, especially on the rachis, that they touch one another, place this bramble in the section Glandulosi series Hystrices of Watson’s arrange- ment. The flowers were well described by Watson (1931): ‘The flowers are very characteristic; white, cup-shaped, with the upper parts of the petals inflexed. The petals are roundish but they taper below into a rather long claw. .. . The white stamens are much longer than the green styles. The young carpels are glabrous.’ The true R. infestus (R. taeniarum) differs from R. infestior in having smaller and more numerous flowers, with pink, notched petals, short stamens and hairy carpels, and is less heavily armed. It belongs to the section Appendiculati in Watson’s classification. R. infestior differs from R. setulosus Muell. & Lefév., which Watson (1958) cited as a synonym, in several ways. P. J. Mueller’s definitive specimen in Lausanne (LAU) (Bois du Roi, Oise, France, 1856, L. V. Lefévre), which I have examined, has obovate terminal leaflets, strongly cuneate in the panicle, and a much weaker armature. R. marianus (Krause) H. E. Weber (R. infestus var. marianus Krause), of which Professor Weber has sent me a specimen collected and determined by himself, also differs most markedly from R. infestior in armature. It has not been recorded for the British Isles. R. infestus is widely distributed in north-western Europe and is known in the British Isles from the following vice counties: 39, 40, 49, 57-60, 62-65, 67, 69-71, 73, 77, 78, 80, 81, 86-89, 96-99, 101, 102, H39. R. infestior is locally common in the south-eastern counties of England. In addition to the holotype I have paratypes in my herbarium from Hayes Common, GR 51/401.657, W. Kent, v.c. 16, Thursley Common, GR 41/9.4, Surrey, v.c. 17, and Yateley Common, GR 41/84.59, and Bramshott Common, GR 41/85.33, N. Hants., v.c. 12. 7. Rubus anglohirtus E. S. Edees, sp. nov. R. hirtus sensu W. C. R. Wats. ex parte, non Waldst. & Kit. Turiones obtuse angulati vel teretiusculi, in apricis rufescentes, pruinosi, breviter et dense pilosi, glandulis stipitatis aciculisque obsiti. Aculei tenues, parvi (saepe diametro caulis dimidio breviores), paulo declinati. Folia quinata pedata. Foliola utrinque viridia, supra strigosa, subtus ad nervos pilosa, in- aequaliter serrata. Foliolum terminale circa 10x 7 cm, obovatum, interdum subincisum, cuspi- NOTES ON BRITISH RUBI, 4 51 datum, basi emarginatum vel cordatum, petiolulo proprio triplo vel quadruplo longius. Petioli pilis illis caulis similibus vestiti et aculeis tenuibus, declinatis vel curvatis armati. Rami floriferi pilis brevibus patentibusque dense vestiti et glandulis stipitatis aciculisque plus minusve aequalibus exasperati. Aculei tenues, subulati, declinati vel curvati. Inflorescentiae laxe elongatae, apice truncatae, ramulis inferioribus longe adscendentibus, superioribus patulis. Pedun- culi pedicellique pilis adpressis glandulisque inaequalibus copiose instructi. Sepala aculeolata, appendiculata, primo reflexa, demum erecto-patentia. Petala alba, late elliptica (14 x 7 mm), ad marginem glabra. Stamina alba stylos pallidos parum superantia. Carpella et receptacula pilosa. Stems bluntly angled or nearly round, dull red in exposure, with many short, simple and stellate hairs, glandular hairs and acicles, pruinose. Prickles slender, small (often not more than half the stem-width), slightly declining. Leaves quinate-pedate. Leaflets light green on both surfaces, strigose above, hairy on the veins below, with somewhat irregular teeth. Terminal leaflet about 10x 7 cm, obovate, sometimes shouldered or incised, with a cuspidate point and emarginate or cordate base, three to four times as long as its petiolule. Petioles clothed like the stem and with slender, declining or curved prickles. Flowering branches densely hairy with short spreading hairs, many short glandular hairs and acicles of more or less equal length. Prickles slender, weak, declining or curved. Panicles lax, with spreading peduncles above the leaves and long, ascending lower branches, truncate. Peduncles and pedicels densely covered with adpressed hairs and many glandular hairs of variable length. Sepals aculeolate, leafy-pointed, at first reflexed, later patent to erect. Petals white, broadly elliptic (14x 7 mm), glabrous on the margin. Stamens white, slightly exceeding the pale styles. Carpels and receptacles pilose. HOLOTYPUS: Ropsley Rise Wood, near Grantham, GR 43/97.34, S. Lincs., v.c. 53, 27/7/1965, E. S. Edees 19015 (herb. E.S.E.) This is one of several taxa which Watson (1958) called R. hirtus Waldst. & Kit., but Professor H. E. Weber, to whom I sent a specimen, told me (in litt. 1975) that it is quite different from the bramble to which Continental batologists have usually applied this name and corresponds with nothing known to him. R. hirtus has not been typified and as generally understood is a notoriously aggregate taxon. Rogers (1900) said he could not define it and quoted Focke as saying that it was a polymorphic species with countless indefinable varieties. I have seen no specimen of R. hirtus named by the original authors, but the bramble described and illustrated by Weihe & Nees (1827) is different from ours. A specimen of R. hirtus forma borealis G. Braun (Herbarium Ruborum Germaniae no. 60) in K is closer to R. anglohirtus but is not identical with it. R. anglohirtus is a constant and easily recognized taxon with a wide distribution in eastern England, occurring in woods from S. Lincs., v.c. 53, to E. Kent, v.c. 15. In addition to the holotype I have paratypes in my herbarium from Twyford Forest, near Bourne, GR 43/95.23, Row Wood, near Bourne, GR 53/07.26, Spring Wood, near Bourne, GR 53/06.23, Ponton Park Wood, near Grantham, GR 43/94.31, all in S. Lincs., v.c. 53, and from Alsa Wood, near Stansted, GR 52/5.2, Birchanger Wood, GR 52/503.225, Broom Wood, Ugley, GR 52/510.285, N. Essex, v.c. 19. ACKNOWLEDGMENTS I am greatly indebted to Professor H. E. Weber, whose recent book (1972) has stimulated interest in the relationship of the British and Continental Rubi and prompted a fruitful correspondence and exchange of specimens. I must also once more express my gratitude to A. Newton for some references and many helpful suggestions and to the curators of the national herbaria mentioned in this paper for the loan of specimens. REFERENCES BABINGTON, C. C. (1869). The British Rubi. London. FockeE, W. O. (1877). Synopsis Ruborum Germaniae. Bremen. Focke, W. O. (1914). Species Ruborum. Monographiae generis Rubi, Prodromus, 3. Bib/thca bot., 83: 1- 274. 52 E. S. EDEES HESLOP-HARRISON, Y. (1968). Rubus L., in Tutin, T. G. et al., eds. Flora Europaea, 2: 7-25. Cambridge. — Less, E. (1852). Botany of the Malvern Hills, 2nd ed. London. Legs, E. (1867). The botany of Worcester. Worcester. NEUMAN, L. M. (1915). Ar Rubus taeniarum Lindeberg identisk med Rubus infestus Weihe? Botaniska Notiser, 1915: 85-91. Rocers, W. M. (1900). Handbook of British Rubi. London. SupreE, H. (1904). Observations sur ‘Set of British Rubi’. Bull. Soc. Etud. scient. Angers, 33: 106-145. Supre, H. (1911). Rubi Europae, (4). Paris. WATSON, W. C. R. (1931). Brambles of Kent and Surrey, 4. Lond. Nat., 10: 68-74. WATSON, W. C. R. (1935). Brambles of Kent and Surrey, 6. Lond. Nat., 14: 59-66. WATSON, W. C. R. (1949). Weihean species of Rubus in Britain. Watsonia, 1: 71-83. Watson, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge. Weser, H. E. (1972). Die Gattung Rubus L. (Rosaceae) im nordwestlichen Europa. Lehre. WEIHE, K. E. A. & NEES VON ESENBECK, C. G. D. (1822-27). Rubi Germanici. Elberfeld. (Accepted May 1975) Watsonia, 11, 53-57 (1976) 53 Chromosome number of Puccinellia maritima (Huds.) Parl. in the British Isles R. SCOTT and A. J. GRAY Institute of Terrestrial Ecology, Colney Research Station, Colney Lane, Norwich ABSTRACT Chromosome counts were made on root-tip cells of Puccinellia maritima plants representing a wide range of localities and growth forms. All plants examined had 2n = 56. INTRODUCTION The most frequently quoted chromosome number for British Puccinellia maritima (Huds.) Parl. is 2n = 63 (Mills 1967, Newton 1965), which is nonaploid if x = 7 for the genus, and consistent with the suggestion (Tutin 1955) that the species usually reproduces vegetatively and may be apomictic. Mills also reported 2” = 56, which is commonly listed for foreign material (Bernstr6ém 1948, Church 1949, Sorensen 1958). Other British counts are 2” = 14, 49, and 77 (Brown-Packer 1961), and from foreign material 2n = 42 (Church 1949), c 60 (Castro & Fontes 1946) and 70 (Wulff 1937, Rodrigues 1953). The counts made by Mills (1967) and Newton (1965) are listed in Table 1, and suggest the possibility of differences in distribution of the octoploid and nonaploid plants in Britain. Because it also seemed possible that the wide range of phenotypic variation observed in the species might correspond to different cytological races, chromosome counts were made on a large number of plants collected as part of a study of infraspecific variation. Plants were sampled from a wide range of habitats and locations (Table 2), different growth forms being included when found adjacent to one another in the field. Growth form differences, often very large, were usually main- tained after a period of collateral cultivation (Table 3). The object of the work presented here was to relate chromosome number to the growth forms and geographical races of the plant. TABLE 1. PREVIOUS LOCALIZED CHROMOSOME COUNTS FOR PUCCINELLIA MARITIMA IN THE BRITISH ISLES Author Location No. of plants Chromosome counted Number 2n A. R. Mills Parkgate, Cheshire, v.c. 58 8 51-62 A. R. Mills Silverdale, W. Lancs., v.c. 60 3 51-56 A. R. Mills Ringhaddy, Down, v.c. H38 5 51-57 A. R. Mills Baldoyle, Down, v..c. H38 1 c 56 A. R. Mills Rye, E. Sussex, v.c. 14 1 c 60 A. R. Mills Tollesbury, N. Essex, v.c. 19 1 c 63 A. R. Mills West Mersea, N. Essex, v.c. 19 4 62-63 A. R. Mills Mudeford, S. Hants., v.c. 11 1 c 63 L. E. Newton Canvey Island, S. Essex, v.c. 18 1 c 63 L. E. Newton Tollesbury (tall plant), N. Essex, v.c. 19 1 c 63 L. E. Newton Tollesbury (dwarf plant), N. Essex, v.c. 19 1 c 63 L. E. Newton Yantlet Creek, W. Kent, v.c. 16 1 c 63 34 R. SCOTT AND A. J. GRAY TABLE 2. SITES OF PUCCINELLIA MARITIMA WITH 2n = 56 DETERMINED Accession Grid Location Habitat type code reference Ps 1 34/465.795 Arnside, Westmorland, v.c. 69 Grazed saltmarsh | nee | J/51.64 Strangford Lough, Down, v.c. H38 Ungrazed saltmarsh P 17 54/355.032 Tetney, N. Lincs., v.c. 54 Tidal riverbank P (28 34/202.758 Roanhead, Furness, v.c. 69b Cattle-grazed saltmarsh P 31 34/432.789 Holme Island, Furness, v.c. 69b_ . Ungrazed saltmarsh P35 34/437.791 Meathop, Westmorland, v.c. 69 Grazed saltmarsh P 44 34/477.828 Gilpin Bridge, Westmorland, v.c. 69 Grazed saltmarsh P57 34/466.798 Arnside, Westmorland, v.c. 69 Grazed saltmarsh P 58 34/452.759 Silverdale, Westmorland, v.c. 69 Grazed saltmarsh P 59 34/450.758 Silverdale, Westmorland, v.c. 69 Grazed saltmarsh Po 34/324.833 Haverthwaite, Furness, v.c. 69b Grazed saltmarsh P 97 34/471.682 Carnforth, W. Lancs., v.c. 60 Grazed saltmarsh P 98 34/471.682 Carnforth, W. Lancs., v.c. 60 Grazed saltmarsh P 100 34/474.713 Warton, W. Lancs., v.c. 60 Grazed saltmarsh P 102 34/484.716 Warton, W. Lancs., v.c. 60 Grazed saltmarsh P 118 53/926.457 Wells, W. Norfolk, v.c. 28 Ungrazed saltmarsh P 119 53/926.457 Wells, W. Norfolk, v.c. 28 Ungrazed saltmarsh P 120 53/926.457 Wells, W. Norfolk, v.c. 28 Ungrazed saltmarsh P 124 51/028.014 Littlehampton, W. Sussex, v.c. 13 Sandflat P 127 63/487.065 Breydon Water, E. Suffolk, v.c. 25 Ungrazed saltmarsh P¥52 51/935.193 Rye Harbour, E. Sussex, v.c. 14 Creek bank P 156 51/444.082 Beddingham, E. Sussex, v.c. 14 Tidal riverbank P 165 40/183.918 Mudeford, S. Hants., v.c. 11 Sandy mudfiat P 167 30/255.907 Axmouth, S. Devon, v.c. 3 Stony, tidal river shore P 177 20/116.579 Lostwithiel, E. Cornwall, v.c. 2 Stony, tidal river shore P 183 21/205.064 Bude, E. Cornwall, v.c. 2 Stony, tidal river shore P 188 31/550.753 Avon Gorge, W. Gloucs., v.c. 34 Ungrazed saltmarsh P 236 34/208.855 Foxfield, Furness, v.c. 69b Grazed saltmarsh P 241 34/184.694 Vickerstown, Furness, v.c. 69b Ungrazed saltmarsh P 254 28/847.853 Morrich More, E. Ross, v.c. 106 Sandflats P 255 16/715.821 Ulva Islands, S. Ebudes, v.c. 102 Gravelly shore P 259 53/555.578 Gibraltar Point, N. Lincs., v.c. 54 Ungrazed saltmarsh P.282 27/096.587 Loch Leven, Argyll Main, v.c. 98 Ungrazed saltmarsh P 295 18/897.547 Loch Torridon, W. Ross, v.c. 105 Grazed saltmarsh P 299 29/554.540 Kyle of Tongue, W. Sutherland, v.c. 108 Grazed saltmarsh P 304 39/166.294 Dunbeath, Caithness, v.c. 109 Quay top P322 37/342.294 Kingoodie, Fife, v.c. 85 Ungrazed saltmarsh P 326 46/082.426 Lindisfarne, Cheviot, v.c. 68 Ungrazed saltmarsh P 329 54/384.018 North Cotes, N. Lincs., v.c. 54 Ungrazed saltmarsh P 332 62/048.212 Wivenhoe, N. Essex, v.c. 19 Ungrazed saltmarsh P 333 23/405.672 R. Cefni, Anglesey, v.c. 52 Ungrazed saltmarsh P.337 33/648.512 Nantwich, Cheshire, v.c. 58 Inland saltmarsh P 342 53/920.448 Wells, W. Norfolk, v.c. 28 Ungrazed saltmarsh P 358 53/939.450 Wells, W. Norfolk, v.c. 28 Ungrazed saltmarsh IN THE PRESENT STUDY METHODS Tillers taken from individual plants in the field were grown in plastic pots containing John Innes No. 3 potting compost and a basal layer of moss-peat. Portions of root-tip c 30 mm long were taken from the peat, where they were grit-free, pre-fixed for 3 hours in dilute 8-hydroxyquinoline at 5°C, fixed for 3 hours in 3:1 ethanol/acetic acid, and hydrolyzed for 10 minutes at 60°C in 1 N HCI, followed by staining in Feulgen reagent for 1.5 hours. After staining, the tips were treated in pectinase (Ostergren & Heneen 1962) to facilitate cell spreading. The preparations were squashed in 45% acetic acid, and temporary mounts were made by ringing coverslips with rubber solution. Whenever possible chromosome counts were taken from replicated root-tips and plants. CHROMOSOME NUMBER OF PUCCINELLIA MARITIMA 55 TABLE 3. GROWTH FORM DIFFERENCES BETWEEN TWO CLONES OF PUCCINELLIA MARITIMA FROM WELLS, NORFOLK 3rd leaf-blade length Maximum vegetative tiller Accession (cm) height from ground (cm) No. of tillers code (13/6/72 (13/6/73 (13/6/72 (13/6/73 (18/4/73 in field) at Norwich) in field) at Norwich) at Norwich) P 348 2.26 3.76 2.68 7.42 19.5 (0.129) (0.108) (0.292) (0.684) (2.53) P 353 8.38 7.54 7.94 14.64 11.25 (0.443) (0.389) (0.534) (1.13) (2.29) Replicated single tillers were grown in non-saline soil outdoors at Norwich from Oct. 1972 to July 1973. Standard error in brackets. CHROMOSOME NUMBERS P. maritima has chromosomes which are relatively long and thin, which makes counting difficult because they are often intermingled with one another in imperfect squashes. In over-spread pre- parations splitting of chromatids frequently occurred, adding another difficulty to counting. In good cells 56 chromosomes were usually present, but a few cells appeared to contain numbers in the range 53-55, suggesting a degree of somatic aneuploidy. All plants, including representatives of extremes of growth form, had a number at or just below 2” = 56, the octoploid (x = 7) level (Fig. 1). Some other counts ranged from 50 to 64, but in all instances where aberrant counts were investigated repetition on new material of the same plants led to the conclusion that 2n = c 56. FicureE 1. A root-tip cell of Puccinellia maritima showing a full complement of chromosomes, 2” = 56 (accession P 35). 56 R. SCOTT AND A. J. GRAY Ficure 2. Chromosome types in Puccinellia maritima. KARYOTYPE Preparations were insufficiently clear for full karyotype analysis but Fig. 2 summarizes five categories of chromosome size and form. About six pairs of small acrocentric chromosomes are present; the rest are metacentric ranging in size up to twice as long as the acrocentrics. At least two pairs of the largest metacentrics have distinct satellites, and identical chromosomes have been seen in other species of the genus Puccinellia. All chromosomes showed bands of more intense staining which matched in several pairs and could be valuable in karyotype analysis. DISCUSSION AND CONCLUSIONS Chromosome number was found to be constant in plants of different growth form coming from a wide range of sites, suggesting that it is not a facet of variation in the species in the British Isles. The widespread occurrence of octoploids (2n = 56) with little aneuploidy is consistent with results of our unpublished breeding experiments which show that P. maritima is an active out-breeder with a low selfing rate. The presence in its karyotype of sets of many similar chromosomes supports the idea that the species is a polyploid, but meiosis appears to be regular (Church 1949), and the plants are highly interfertile. Our evidence suggests that British plants are octoploid, but allowing for difficulties with counting it is not possible to rule out the existence of aneuploids. The wide range of morpho- logical variability of the species is likely to be the produce of environmental selection on popula- tions whose gene systems perform normal segregation and recombination, and not the expression of isolated lines maintained after a breakdown of sexuality. ACKNOWLEDGMENTS We thank W. E. Hughes, B. M. G. Jones, and A. R. Mills for information on Puccinellia. L. F. La Cour gave valuable advice on methods, and we are grateful to Miss N. Meek whose patient work added many new counts during 1974. F. T. Last and D. S. Ranwell made valuable comments on the text. REFERENCES BERNSTROM, P. (1948). In Love, A. & Love, D. Chromosome numbers of northern plant species. Rep. Dep. Agric. Univ. Reykjavik, Ser. B, 3: 9-131. BRown-PAcCKER, A. K. (1961). In Love, A. & Léve, D. Chromosome numbers of central and northwest European plant species. Op. bot. Soc. Lund., 5: 1-581. CHROMOSOME NUMBER OF PUCCINELLIA MARITIMA 57 Castro, D. DE & Fontes, F. C. (1946). Primeiro contacto citoloégico com a flora halofila dos salgados de Sacavém. Bolm Soc. broteriana, 15: 38-46. CuHurcH, G. L. (1949). A cytotaxonomic study of Glyceria and Puccinellia. Am. J. Bot., 36: 155-165. Mitts, A. R. (1967). A cytotaxonomic study of variation in Puccinellia maritima in the British Isles. M.Sc. thesis, University of Liverpool. Newton, L. E. (1965). Taxonomic studies on British species of Puccinellia. M.Sc. thesis, University of London. OSTERGREN, G. & HENEEN, W. K. (1962). A squash technique for chromosome morphological studies. Hereditas, 48: 332-341. RopriGcues, J. E. DE (1953). Contribuigéo para o conhecimento carioldgica das haldfitas e psamofitas literais. Diss. Dout. Coimbra. Univ. SORENSEN, TH. (1958). In JORGENSEN, C. A., SORENSEN, TH. & WESTERGAARD, M. The flowering plants of Greenland. A taxonomical and cytological survey. K. danske Vidensk. Selsk., Biol. Skr., 9(4): 1-172. TuTIN, T. G. (1955). Species problems in plants with reduced floral structure, in LousLey, J. E., ed. Species studies in the British flora, pp. 21-26. London. WuLrF, H. D. (1937). Karyologische Untersuchungen an der Halophytenflora Schleswig-Holstein. Jb. wiss. Bot., 84: 812-840. ROYAL IRISH ACADEMY Robert Lloyd Praeger Fund for Field Natural History Grants, not normally exceeding £50 in any one year, will be awarded for field work relevant to the Natural History of Ireland. Application forms, which should be returned before February 15, may be obtained from The Secretary, Royal Irish Academy, 19 Dawson Street, Dublin 2, who will also be glad to give further information. Watsonia, 11, 59-65 (1976) 59 Nomenclature of the British taxa of the Carex muricata L. aggregate R. W. DAVID 41 Barton Road, Cambridge ABSTRACT The nomenclatural problems surrounding the Carex muricata L. aggregate are discussed. The taxa as at present understood should be known as C. spicata Hudson, C. lumnitzeri (Rouy) Krecz., C. muricata L. subsp. muricata, C. muricata L. subsp. pairaei (F. W. Schultz) Celak., C. polyphylla Kar. & Kir., C. leersiana Rauschert, C. divulsa Stokes and C. chabertii F. W. Schultz. The Carex muricata L. aggregate comprises all the European taxa of Carex section Muehlen- bergianae Tuckerm., and of these C. polyphylla Kar. & Kir., C. divulsa Stokes, C. spicata Hudson and C. muricata L. (to quote the names included in Dandy (1958)) are commonly recognized as British plants. Their nomenclature, however, has undergone a multiplicity of confusions. This has been primarily due to uncertainty as to what is the Linnaean type; but the initial error has been compounded by subsequent mistakes in determination (especially confusions with C. otrubae Podp.), by inadequate descriptions that often emphasize characters that are in fact indecisive, and by differing opinions as to the rank of the taxa included within the aggregate. Linnaeus (1753) described C. muricata as follows: ‘muricata. 8. CAREX spiculis subovatis sessilibus remotis androgynis, capsulis acutis divergentibus spinosis. Fl. suec. 753. Gmel. sib. I p. 148.’ This original diagnosis could apply to several taxa in the aggregate; but Linnaeus’ herbarium (LINN) contains a Swedish specimen marked in Linnaeus’ own handwriting ‘8 muricata’. The use of the identical number and name directly connects the diagnosis and the specimen. Nevertheless nineteenth-century botanists failed to notice this connection. Most of them, for example Koch (1837), merely lumped all taxa of the aggregate under the Linnaean name. The case of Crépin (1859) is instructive, for it illustrates both the failure of the earlier botanists to find satisfactory criteria for the distinguishing of the individual taxa, and the distinctiveness of the taxa when such criteria are found. Crépin cited two varieties of ‘C. muricata’: « genuina Grenier & Godron, and f virens Koch. From each of these he segregated a subvariety, incrassata, distinguished by the thickened base of its utricle. Such utricles are found only in the taxon isolated by Hoppe (Sturm 1833) as C. contigua and now known as C. spicata Hudson. Evidently each of the varieties cited by Crépin was in itself a conglomerate of several taxa of the aggregate, including C. spicata, which he correctly separated out from each of the two conglomerates. Other attempts were made, as for example by Andersson (1849) and by Neuman (1901), to distinguish the individual taxa of the aggregate at least as subspecies or varieties, but these attempts appear to be no more soundly based than those of Grenier & Godron or of Koch. There was, however, through the century, a growing tendency, exemplified in Ascherson & Graebner (1902), to apply the Linnaean name not in its original sense but to C. contigua. Nelmes (1942) following Mackenzie (1935), produced evidence that the correct prior name for C. contigua is C. spicata Hudson. Though that evidence is not very weighty, the identification and the name have been generally accepted. To the original uncertainty about the identity of C. muricata a further complication was added by Linnaeus’ Swedish pupil, Johann Andreas Murray, who (Murray 1770) described Carex echinata. Formerly in LINN, but in January 1963 transferred to BM, is a sheet of four stems collected by Murray near Gottingen and inscribed by him ‘Carex echinata’. These specimens in fact belong to a taxon of the C. muricata aggregate identical with or near to Linnaeus’ type-specimen of C. muricata. Attempts to resolve this contradiction have followed two opposing lines. On the one hand, Kiikenthal (1905, 1909) took C. echinata Murray to be the correct name of C. muricata 60 R. W. DAVID and rejected the latter altogether as a nomen delendum on the grounds that it covered several members of the aggregate. In support of this view he observed (1909, p. 156 note 2) that the Linnaean herbarium contained, in addition to the specimen labelled C. muricata, an unnamed specimen of C. divulsa (which is true) and also an unnamed specimen of C. contigua. There is, however, no specimen of this last taxon in Linnaeus’ own collection. It was probably in one of the subsidiary collections (e.g. Murray’s) that have lately been transferred by the Linnean Society to other herbaria. On the other hand Mackenzie (1923), arguing with some force and citing not only Linnaeus’ Species Plantarum but also his Flora Suecica which describes C. muricata as a plant of wet ground, took C. muricata L. to be the correct name for C. echinata Murray, which he dis- missed as a later synonym. In this Mackenzie was followed by Kreczetovicz (1935) in Flora URSS. Britten (1907), followed by Kovacs (1910), Briquet (1910), Fernald (1917), and de Langhe (1944), had shown, however, that Murray’s published description of C. echinata referred without any possible doubt to the sedge otherwise known as C. stellulata Good.; and Nelmes (1959) implicitly rejected Mackenzie’s argument, maintaining that, as Linnaeus labelled both his description and his specimen with the same reference number, ‘8 muricata’, no confusion exists apart from the misidentifications of others, and that the name C. muricata should stand. Nelmes’ conclusion is surely right, although Hylander (1966), who appears to have been unaware of Nelmes’ 1959 paper (it was published in Russian), again rejected the name C. muricata as a nomen ambiguum. The question remains: what is in fact the plant labelled C. muricata by Linnaeus? Nelmes (1947, emended 1959) distinguished a northern and eastern taxon growing on limestone, and a western taxon preferring acid sands and gravels. Linnaeus’ type-specimen of C. muricata belongs to the northern taxon, although the glumes, which should be dark brown, are now quite bleached and the specimen is a less robust form than that described by Hylander (1966) as C. pairae [sic] subsp. borealis. As another example of the northern taxon Nelmes (1959) claimed C. loliacea sensu Schkuhr, non L. nec Schreber, as illustrated (the description is borrowed and applies to C. loliacea L.) in Plate Ee No. 91 of Schkuhr (1801). Two other names, C. nemorosa Lumn. and C. cuprina (Sandor ex Heuffel) Nendtvich ex A. Kerner, have also been referred to the northern taxon. Schultz (1871) thought that he had shown C. nemorosa to be synonymous with C. loliacea Schkuhr non L. and therefore (in his eyes) with his own C. pairaei of which it might, he thought, be the correct prior name. But in BM there is a specimen from the herbarium of J. J. Roemer (1763-1819) inscribed by him ‘Carex 32 nemorosa Lumnitzer ... Ab ipso accepi beatae memoriae D. Lumnitzer’; and this plant appears to be an immature specimen of that currently known as C. otrubae. It is certainly neither C. muricata L. nor its western relative. The name ‘C. cuprina (Sandor) Nendtvich’ was adopted by Kreczetovicz (1935) as the correct name of the northern taxon after he had taken C. muricata to be the correct name for C. echinata. Nendtvich’s name was not accompanied by a description and was therefore discarded by Hylander (1966) as a nomen nudum; but it was based on a Hungarian specimen in Sandor’s herbarium which had been validly published, with a description, as C. nemorosa B cuprina Sandor. Sandor’s specimen, however, as reported by Rauschert (1973), has lately been determined by Soo as C. otrubae, of which C. cuprina would accordingly appear to be the correct prior name. Kikenthal (1909, p. 156) is justified in his note: ‘Alia quoque nomina non pauca quoad synony- miam semper in dubio remanebunt, cum species supra laudatae ante cl. Fr. Schultz, Etude sur quelques Carex (1868), Flora LIII. (1870) et LIV. (1871) non certe distinctae sint, insuper confusio cum C. vulpina timenda.’ Both C. nemorosa Lumn. and ‘C. cuprina (Sandor) Nendtvich’ are included in Kiikenthal’s list of nomina dubia and both should now disappear from the synonymy of the C. muricata aggregate. The history of the naming of the western taxon is even more complicated. In June 1868 Michel Paira, of Geudertheim in Bas-Rhin (Alsace), sent to F. W. Schultz specimens of a sedge that grew plentifully in his neighbourhood. Schultz confessed that till then he had overlooked it, recognized that it differed both from C. spicata (which he then knew as C. muricata) and from C. divulsa, and proposed to describe it as a new species under the name C. pairaei in honour of its discoverer. As ‘Paira’ and ‘Pairaeus’ are equally acceptable Latin forms of Paira’s name, there is no need to make, as Hylander did, an orthographical correction to ‘pairae’. Meanwhile there was further correspondence between the two men, in the course of which Paira said that he had taken his sedge to be the same as C. loliacea sensu Schkuhr non L.; and to show what he meant he sent Schkuhr’s illustrations (Schkuhr 1801) to Schultz who did not himself NOMENCLATURE OF THE CAREX MURICATA L. AGGREGATE 61 possess the book. Schultz (1868a) thereupon recorded in print: ‘Die [in Schkuhr] Tafel Ee No 91, als C. loliacea abgebildete Pflanze entspricht vollkommen der von Hrn. Paira erhaltenen Carex’; and again (1868b): ‘Ayant comparé le dessin de Schkuhr et les échantillons, j’ai reconnu sans hesiter que la resemblance était exacte.’ Nelmes (1947) took C. pairaei to be the western taxon; and well he might, for Schultz’s full Latin description (1868b) contains at least two sentences (‘habitat ... in montibus graniticis’ and “Carex muricata [i.e. C. spicata Huds] differt ... tempore florenti, in iisdem locis, multo preco- ciori’) that apply only to the western taxon and not at all to the northern. By 1959 Nelmes’ co!- league, A. A. Bullock, had called his attention to the fact that Schultz, in his first paper (1868a), had equated his C. pairaei with C. loliacea sensu Schkuhr and that, as the actual specimen described by Schultz could not be certainly identified, Schkuhr’s illustration was the only type upon which Schultz’s name could be based. As Schkuhr’s plant was in Nelmes’ view the northern taxon, it followed that C. pairaei, based upon it, could by the International Code of Botanical Nomenclature be regarded only as yet another synonym for that taxon (= C. muricata L. sensu stricto). In that case the western taxon still lacked a name, and Nelmes (1959) supplied it by describing the western plant as C. bullockiana Nelmes. Now it is true that Schultz’s first paper (1868a) is headed ‘Carex muricata var. 8 Schkuhr (C. loliacea Schk., non Linné) als gute Art aufgestellt’ and that he positively identified Paira’s sedge with Schkuhr’s illustration, though he did question whether the enlargement of the utricle can be a true representation. Yet in the same paper he made it very clear that what he set out to describe were the specimens sent to him by Paira, and he announced his intention to do two things: to issue a full description of Paira’s plant, and to preserve the specimens as vouchers ‘in der 12 Centurie meines herbarium normale.’ In this first paper (Schultz 1868a) there is in fact no adequate description or reference to one, so that C. pairaei is there a nomen nudum. One must conclude that Schkuhr’s illustration (even if the discrepancy in the utricle is disregarded) was not intended by Schultz to be a type; as explained above, he intended Paira’s specimens to be the types. It may also be added that the source of Schkuhr’s plant is unknown. He obtained it from an unnamed friend and the illustration did not indicate habitat or time of flowering. The only diagnostic characters that could be shown in a plate are the degree of divergence of the ripe utricles, their shape, and the colour of the glumes. On the first point Schkuhr’s plate is ambivalent: the utricles are somewhat divergent but not more so than can be found in the western taxon. The illustration of the utricle is, as Schultz remarked, unsatisfactory: it does not correspond with that of either taxon. On the last point the evidence varies from copy to copy of the book according to the individual colourist responsible. For example, the copy in the Department of Botany, British Museum (Natural History), shows dark glumes contrasting with grey-green utricles—the northern taxon. The two copies owned by the Linnean Society show glumes and utricles that are almost concolorous—the western taxon. It is really not possible to say whether the plant illustrated is the one or the other. Valid publication of C. pairaei was achieved, as Schultz had promised, in his Etude sur quelques Carex (Schultz 1868b). This contains a full Latin description, together with a drawing by Paira, both of which can only refer to the western taxon, while of the ‘quelques cents exemplaires’ received from Paira (Schultz 1868b) those distributed to a number of major herbaria are certainly that plant. They constitute genuine type-material, and I choose as lectotype the sheet (‘déc. et rec. M. Paira’) in K dated 27 June and 9 July 1868. C. pairaei can therefore be retained as the name of the western taxon, while C. bullockiana, which (perhaps because it was published in a Russian journal) has never received much currency, becomes a later synonym. In fact I am of the opinion, which is shared by Mr A. O. Chater, that the two taxa, while clearly distinct, do not merit more than subspecific rank. In that case the name of the northern taxon would be C. muricata L. subsp. muricata, and that of the western taxon C. muricata L. subsp. pairaei (F. W. Schultz) Celak. It is ironical that the correctness of Celakovsky’s combination is accidental, for his C. muricata is certainly not Linnaeus’ and his C. pairaei is probably not Schultz’s. C. muricata subsp. pairaei is not uncommon in Britain. C. muricata subsp. muricata is a rare plant there, and has at present been refound in only one of the four localities recorded by Nelmes (David & Kelcey 1975). A plant apparently related to C. spicata, but of greater stature and with widely interrupted inflorescences, was described as C. muricata Race II lumnitzeri by Rouy (1912). Rouy noted that it 62 R. W. DAVID has ‘écailles et utricules concolores, d’un vert blanchatre.’ This is more a character of C. leersii F. W. Schultz (see below) and Rouy may have been referring to that taxon, though he also listed C. muricata Race III leerseana, with uncharacteristic ‘écailles + brunatres.’ C. lumnitzeri was, however, accorded specific rank by Kreczetovicz (1935), who reported it to be widespread in western Europe and equated it with C. contigua var. y longissima Tauscher ex Vollmann (Vollmann 1903). Kiikenthal (1909) followed Vollmann in giving the plant this varietal rank but cited only a restricted distribution (Hungary). In Britain I have seen only two plants (in a single locality) that answer to the description of C. lumnitzeri, and there was some likelihood that they were hybrids between C. spicata and C. otrubae. A specimen from Tauscher’s own herbarium, collected by him on 26 July 1869 and now in K (ex herb. Churchill), has no mature fruit and looks as if it might also be of this hybrid origin. The one member of the aggregate whose name has never really been in doubt is C. divulsa Stokes (published in 1787), though for a time the name was given a later authority, C. divulsa Good. (published in 1794), which allowed the problematical C. virens Lam. (published in 1791) to qualify as a claimant for priority; and the plant, like all other members of the aggregate, has sometimes been reduced to subspecific or varietal rank. Another taxon that has been put forward as the plant intended by Lamarck under his C. virens is the one later known as C. leersii F. W. Schultz. Lamarck’s description is, however, quite in- adequate and his specimen was reported by Winter (1870) to be even then in such poor condition that no conclusion whatever could be drawn from it. One may hope that the last to be heard of C. virens is the requiescat pronounced by Vollmann (1903) on this ghost-name that has troubled taxonomic discussion for so long and to such little effect. C. leersii F. W. Schultz is almost certainly what Hoppe understood by C. muricata and illustrated under that name in Sturm (1833). The description by Schultz (1870) is, however, the first in which the plant was clearly differentiated, though the name C. J/eersii is, as a species, invalidated by Willdenow’s earlier use of it as an illegitimate substitute for C. echinata Murray. Nelmes (1947) followed Kreczetovicz (1935) and Kiikenthal (1911) in identifying Schultz’s sedge with C. poly- phylla Kar. & Kir. (Karelin & Kirilov 1841). Kreczetovicz seems himself to have had two opinions about this, for in a note headed ‘Herbarium Florae Asia Medie ab Universitate Asiae Mediae editum Fasc. XXIII Julio a 1934’ and attached to a specimen in K of C. polyphylla from its original locality in the Targabatai mountains, he wrote: ‘Cette espéce appartient a une série avec le C. Leersii F. Schultz de l’Europe occidentale duquel elle se distingue par les épillets fortement divariqués, de plus larges feuilles a ligule aplatie et une autre forme des utricules.’ Unfortunately Kreczetovicz did not particularize the difference in the shape of the utricles; but those of the specimen are very large (over 5 mm long) and their form, with rounded base and narrowly tapered beak, is more akin to that found in C. spicata than to the smaller (4-5 mm), neater ‘diamonds’ of C. leersii. The true C. polyphylla is not in Britain and probably not in Europe. If Schultz’s sedge is to have specific rank, its name must be C. leersiana Rauschert, but it may be no more than a subspecies of C. divulsa and would then be subsp. Jeersii (Aschers. & Graebn.) W. Koch. The dividing line between these two taxa is the most difficult to draw in the whole taxo- nomy of the aggregate. Lastly there is C. chabertii F. W. Schultz, actually named ‘Chaberti’ in Schultz (1871). Schultz’s description has not received proper attention, and the name, from the time of Kneucker’s col- lections (late nineteenth century) onwards, has been incorrectly applied to a number of plants intermediate between C. divulsa and C. leersiana and possibly of hybrid origin, as they do not seem to mature fruit. The true C. chabertii, as seen in specimens in K that originated from Schultz, is characterized by a comparatively short and closely-packed inflorescence of long (over 5 mm), upward-pointing (i.e. not patent) utricles, and appears to have been reported only from the Rhine- land and Palatinate, Germany, from the département of Dréme and the Pyrenees, France, and possibly from Bulgaria. As it is said to be a very local plant of woods on dry and stony ground in the mountains it is unlikely to occur in Britain. Hybrids within the aggregate, and between members of the aggregate and sedges in other sections of subgenus Vignea, are reported from time to time in Britain and elsewhere. The majority if not quite all of them are more probably atypical plants of one or other of the taxa; and of the specimens that I have seen so far, either in the field or in herbaria, the only ones of whose hybrid origin I am convinced have been the cross between C. divulsa and C. otrubae. This is occasionally NOMENCLATURE OF THE CAREX MURICATA L. AGGREGATE 63 found in southern England. The possibility of other hybrids has been indicated under C. lumnitzeri and C. chabertii above. ACKNOWLEDGMENTS This paper is part of a wider study of the C. muricata aggregate, in which I am collaborating with Mr J. G. Kelcey; and it owes much to discussions with him. For general guidance in its prepara- tion I am deeply indebted to Mr A. O. Chater, Mr A. C. Jermy, and especially to Mr P. D. Sell who has also set in order the synonymy and references. For particular advice on the difficult nomenclatural problem of Carex pairaei I am grateful to Mr J. E. Dandy, Mr R. Ross, and Dr W. T. Stearn. REFERENCES ANDERSSON, N. J. (1849). Cyperaceae Scandinaviae, p. 65. Stockholm. ASCHERSON, P. F, A. & GRAEBNER, K. O. P. P. (1902). Synopsis der mitteleuro pdischen Flora, 2(2): 38-43. Leipzig. BRITTEN, J. (1907). (Editorial note). J. Bot., Lond., 45: 163. BRIQUET, J. (1910). Prodrome de la flore corse, 1: 199. Geneva, Basle and Lyons. CrEPIN, F. (1859). Notes sur quelques plantes rares ou critiques de Belgique. Bull. Acad. r. Belg. Cl. Sci., sér. 2, 7: 112. Danpy, J. E. (1958). List of British vascular plants, p. 155. London. Davin, R. W. & KEtcey, J. G. (1975). Carex muricata sensu Nelmes and Carex bullockiana Nelmes. Wat- sonia, 10: 412-414. De LANGHE, J. E. (1944). Sur le groupe du Carex muricata L. en Belgique. Bull. Soc. r. Bot. Belg., 76: 47-48. FERNALD, M. L. (1917). Some new or critical plants of eastern North America. Rhodora, 19: 154. HyYLANpDER, N. (1966). Nordisk Karlvdxtflora, 2: 84-88. Stockholm. KARELIN, G. S. & KiriLov, I. P. (1841). Enumeratio plantarum anno 1840 in regionibus Altaicis et con- finibus collectarum. Byull. Mosk. Obshch. Ispyt. Prir., 14: 859. Kocu, W. D. J. (1837). Synopsis florae germanicae et helveticae, p. 751. Frankfurt. Kovacs, B. (1910). Carex echinata Murr. (1770) még sem mas mint C. stellulata Good. (1794). Magy. bot. Lap., 9: 126-131. KRECZETOVICZ, V. I. (1935). Carex: Muehlenbergia, in Komarov, V. L., ed. Flora URSS, 3: 153-156. Leningrad. KUKENTHAL, G. (1905). Carex fulva Good. und Carex echinata Murr. Allg. bot. Z., 11: 45. KUKENTHAL, G. (1909). Carex Sect. 22, Muehlenbergianae Tuckerm., in ENGLER, H. G. A., ed. Das Pflanzen- reich, 38 (1V.20) (Cyperaceae-Caricoideae): 152-165. Leipzig. KUKENTHAL, G. (1911). Cyperaceae Sibiriae subfamilia Caricoideae (Pax.). Russk. bot. Zh., 3-6: 54-56. Linnaeus, C. (1753). Species Plantarum, ed. 1, p. 974. Stockholm. MACKENZIE, K. K. (1923). Notes on Carex, XII, 2. Bull. Torrey bot. Club, 50: 346-347. MACKENZIE, K. K. (1935). North American flora, 18(1): 41-62. New York. Murray, J. A. (1870). Prodromus designationis stirpium Gottingensium, p. 76. Gottingen. NELMES, E. (1942). Notes on British Carices, VI. J. Bot., Lond., 80: 107-109. NELMES, E. (1947). Two critical groups of British sedges. Rep. botl Soc. Exch. Club Br. Isl., 13: 99-105. NELMES, E. (1959). De Caricibus Britannicis notae, IX. Bot. Mater. Gerb. bot. Inst. V. A. Komarova, 19: 74-78. NEUMAN, L. M. (1901). Sveriges Flora (Fanerogamerna), p. 716. Lund. RAUSCHERT, S. (1973). Nomenklatur der Fern- und Blutenpflanzen Deutschlands, III. Reprium nov. Spec. _ Regni veg., 83: 660. Rouy, G. (1912). Flore de France, 13: 412-413. Paris. Scukunr, C. (1801). Beschreibung und Abbildung der Arten von Riedgrdsern. Wittenberg. SCHULTZ, F. W. (1868a). Carex muricata var. B Schkuhr (C. loliacea Schk. non Linné) als gute Art auf- gestellt. Flora, Jena, 51: 302-303. SCHULTZ, F. W. (1868b). Etude sur quelques Carex. Haguenau. SCHULTZ, F. W. (1870). Bemerkungen tiber einige Carex. Flora, Jena, 53: 458-459. ScHULTZz, F. W. (1871). Zusatze und Verbesserungen zu den, in der Flora 1870, Nr 29, Seite 458 bekannt gemachten Bemerkungen iiber einige Carex ... Flora, Jena, 54: 21-32. \ 64 R. W. DAVID — SturM, J. (1833). Deutschlands Flora, fasc. 61. Niirnberg. VOLLMANN, F. (1903). Der Formenkreis der Carex muricata und seine Verbreitung in Bayern. Denkschr. K. bayer. bot. Ges. Regensb., n.s. 2, 8: 55-90. WINTER, F. (1870). [Review of T. F. Marsson’s Bemerkungen tiber die Flora von Neu-Pommern.] Flora, Jena, 53: 455. APPENDIX MAIN SYNONYMS OF TAXA CONSIDERED This list includes those names that have significantly affected taxonomic discussion or that mark clear stages in the taxonomic argument. Many nomina dubia, and such names as C. orsiniana Tenore and C. /itigiosa Chaubard, which appear to be merely the result of error, have been omitted. Names not validly published are enclosed in square brackets. CAREX SPICATA Hudson, Fl. Angl., 349 (1762) C. contigua Hoppe in Sturm, Deutschl. Fl., fasc. 61 (1833) [C. muricata sensu Koch, Syn. Fl. germ., 751 (1837) pro parte, non L. (1753)] C. muricata var. genuina subvar. incrassata Crépin, Bull. Acad. r. Belg. Cl. Sci., sér. 2,'7: 112 (1859) C. muricata var. virens subvar. incrassata Crépin, Bull. Acad. r. Belg. Cl. Sci., sér. 2, 7: 112 (1859) [C. muricata sensu Aschers. & Graebn., Syn. mitteleur. Fl., 2(2): 38 (1902), non L. (1753)] CAREX LUMNITZERI (Rouy) V. Krecz. in Komarov, Fl. URSS, 3: 154 (1935) C. contigua var. longissima Tauscher ex Vollmann, Denkschr. K. bayer. bot. Ges. Regensb., 8: 81 (1903) C. muricata race lumnitzeri Rouy, Fl. Fr., 13: 412 (1912) C. muricata subsp. lumnitzeri (Rouy) Sood in Sod & Jav., Magyar Nov. Kéz., 2: 903 (1951) [C. spicata subsp. lumnitzeri (Rouy) Soo in Soo & Karpati, Magyar Fl., 724 (1968) sine basion.] C. spicata subsp. lumnitzeri (Rouy) So6, Reprium nov. Spec. Regni veg., 85: 438 (1974) CAREX MURICATA L. subsp. MURICATA C. muricata L., Sp. Pl., 974 (1753) [C. loliacea sensu Schkuhr, Riedgrdser, 22, tab. Ee no. 91 (1801) quoad tab. exclud. descript. et syn.] [C. echinata sensu Kiikenthal in Engler, Pflanzenreich, 38 (IV.20): 160 (1909), non Murray (1770)] [C. cuprina sensu V. Krecz. in Komarov, Fl. USSR, 3: 155 (1935), non Sandor ex Heuffel (1862)] C. pairae subsp. borealis Hyl., Nord. Karlvaxtfl., 2: 386 (1966) subsp. PAIRAEI (F. W. Schultz) Celak., Prodr. Fl. Bohm., 4: 731 (1881) [C. pairaei F. W. Schultz, Flora, Jena, 51: 302 (July 1868) nom. nud.) C. pairaei F. W. Schultz, Etude quelques Carex, 9 (September 1868) C. bullockiana Nelmes, Bot. Maer. Gerb. bot. Inst. V. A. Komarova, 19: 77 (1959) CAREX POLYPHYLLA Kar. & Kir., Byull. Mosk. Obshch. Ispyt. Prir., 14: 859 (1841) CAREX LEERSIANA Rauschert, Reprium nov. Spec. Regni veg., 83: 660 (1973) [C. muricata sensu Hoppe in Sturm, Deutschl. Fl., fasc. 61 (1833), non L. (1753)] C. leersii F. W. Schultz, Flora, Jena, 53: 459 (1870), non Willd. (1787) C. muricata var. leersii Kneucker in Seubert-Klein, Excurs. Fl. Baden, 52 (1891) C. muricata subsp. leersii Aschers. & Graebn., Syn. mitteleur. Fl., 2(2): 40 (1902) C. echinata subsp. leersii Kiikenthal in Engler, Pflanzenreich, 38 (1V.20): 161 (1909) C. pairaei var. leersii Kiikenthal, Russk. bot. Zh., 3-6: 55 (1911) C. divulsa subsp. leersii (Aschers. & Graebn.) W. Koch, Mitt. bad. Landesver. Naturk., 11: 259 (1923) [C. polyphylla sensu V. Krecz. in Komarov, Fl. URSS, 3: 155 (1935), non Kar. & Kir. (1841)] CAREX DIVULSA Stokes in With., Arr. Br. Pl., ed. 2, 2: 1035 (1787) ? C. virens Lam., Encycl. Méth. Bot., 3: 384 (1792) C. muricata B divulsa (Stokes) Wahlenb., K. svenska Vetensk-Akad. Handl., 24: 153 (1803) [C. guestphalica Boenn., Vat. Sem. Hort. Monast. (1829) nom. nud.] NOMENCLATURE OF THE CAREX MURICATA L. AGGREGATE 65 Vignea guestphalica Boenn. ex Reichenbach, F/. Germ. Excurs., 130 (1830) Carex guestphalica (Boenn. ex Reichenbach) C. F. Lang, Flora, Jena, 26: 147 (1843) CAREX CHABERTII F. W. Schultz, Flora, Jena, 54: 21 (1871) C. virens var. duriaei F. W. Schultz, Grundz. Phyt. Pfalz, 161 (1863) C. duriaei (F. W. Schultz) F. W. Schultz, Flora, Jena, 53: 458 (1870), non Steudel (1841) C. divulsa subsp. chabertii (F. W. Schultz) Aschers. & Graebn., Syn. mitteleur. Fl., 2(2): 42 1902 (Accepted April 1975) BSBI ABSTRACTS ABSTRACTS FROM LITERATURE RELATING TO THE VASCULAR PLANTS OF THE BRITISH ISLES Compiled by D. H. KENT. Published annually Includes abstracts of botanical papers published in many journals throughout the Temperate World. Subject headings include History, Biography, Floras and Catalogues, Chromosome Surveys, Systematics, etc. The Systematic Section, which is the bulk of the periodical, is arranged alpha- betically under families and genera. THE TARAXACUM FLORA OF THE BRITISH ISLES By A. J. RICHARDS A Supplement to Watsonia, Journal and Proceedings of the Botanical Society of the British Isles, vol. 9, 1972 The genus Taraxacum, commonly known as Dandelions, has been largely neglected in the British Isles, although it is of botanical, ecological and economic importance. Most accounts of the genus in this country consider only four species, which are too general in scope to be of much service. In the present account the author shows that, due to the apomictic (non-sexual) means of reproduction usual in the genus, a much larger number of clear-cut species can be recognised in Britain, 132 in all, and that these demonstrate characteristic ecological and distributional behaviour. A substantial introduction, with notes on the taxonomic history, biology, genetics, taxonomy, distribution, identification, cultivation and collection of the British species, is followed by a key to the species. In the main account, each Species is treated individually, with nomenclature, a full description, breeding behaviour, ecology, British distribution (including a vice-county list), European distribution and general notes and comments repeated in each case. Each species is illustrated with line drawings and there is a full index. Although our knowledge of the British species of the genus is still incomplete, it is hoped that this work will form a standard reference for a number of years and thus should be valuable to Universities, Colleges, Field Stations, Museums and to all botanists interested in the higher plants of the British Isles. Details from E. W. CLASSEY LTD, PARK ROAD, FARINGDON, BERKSHIRE, SN7 7DN Watsonia, 11, 67-74 (1976). 67 Short Notes DIPSACUS STRIGOSUS WILLD. IN CAMBRIDGESHIRE, V.C. 29 Dipsacus strigosus Willd., a native of southern Russia and western Asia, was reported as an alien in Britain by Hansen (1963), and a later note by Gerrans (1964) tabulated the characters that separate it from D. pilosus L., which it closely resembles. D. strigosus is most easily distinguished from D. pilosus by the capitulum size, 30-45(-60)mm wide (15—25mm in D. pilosus) and the length of the receptacular bracts, 15—20(—30)mm (7—22mm in D. pilosus). These measurements are from Cambridgeshire material. Hansen’s records show that the plant has occurred as an alien in the vicinity of Oxford, Cambridge, Kew and Bristol, which suggests that it originated from the Botanic Gardens in the first three localities. Examination of material in the Cambridge University Herbarium (CGE) reveals three additional records from v.c. 29: ‘Hinton, Cambridge 18 July 1828 J. S. Henslow’ (as D. pilosus) ‘Waste ground W. of Botanical Laboratory, Cambridge 18 July 1921 A. J. Crosfield’ (as D. pilosus) ‘Churchyard, Little St. Mary’s church, Cambridge 1967 S. M. Walters’ (as D. strigosus). Stock from this locality is now growing in the Cambridge Botanic Garden; otherwise the plant has not apparently been grown there in recent years. In addition to these, in July 1974, J. O. Mountford found six plants of what proved to be D. strigosus on an earth tip on the west side of the Gogmagog golf course, 34 miles south-east of Cambridge. Crosfield’s record above ties in with two given by Hansen: “Waste ground near Botanic Garden, Cambridge 1863 F. J. Hanbury’ (BM) and ‘Waste ground, Cambridge 1908 R. S. Adamson’ (BM); and also with a specimen collected by N. D. Simpson in 1913 on Coe Fen (BM). Coe Fen lies directly west of the Cambridge Botanic Garden. These records suggest that the plant has occurred as an alien in the south-western part of the City of Cambridge over a period of at least 112 years. Little St Mary’s, where the plant was first recorded by W. B. Gourlay in 1949 (as D. pilosus—in card index of Cambridgeshire Flora in CGE), still supported the species in the less tended parts of the churchyard in 1974. The Hinton record is the earliest so far traced for this species in the British Isles. Recent searches in the field have revealed that the plant still occurs in Lime Kiln Close, Cherry Hinton (24 miles south-east of Cambridge city centre), presumably the area that Henslow referred to. In 1972 and 1973 there was one large flowering plant, in 1974 five flowering plants were found, and 23 rosettes were counted in February 1975. This increase is best interpreted as a response to the opening up of the habitat (heavily cloaked with Clematis vitalba) by children playing in the pit and making new paths. Thus D. strigosus may have been established at Cherry Hinton for 147 years, though recorded as D. pilosus for much of that time. It certainly justifies a place in our Floras. Perring et al. (1964), who did not distinguish D. strigosus, listed five recent records for ‘D. pilosus’. Two of these (Little St Mary’s and Cherry Hinton) are D. strigosus and of the others only that at Hildersham (9 miles south-east of Cambridge) can be confirmed as true D. pilosus by a specimen in CGE. D. pilosus has recently been rediscovered in Bottisham Park (7 miles north-east of Cambridge). It will be interesting to record the persistence of D. strigosus in its present sites in v.c. 29 and any further spread. Does it persist elsewhere, e.g. near Oxford, Kew or Bristol? Is it still un- recognized in other sites? ACKNOWLEDGMENTS I should like to thank Miss M. B. Gerrans, Mr P. D. Sell and Dr S. M. Walters for help in preparing this note. 68 SHORT NOTES REFERENCES GERRANS, M. B. (1964). Dipsacus. Proc. bot. Soc. Br. Isl., 5: 231. HANSEN, A. (1963). Dipsacus strigosus Willd. Proc. bot. Soc. Br. Isi., 5: 123. PERRING, F. H., SELL, P. D., WALTERS, S. M. & WuITEHOUSE, H. L. K. (1964). A Flora of Cambridgeshire. Cambridge. A. C. LESLIE RUBUS PLATYACANTHUS MUELL. & LEFEV. AND ITS ALLIES IN BRITAIN In Newton (1974) I gave specific rank to Rubus carpinifolius var. laxus Sudre under the name R. laxissimus, based on A. Ley’s gathering from Mitcheldean Meend (v.c. 34) named by Sudre and issued as his Rubi rari no. 3. At the same period Professor H. E. Weber was researching the R. carpinifolius group of brambles, during which he referred several of the sheets in BM cited by me as examples of R. laxissimus to R. platyacanthus Muell. & Lefev. Further, he was of the opinion that many British and German specimens named R. carpinifolius by Focke, Rogers and others in the past should also be called R. platyacanthus, claiming in support the view of A. Neumann, a distinguished German batologist, who had verified R. platyacanthus in its locus classicus in France. Weber (1973) has renamed as R. adspersus Weihe ex Weber the plant formerly known as R. carpinifolius Weihe & Nees, since this name had previously been used (by J. & C. Presl) for a different taxon. In this article he described the features distinguishing the two taxa, emphasizing the stouter, denser stem-prickles, the ovate, gradually acuminate terminal leaflet and gland-dotted bracts in the panicle as characters of R. adspersus, and the elliptical, shortly cuspidate leaflets with rounded rather than subcordate bases, the sparser, rather finer stem armature and the eglandular panicle-bracts in R. platyacanthus. In the light of these investigations, and following inspection in company with Professor Weber of both species growing in Germany, I have re-examined many British sheets of R. laxissimus and R. carpinifolius with the holotype of R. platyacanthus (Lefévre, Forét de Retz, France, 1856, as R. nitidus, hb. P. J. Miiller in LAU) in front of me. Most of the British material named R. carpinifolius is indeed exactly R. platyacanthus, and despite slight variations in terminal leaflet- shape and indumentum in R. laxissimus, and the consistently broad panicles, I am now convinced that these features are not sufficient to support a distinct taxon at specific rank; R. laxissimus is also to be included within R. platyacanthus. What is the relative status of R. platyacanthus and R. adspersus in Britain? Examination of sheets in several herbaria indicates that R. platyacanthus is by far the more widespread and common of the two; I have seen specimens either in the field or herbarium from v.c. 11-19, 21-24, 27, 30, 34-36, 38-40, 43, 53-59 and 62-64. R. adspersus appears to be confined to the areas of Flitwick Moor (v.c. 30), Milford Heath (v.c. 17, the specimens of ‘R. carpinifolius flore roseo’ collected and issued as Set of British Rubi no. 28), Holme and Woodwalton Fens (v.c. 31), and the ancient Knutsford Heath (v.c. 58); doubtless it is present in other localities, but before this can be established a more comprehensive review of the available specimens will be necessary. In the herbarium, specimens of the two are sometimes difficult to distinguish, particularly on badly collected or incomplete sheets. In the field, however, they can often be distinguished at a glance: R. platyacanthus exhibits a uniformly fresh green appearance, whereas R. adspersus has a characteristic yellowish-green cast (at least in the open) due to the reddish-spotted pigmentation of stems and panicles reflected in the name adspersus; also the petals are pale pink or pink-tinted, unlike those of R. platyacanthus which are nearly always pure white. REFERENCES Newton, A. (1974). Five brambles from South Wales. Nature Wales, 14: 24-35. Weser, H. E. (1973). Zur Systematik und Nomenklatur des Rubus adspersus. Osterr. bot. Z., 122: 275-282. A. NEWTON SHORT NOTES 69 NOTES ON SOME WOOL-ALIEN CYPERACEAE FROM BLACKMOOR, N. HANTS. The Cyperaceae form a huge natural family, with some 90 genera comprising 4,000 species, and are especially well represented in both temperate and warmer regions of the world where sheep are reared. Very few species have been recorded as wooi-aliens in the British Isles, e.g. one species of Cyperus by Hayward & Druce (1919), two species of Cyperus by Lousley (1961), one species each of Cyperus and Carex by Dony (1969) and two species of Carex by Jermy & Tutin (1968), although on the Continent Probst (1949) listed about 20 species, and several other more recent records are scattered through the literature. The rather surprising lack of records is probably due to the ease with which different species in the field may be overlooked, because in the early vegetative stage many Cyperaceae look alike, and also to the difficulty in getting mature specimens for identification. During the past six years (1969-1974) over 30 species have been collected from Blackmoor, N. Hants (v.c. 12), and these are listed below together with a few earlier records. Relatively few species were seen flowering in the field, the remainder having been cultivated under glass to protect them from frost. Many proved difficult to grow successfully, and took several years to produce even one inflorescence. For satisfactory determinations it is essential to obtain mature fruiting material with good rootstocks, especially in Cyperus and Carex. Several plants have not yet flowered and remain undetermined. A number of Carex species which are common British natives have been included, since these occurred regularly with undoubted wool-aliens. The geographical distributions of the species are mostly taken from the monographs by Kiikenthal (1909, 1935), but it is likely that they have been introduced into many other countries, e.g. Australia and S. America. For Carex, the more recent accounts by Nelmes (1944), on Australian, and by Moore & Edgar (1970), on New Zealand species, have proved very useful. Voucher specimens are in my herbarium, and also some (K) at Kew. CYPERUS L. 550 species . clarus 8. T. Blake Australia 1971 (K) det. SSH . congestus Vahl Originally S. Africa, widespread Frequent (K) det. SSH . dactylotes Benth. Australia 1973 det. SSH . eragrostis Lam. (C. vegetus Willd.) Originally S. America, widespread Frequent det. SSH esculentus L. Widespread 1973 conf. SSH gunnii Hook. f. Australia 1971, 1972 (K) conf. SSH longus L. var. tenuiflorus (Rottb.) Kiik. Widespread AB and MMcCW 1973 det. JEL luzulae (L.) Retz. S. America 1973 (K) conf. SSH ovularis (Michx.) Torr. U.S.A. 1971 det. TBR rigidifolius Steud. E. Africa 1973 (K) conf. SSH rotundus L. Cosmopolitan 1971 det. SSH rutilans (C. B. Clarke) Maiden & Betche Australia ACL, TBR 1969 (K) det. DMN sporobolus R. Br. N. Australia 1973 conf. SSH tenuis Schwartz S. and tropical Africa, Mexico, S. America 1971 (K) det. SSH ustulatus A. Rich. New Zealand 1971 det. SSH vaginatus R. Br. Australia MMcCW 1964; 1970, 1973 conf. SSH KYLLINGA Rottb. 60 species K. brevifolia Rottb. Widespread in warm regions 1971, 1973 det. SSH K. erecta Schum. Africa 1972 (K) det. SSH K. odorata Vahl Africa and America AB 1973 conf. SSH SCIRPUS L. 300 species S. holoschoenus L. Europe, Asia and the Mediterranean MMcCW c 1969 S. nodosus Rottb. S. Africa, S. America, Australia and New Zealand TBR, ACL 1971 (K) conf. SSH ELEOCHARIS R. Br. 200 species E. nodulosa (Roth) Schultes S. America 1971 det. TBR BULBOSTYLIS Kunth 100 species B. humilis Kunth S. Africa MMcCwW 1964; 1972, 1973 (K) conf. SSH FEO OO SAA AO ao Os 70 SHORT NOTES B. striatella C. B. Clarke E.andS. Africa 1969, 1973 det. EJC CAREX L. Over 1,500 species C. appressa R. Br. Australia, New Zealand, New Guinea and New Caledonia MMcCW 1968; 1971 C. devia Cheesem. New Zealand 1971 det. SSH C. flacca Schreb. Europe, W. Asia, N. America, introduced in New Zealand Frequent C. flagellifera Col. New Zealand 1972 det. SSH C. hirta L. WN. temperate regions 1971 det. EJC C. hubbardii Nelmes Queensland 1972 conf. SSH C. inversa R. Br. Australia and New Zealand 1971 C. muricata L. N. temperate regions Frequent conf. SSH C. ovalis Good. N. temperate regions, introduced in New Zealand Frequent C. secta Boott New Zealand MMcCW 1964 det. SSH C. solandri Boott New Zealand MMcCW 1970 det. SSH C. tereticaulis F. Muell. Australia MMcCW 1968; 1972 conf. SSH C. virgata Sol. ex Boott New Zealand MMcCwW 1968; 1969 (K) det. SSH C. vulpinoidea Michx. N. America, Colombo, naturalized in Europe 1968 (K) det. SSH Key to initials used in the list: AB _ A. Brewis EJC E. J. Clement SSH — S. S. Hooper ACL A.C. Leslie JEL J. E. Lousley DMN D. M. Napper TBR T. B. Ryves MMcCW M. McCallum Webster ACKNOWLEDGMENTS I am grateful to Miss S. S. Hooper for identifying many of the specimens, to Lady Anne Brewis for permission to visit Blackmoor Fruit Farm, and to Miss M. McCallum Webster for giving me some of her interesting collections of Carex. Some of the species listed were also collected by other members of the London Natural History Society. REFERENCES Dony, J. G. (1969). Additional notes on the flora of Bedfordshire. Proc. bot. Soc. Br. Isl., 7: 523-535. HAYWARD, I. M. & Druce, G. C. (1919). The adventive flora of Tweedside. Arbroath. JermMy, A. C. & TuTIN, T. G. (1968). British sedges. London. KUKENTHAL, G. (1909). Das Pflanzenreich, 38 (IV, 20). Cyperaceae: Caricoideae. Leipzig. KUKENTHAL, G. (1935). Das Pflanzenreich, 101 (IV, 20). Cyperaceae: Scirpoideae-Cypereae. Leipzig. LousLey, J. E. (1961). A census list of wool aliens found in Britain, 1946-1960. Proc. bot. Soc. Br. Isl., 4: 221-247. Moore, L. B. & Epaar, E. (1970). Flora of New Zealand, 2. Wellington. NELMES, E. (1944). A key to the Australian species of Carex (Cyperaceae). Proc. Linn. Soc. Lond., 155: 277-285. Prost, R. (1949). Wolladventivflora Mitteleuropas. Solothurn. T. B. Ryves NOTES ON SOME WOOL-ALIEN MALVACEAE The Malvaceae form a moderately large family (1,000 species in 75 genera) in tropical and temperate regions. Many species have big, handsome flowers and are commonly grown in gardens, including Hibiscus trionum L., which is also one of our most attractive wool-alien plants. Probst (1949) listed 26 species of Malvaceae which were associated with shoddy on the Continent, and 14 species from Britain were listed by Lousley (1961). Most of the latter have recurred in the SHORT NOTES 71 past six years in the two areas at present under review, but the purpose of this note is to give records only of species which were either not included or noted only once in Lousley’s list. Several of them have also been recorded more recently by Dony (1969). Modiola appears to have been previously recorded as a British wool-alien only by Hayward & Druce (1919), from Galashiels 60 years ago. The commonest wool-alien species were Malva parviflora, M. sylvestris and M. neglecta (the two latter, however, possibly being native weeds), followed by Sida spinosa, S. cordifolia and Pavonia. The Sida species hardly ever flowered in the field, although in the exceptionally fine hot summer of 1973 several very large plants of S. rhombifolia flowered freely and set good fruit at Blackmoor. Most of the alien Malvaceae were sensitive to slight or moderate frost, but nearly all the attempts made to grow them on through the winter were abortive, and several new species died before they could be induced to flower and hence identified. Exceptionally, Pavonia and Modiola proved relatively easy to cultivate and grew strongly under glass. The following 12 species were found at Blackmoor, N. Hants., v.c. 12, and Maulden, Beds., v.c. 30. Records are my own unless otherwise stated. ABUTILON Miller Over 100 species A, malvifolium (Benth.) Domin Australia Blackmoor 1973 MALOPE L. 4 species M. trifida L. W. Mediterranean Maulden 1969 E. J. Clement & T. B. Ryves MALVA L. 40 species M. neglecta Wallr. Europe Blackmoor and Maulden Common, perhaps native M. sylvestris L. Europe Blackmoor and Maulden Common, perhaps native MALVASTRUM A. Gray 12 species M. multicaule (Schlecht.) Britton S. America Blackmoor and Maulden Single plants in several years M. peruvianum (L.) A. Gray (?) S. America Blackmoor 1970 det. C. C. Townsend MODIOLA Moench #1 species M. caroliniana (L.)G. Don America, naturalized in Australia, S. Africa and Asia Blackmoor 1970, 1973 PAVONIA Cav. 200 species P. urens Cay. Africa Blackmoor’ A few plants in most years SIDA L. 200 species S. cordifolia L. Tropics Blackmoor A few plants in most years S. rhombifolia L. Tropics Blackmoor 1973, 1974 S. spinosa L. Tropics Blackmoor A few plants in most years UROCARPIDIUM Ulbr. 11 species U. shepardae (Johnst.) Krapov. S. America Maulden 1969 E. J. Clement, J. L. Mason & T. B. Ryves det. C. C. Townsend ACKNOWLEDGMENTS I wish to thank Lady Anne Brewis for permission to visit Blackmoor, Dr and Mrs J. G. Dony for their help in locating good areas to search in Bedfordshire, and various members of the London Natural History Scciety for general assistance in finding specimens. REFERENCES Dony, J. G. (1969). Additional notes on the flora of Bedfordshire. Proc. bot. Soc. Br. Isl., 7: 523-535. Haywarp, I. M. & Druce, G. C. (1919). The adventive flora of Tweedside. Arbroath. LousLey, J. E. (1961). A census list of wool aliens found in Britain, 1946-1960. Proc. bot. Soc. Br. Isl., 4: 221-247. ProsstT, R. (1949). Wolladventivflora Mitteleuropas. Solothurn. T. B. RyYvVEs 72 SHORT NOTES NOTES ON TWO ALIEN VULPIAS IN BRITAIN Townsend (1959) first reported the presence in Britain of Vulpia australis (Steudel) Blom, an alien from South America. This addition to the British list was based on a gathering by C. W. Bannister and C. C. Townsend in 1955 from the docks at Sharpness, W. Gloucs., v.c. 34, of which we have seen material in K and herb. J. E. Lousley. Townsend appears to have identified V. australis by comparing his specimens with material so-determined in K, where there are several sheets from South America of plants agreeing with the Sharpness specimen and labelled V. australis (in particular by the American agrostologist A. S. Hitchcock). Such plants do not, however, agree with the original description nor with the type-specimen of this species. V. australis (Steudel) Blom is based on Festuca australis [Nees ex] Steudel, which was in turn based on Festuca tenella var. « Nees, non Willd. The description given by Nees (1829) mentions few characters which one could use to separate the closely related species in the section Vulpia of the genus Vulpia, but he described the lower glume as half as long as the lowest lemma on the same side, and the upper glume as about as long as the lowest lemma on the same side. Taken in their strict application these features agree very closely with those of V. bromoides (L.) S. F. Gray, but less so with those of other related species or with those of the Sharpness specimen. The only specimen cited by Nees was collected by Sellow at Montevideo and seen by Nees in B. No such specimen exists at B now, but a duplicate of it, sent from B in 1840 (after Nees’ publication), is at K. It is clearly V. megalura (Nutt.) Rydb., a taxon to which Nees’ description does not strictly apply, and quite different from the Sharpness plant. The true identity of V. australis as understood by Nees is thus uncertain, and will be discussed by us more fully in a later paper. The Sharpness specimen is a typical 1epresentative of the plant generally known as V. hybrida (Brot.) Pau or V. broteri Boiss. & Reut., a western Mediterranean species which is widely naturalized in South America, and Townsend’s (1959) description gives a good idea of the differences between it and V. bromoides. Several other British specimens, mostly wool-aliens, have been identified as V. australis since 1959, and the name has been used in the literature (e.g. Lousley 1961). All specimens we have seen so-labelled have been referable to V. broteri, or (presumably representing mis-identifications) to V. bromoides, V. myuros (L.) C. C. Gmel. or V. megalura, all of which occur as aliens in South America. We have seen material of V. broteri from Scilly, v.c. la; N. Hants., v.c. 12; W. Kent, v.c. 16; Beds., v.c. 30; and W. Gloucs., v.c. 34. The correct name for V. broteri (V. hybrida) is, however, Vulpia muralis (Kunth) Nees, a combination based on Festuca muralis Kunth, which was in turn based by Kunth on his own earlier concept of F. myuros L. (V. myuros) as it occurred in South America, but which he came to believe was a distinct species. There is an isotype of F. muralis (from garden walls, Quito, Ecuador) in BM, and it is a perfect match for the European V. broteri. The name F. muralis was published in 1822 and easily predates V. broteri or V. hybrida. Vulpia megalura, the second subject of this note, has been known as a wool-alien species in Britain for many years. It was described (as Festuca megalura Nutt.) from Californian material, and it is very widespread in both North and South America. Nevertheless it is scattered throughout the European and North African range of V. myuros, to which it is very closely related, in places where it is most unlikely to have been introduced (in contrast to its casual status in Britain), and we are convinced that it was introduced to America from the Mediterranean region. The same conclusion has been recently reached by Lonard & Gould (1974). V. megalura differs from V. myuros only in its lemmas, which are ciliate distally. Such variation is found in several species of Vulpia, and we fully agree with Lonard & Gould (1974) that the two taxa are conspecific. Lonard & Gould included V. megalura in V. myuros var. hirsuta Hackel. However, we consider that the rank of forma is more appropriate, and the combination V. myuros f. hirsuta (Hackel) Blom already exists (Blom 1934). But the basionym V. myuros B hirsuta Hackel referred to a Portuguese plant with dorsally hairy (not marginally ciliate) lemmas (Hackel 1880). Plants of this sort are also scattered throughout the range of V. myuros, and we consider them to represent a distinct forma. Therefore we wish to make a new combination for plants previously known as V. megalura: Vulpia myuros (L.) C. C. Gmel. f. megalura (Nutt.) Stace & Cotton, comb. et stat. nov. Festuca megalura Nutt., J. Acad. Nat. Sci. Philadelphia, 2(1): 188 (1847) It is interesting to note that these two plants, V. muralis and V. myuros f. megalura, are both SHORT NOTES 73 Mediterranean taxa which have become naturalized in America, were both first named in America, and have both been re-introduced into Europe as wool-aliens from America. REFERENCES Biom, C. (1934). Uber einige Vulpia-Arten. Acta Horti gothoburg., 9: 153-164. HACKEL, E. (1880). Catalogue raisonné des Graminées du Portugal, p. 24. Coimbra. LonarD, R. I. & GouLp, F. W. (1974). The North American species of Vulpia (Gramineae). Madroji, 22: 217-230. LousLEyY, J. E. (1961). A census list of wool aliens found in Britain, 1946-1960. Proc. bot. Soc. Br. Isl., 4: 221-247. Nees, C. G. (1829). Agrostologia brasiliensis, p. 474. Stuttgart & Tiibingen. TOWNSEND, C. C. (1959). Vulpia australis (Nees ex. Steud.) Blom. Proc. bot. Soc. Br. Isl., 3: 289-290. C. A. STACE & R. COTTON FESTUCA GLAUCA AUCT. AND FESTUCA CAESIA SM. In a recent paper (Trist 1973), I discussed the glaucous fescue confined to a limited area of the Suffolk Breckland. It had been considered that one species with two varieties could be identified, but the results of a field study showed that only one taxon should be recognized. As pointed out in that paper, the uncertainty of the identification of this glaucous Breckland fescue has been a contention of botanists for over 160 years. Smith (in Sowerby & Smith 1808), who studied this plant and collected it from the Breckland site, gave it specific rank as Festuca caesia Sm. and after much study concluded that it was quite distinct from Festuca ovina L. and Festuca duriuscula auct. (F. longifolia Thuill.). Hackel (1882) placed both of the varieties referred to by Trist (1973) under Festuca ovina sensu lato, as F. ovina subsp. eu-ovina var. glauca (Lam.) Hack. and F. ovina subsp. eu-ovina var. glauca subvar. caesia (Sm.) Hack. Richter (1890) similarly subordinated Smith’s F. caesia to F. glauca Lam. as F. glauca Lam. e. caesia (Sm.) K. Richter, which has been regarded as a subspecies by some (e.g. Auquier 1973) and as a variety by other authors (e.g. Hubbard 1954). In the circumstances, it seems reasonable to doubt that a comparison of the English and Continental glaucous fescues has been fully made; in fact it must be admitted that a full under- standing of the taxonomy of this group of glaucous fescues has still not been reached. Dr P. Augquier of Liége University (1973 in litt.) has told me that the date of Lamarck’s description of F. glauca was 1788 and nct 1786, as previously thought. This was one year later than the notes published on F. glauca by Villars (1787). Villars would have known of Lamarck’s work, but were the plants described by each of them the same or different? Auquier (1973 in litt.) also informed me that his studies have conclusively shown that F. glauca sensu Villars has distinct differences from the Breckland plants and also from the plants of the locus typicus of Lamarck in the Massif Central; Lamarck’s plants also differ from those found in the Suffolk Breckland. Auquier (1973) discussed the Suffolk plants and those from the loci typici of Villars and Lamarck and presented the results of a very detailed study. He has made a valuable contribution to our knowledge of this group of glaucous fescues. It now appears clear that the two fescues of Villars and Lamarck cannot both be referred to the same taxon; also it is shown that the Breckland fescue is different from either. It therefore follows that the name of the Breckland plant must be reconsidered. In considering the present state of our knowledge of the F. ovina group, Auquier (1973 in litt.) has suggested that the Breckland fescue be referred to the original name given by J. E. Smith, 1.e. F. caesia Sm. ACKNOWLEDGMENTS I am grateful to Dr P. Auquier for information through his publications and correspondence, which gave invaluable guidance, and also to Dr C. E. Hubbard for his continued interest and assistance in this complicated group of fescues. 74 SHORT NOTES REFERENCES AUQUIER, P. (1973). Qu’est-ce que le Festuca caesia Sm. (Poaceae)? Lejeunia, n.s., 70. HACKEL, E. (1882). Monographia festucarum europaearum. Kassel & Berlin. HUBBARD, C. E. (1954). Grasses, p. 111. Harmondsworth. LAMARCK, J. B. (1788). Encyclopédie méthodique, 2: 459. Paris & Liége. RICHTER, K. (1890). Plantae Europeae, 1: 95. Leipzig. SOWERBY, J. & SMITH, J. E. (1808). English botany, 27: Plate 1917. London. Trist, P. J. O. (1973). Festuca glauca Lam. and its var. caesia (Sm.) Richt. Watsonia, 9: 257-262. VILLARS, D. (1787). Histoire des plantes de Dauphiné, 2: 99, in obs. Lyon & Paris. P. J. O. TRist Watsonia, 11, 75-82 (1976). 75 Plant Records Records for publication must be submitted in the form shown below to the appropriate vice-county Recorder (Watsonia, 8: 435-447 (1971), and not to the Editors. Records are arranged in the order given in the List of British Vascular Plants by J. E. Dandy (1958) and his subsequent revision (Watsonia, 7: 157-178 (1969)) but Taraxacum is arranged according to A. J. Richards (Watsonia, 9, Suppl. (1972)). With the exception of collectors’ initials, herbarium abbreviations are those used in British Herbaria by D. H. Kent (1958). The following signs are used: *before the record: to indicate a new vice-county record. tbefore the species number: to indicate that the plant 1s not a native species of the British Isles. tbefore the record: to indicate a species which, though native in some parts of the British Isles, is not so in the locality recorded. [] enclosing a previously published record: to indicate that the record should be deleted. 1/5. LYCOPODIUM ALPINUM L. *84, Linlithgow: Broxburn, GR 36/08.72. Shale bing. I. R. Bonner, 1974, field record. 4/9 x 5. EQUISETUM ARVENSE L. x E. FLUVIATILE L. *35, Monmouth: Henllys, GR 31/26.92. Fen. T. G. Evans, 1974, herb. T.G.E., det. A. E. Wade. (Nature Wales, 14: 197 (1975)). 5/1. OSMUNDA REGALIS L. 109, Caithness: Dunnet Head, GR 39/2.7. E. R. Bullard, 1974, field record. Ist record since 1856. 21/3. DRYOPTERIS ABBREVIATA (DC.) Newm. *107, E. Sutherland: Carn Dearg, GR 29/37.39. A. G. Kenneth, 1973, BM. 21/6. DRYOPTERIS CARTHUSIANA (Vill.) H. P. Fuchs 78, Peebles: Broughton, GR 36/11.32. Cleared woodland. J. Dyce, 1972, field record. Ist localized post-1930 record. 21/a. DRYOPTERIS ASSIMILIS S. Walker 107, E. Sutherland: Creag na h-Uidhe, GR 29/38.37. A. G. Kenneth, 1973, BM. Ist record since 1891. 25/1/1. POLYPODIUM VULGARE L. *82, Haddington: Gifford, GR 36/54.68. *84, Linlith- gow: Linlithgow golf course, GR 26/98.76. Both records E. P. Beattie, 1972, E, det. J. A. Crabbe & A. C. Jermy. 25/1/2. POLYPODIUM AUSTRALE Fée *1, W. Cornwall: Padstow, GR _ 10/91.75. Stone embankment. M. Rickard, 1972, field record. 25/1/3 x 1. POLYPODIUM INTERJECTUM Shivas x P. VULGARE L. *82, Haddington: Traprain Law, near East Linton, GR 36/57.74. E. P. Beattie, 1972, E, det. J. A. Crabbe & A. C. Jermy. ¢27/1. AZOLLA FILICULOIDES Lam. *1, W. Cornwall: Ding Dong, near Penzance, GR 10/44.34. Pool in old quarry. B. Sturdy, 1971, field record. 29/1b. OPHIOGLOSSUM VULGATUM L. subsp. AMBIGUUM (Coss. & Germ.) E. F. Warb. *70, Cumberland: Ravenglass Nature Reserve, GR 34/06.96. Dune-slack. T. Dargie, 1972, BM, det. A. C. Jermy. 46/12b. RANUNCULUS FLAMMULA L. subsp. scoticus (E. S. Marshall) Clapham *96b, Nairn: Loch Loy, GR 28/92.58. M. McC. Webster, 1974, field record. 46/17. RANUNCULUS OMIOPHYLLUS Ten. *H14, Laois: Gales Hill, Arless, IGR $62.82. F. H. Perring & D. A. Webb, 1974, field record. 76 PLANT RECORDS +58/6. PAPAVER SOMNIFERUM L. *78, Peebles: Horsbrugh quarry, GR 36/28.39. B. Morrison, 1973, field record. Ist localized record. 66/4. FUMARIA BASTARDII Bor. *66, Durham: Piercebridge, GR 45/20.15. Excavations. M. Burnip, 1974, herb. G. G. Graham. Ist definite record. +71/1. HtRSCHFELDIA INCANA (L.) Lagr.-Foss. *35, Monmouth: Newport Docks, GR 31/30.85. T. G. Evans, 1973, herb. T.G.E. Long established and widespread. 498/3. BARBAREA INTERMEDIA Bor. *68, Cheviot: near Shipleylane, N.W. of Alnwick, GR 46/14.18. Roadside. R. S. G. Thompson, 1969, 1971 specimen herb. G. A. Swan. 102/2 x 1. RoRIPPA MICROPHYLLA (Boenn.) Hyland. x R. NASTURTIUM-AQUATICUM (L.) Hayek *84, Linlithgow: Union Canal, Winchburgh, GR 36/08.75. I. R. Bonner, 1974, field record. 4105/1. ERYSIMUM CHEIRANTHOIDES L. *107, E. Sutherland: Rovie Lodge, Strath Fleet, GR 29/71.03. Garden weed. L. S. Garrad, 1972, field record, det. D. McClintock. 115/1. HypeERICUM ANDROSAEMUM L. *107, E. Sutherland: Badbeithe, Loch Migdale, GR 28/63.9191. O. J. Kirby, 1975, field record. 133/6. STELLARIA PALUSTRIS Retz. *78, Peebles: Putt’s Pool, Innerleithen, GR 36/34.37. Oxbow of R. Tweed. D. J. McCosh, 1974, herb. D.J.M. 136/3. SAGINA MARITIMA Don *61, S.E. Yorks.: Hull docks, GR 54/14.28. E. Chicken, 1965, herb. E.C., det. G. A. Matthews. 160/4. SALICORNIA RAMOSISSIMA Woods *111, Orkney: Otterswick, Sanday, GR 57/69.42. E. R. Bullard, 1974, LTR, det. T. G. Tutin. 168/7. GERANIUM SANGUINEUM L. *12, N. Hants.: The Lunway, near South Wonston, GR 41/47.36. Lane near old railway. A Brewis, 1972, field record. Probably introduced. 168/15. GERANIUM LUCIDUM L. *78, Peebles: Rachan Home Farm, Broughton, GR 36/12.34. Roadside wall. R. Hunter, 1971, field record. Probably introduced. 187/2. ULEX GALLU Planch. *109, Caithness: Hill of Mid Clyth, GR 39/28.37. E. R. Bullard, 1974, E, conf. M. C. F. Proctor. +207/g. LATHYRUS GRANDIFLORUS Sibth. & Sm. *82, Edinburgh: near North Elphinstone, GR 36/39.71. Along hedge. N. F. & O. M. Stewart, 1974, field record. #211/11/139. RuBus pRocERUS P. J. Muell. *82, Haddington: S. of Luffness Convent, Aberlady, GR 36/47.80 *83, Edinburgh: Cramond Island, GR 36/1.7. *84 Linlithgow: Carmelhill, GR 36/10.75. All records E. P. Beattie, 1966, E, det. B. Miles. 211/11/178. RuBUS CRINIGER (E. F. Linton) Rogers *66, Durham: Eastgate, GR 35/95.38. Railway embankment. G. G. Graham, 1974, DHM, det. A. Newton. 211/11/212. RuBUS ECHINATUS Lindl. *1, W. Cornwall: Crookhorn, St Just-in-Penwith, GR 10/38.30. Moorland. E. S. Edees, 1974, field record. 212/14 x 15. POTENTILLA ANGLICA Laichard x P. REPTANS L. *66, Durham: Monkton Burn, Jarrow, GR 45/32.63. G. G. Graham & J. W. Patrick, 1974, herb. G.G.G., det. B. Matfield 1st definite record. 220/2. ALCHEMILLA CONJUNCTA Bab. +*84, Linlithgow: Philpstoun, GR 36/05.77. Shale bing. I. R. Bonner, 1974, herb. I.R.B. 220/3/1. ALCHEMILLA GLAUCESCENS Wallr. +*83, Edinburgh: Western Dock, Leith, GR 36/25.77. O. M. Stewart, 1973, E, det. M. E. Bradshaw. PLANT RECORDS 77 220/3/9. ALCHEMILLA GLOMERULANS Buser *107, E. Sutherland: Creag na h-Uidhe, GR 29/38.37. Rock ledges. A. G. Kenneth, 1973, herb. A.G.K., det. S. M. Walters. $224/1. ACAENA ANSERINIFOLIA (J. R. & G. Forst.) Druce *78, Peebles: Talla reservoir, GR 36/10.22. R. C. L. Howitt, 1972, field record. $225/3. ROSA MULTIFLORA Thunb. *83, Edinburgh: Ratho, GR 36/13.70. By canal. E. P. Beattie, 1968, E, det. D. McClintock. 225/4. ROSA PIMPINELLIFOLIA L. 78, Peebles: Gameshope burn, GR 36/13.19. On rocks. D. J. McCosh, 1972, herb. D.J.M. Ist record since 1897. 7227/4. COTONEASTER MICROPHYLLUS Wall. ex Lindl. *78, Peebles: Dawyck, GR 36/16.34. Scree in woodland. D. J. McCosh, 1972, herb. D.J.M., det. N. K. B. Robson. Well-naturalized. 229/1. CRATAEGUS LAEVIGATA (Poir.) DC. *68, Cheviot: near Rugley Wood, Alnwick, GR 46/17.11. By disused railway. R. S. G. Thompson, 1971, 1973 specimen herb. G. A. Swan, det. A. D. Bradshaw. Ist definite record. 237/1. CRASSULA TILLAEA L.-Garland *96, Easterness: by Caledonian Canal, Torvaine, GR 28/65.43. M. Barron, 1974, herb. E. R. T. Conacher, det. H. J. M. Bowen. 239/8. SAXIFRAGA TRIDACTYLITES L. 68, Cheviot: near Ratcheugh, Alnwick, GR 46/22.14. Roadside wall. R. S. G. Thompson, 1965, field record. Ist record for 100 years. 254/5. EpILOBIUM ROSEUM Schreb. *78, Peebles: Broughton station, GR 36/11.36. Old railway line. R. Hunter, 1970, herb. D. J. McCosh, det. T. D. Pennington. 4254/6 x 2. EPILOBIUM ADENOCAULON Hausskn. x E. PARVIFLORUM Schreb. *67, S. Northumberland: Brunton Quarry, N. of Hexham, GR 35/92.70. G. A. & M. Swan, 1971, herb. G.A.S., det. T. D. Pennington. 259/2. MYRIOPHYLLUM SPICATUM L. *109, Caithness: Loch Watten, GR 39/2.5. E. R. Bullard, 1974, field record. 1st localized record. 261/1. HrpPURIS VULGARIS L. 78, Peebles: Putt’s Pool, Innerleithen, GR 36/34.37. Oxbow of R. Tweed. D. J. McCosh, 1974, herb. D.J.M. Ist post-1930 record. 294/2. PIMPINELLA MAJOR (L.) Huds. *68, Cheviot: Christon Bank, N. of Alnwick, GR 46/21.22. Railway bank. R. S. G. Thompson, 1962, 1971 specimen herb. G. A. Swan. 309/3. PEUCEDANUM OSTRUTHIUM (L.) Koch 68, Cheviot: Howick Pond, N.E. of Alnwick, GR 46/25.17. R. S. G. Thompson, 1972, herb. G. A. Swan. Ist record for over 50 years. +319/14. EUPHORBIA URALENSIS Fisch. ex Link *33, E. Gloucs.: between Upper Slaughter and Lower Swell, GR 32/16.24. Roadside. M. J. Cook, 1974, field record. +320/20. POLYGONUM SACHALINENSE F. Schmidt *68, Cheviot: near Bilton, W. of Alnmouth, GR 46/22.11. R. S. G. Thompson, 1966, 1972 specimen herb. G. A. Swan. 343/5. SALIX TRIANDRA L. *78, Peebles: Hallyne, GR 36/19.40. R. C. L. Howitt, 1972, herb. D. J. McCosh. Possibly introduced. 343/14. SALIX NIGRICANS Sm. *78, Peebles: 3 miles N. of West Linton, GR 36/12.54. R. C. L. Howitt, 1973, herb. D. J. McCosh. Ist localized record. 343/17. SALIX LAPPONUM L. *107, E. Sutherland: Creag Mhor, GR 29/7.2. D. A. Ratcliffe, 1962, field record. 354/1. ARCTOSTAPHYLOS UVA-URSI (L.) Spreng. 78, Peebles: Polmood burn, 6 miles S. of Broughton, GR 36/13.26. Calluna hillside. C. O. Badenoch, 1973, field record. Ist record since LIT 78 PLANT RECORDS 4395/1. PENTAGLOTTIS SEMPERVIRENS (L.) Tausch *78, Peebles: Traquair, GR 36/3.3. C. Morrison, 1972, field record. 406/4. CALYSTEGIA SOLDANELLA (L.) R.Br. *67, S. Northumberland: mouth of Meggie’s Burn, Blyth, GR 45/32.78. Sand-dunes. J. D. Parrack, 1973, herb. G. A. Swan. 430/20b. VERONICA HEDERIFOLIA L. subsp. LUCORUM (Klett & Richt.) Hartl *33, E. Gloucs.: Naunton, GR 42/11.23. Garden weed. K. E. Higgs, 1973, field record, det. P. M. Benoit. 437/1. PARENTUCELLIA VISCOSA (L.) Caruel *68, Cheviot: Denwick Quarry, near Alnwick, GR 46/21.14. R. S. G. Thompson, 1971, herb. G. A. Swan. 440/8. OROBANCHE MINOR Sm. *H39, Antrim: The Curran, Larne, GR 34/41.02. Waste ground. D. S. Ledsham, 1973, BEL. 493/1. ADOXA MOSCHATELLINA L. 1, W. Cornwall: Carnaton Woods, St Mawgan in Pydar, GR 10/87.65. River bank. L. Grigg, 1975, field record. Ist post-1930 record. 494/5. VALERIANELLA DENTATA (L.) Poll. 66, Durham: High Goosepool, Middleton-St- George, GR 45/36.14. Farm-track. M. Burnip, 1974, herb. G. G. Graham, conf. C. C. Townsend. Ist record this century. 7506/4 x 8. SENECIO SQUALIDUS L. x S. VULGARIS L. *83, Edinburgh: Granton, GR 36/22.77. O. M. Stewart, 1974, E, det. J. E. Lousley. 4506/13. SENECIO FLUVIATILIS Wallr. *109, Caithness: between Ulbster & Thrumster, GR 39/32.42. Roadside. E. R. Bullard, 1974, E, conf. I. Hedge. 4509/3. PETASITES JAPONICUS (Sieb. & Zucc.) F. Schmidt *34, W. Gloucs.: Lasborough, GR 31/81.93. Streambank. B. M. Mack, 1974, field record. *67, S. Northumberland: by R. Wansbeck, near Bothal, GR 45/23.86. D. M. McArthur, 1974, herb. G. A. Swan. 555/1. MYCELIS MURALIS (L.) Dumort. *96b, Nairn: Holme Rose, GR 28/81.48. M. McC. Webster, 1974, E. §58/1/28. HiERACIUM CENTRIPETALE F. J. Hanb. *78, Peebles: Cape Law, Talla, GR 36/14.14. D. J. McCosh, 1974, herb. D.J.M., det. P. D. Sell. 558/1/47. Hi&RACIUM SCHMIDTII Tausch *78, Peebles: by R. Lyne, West Linton, GR 36/14.52. D. J. McCosh, 1974, herb. D.J.M., det. P. D. Sell. 558/1/53. HIERACIUM ARGENTEUM Fries *78, Peebles: by Gameshope burn, Talla, GR 36/13.19. D. J. McCosh, 1972, herb. D.J.M., det. P. D. Sell. 558/1/66. HiERACIUM SAXORUM (F. J. Hanb.) Sell & West *78, Peebles: by Gameshope burn, Talla, GR 36/13.19. D. J. McCosh, 1974, herb. D.J.M., det. P. D. Sell. 558/1/94. H1ERACIUM DURICEPS F. J. Hanb. *78, Peebles: N. of Megget Water, GR 36/19.23. D. J. McCosh, 1974, herb. D.J.M., det. P. D. Sell. 558/1/99. HIERACIUM GRANDIDENS Dahlst. *78, Peebles: Polmood, GR 36/11.27. D. J. McCosh, 1972, herb. D.J.M., det. P. D. Sell. 558/1/138. HIERACIUM CRAVONIENSE (F. J. Hanb.) Roffey *78, Peebles: Tweedsmuir, GR 36/09.24. D. J. McCosh, 1972, herb. D.J.M., det. P. D. Sell. 558/1/158. HtiERACIUM DIAPHANUM Fr. *61, S.E. Yorks.: Sledmere, GR 44/94.67. E. Chicken, 1974, herb. E.C., det. C. E. A. Andrews. 558/1/163. HIERACIUM STRUMOSUM (W. R. Linton) A. Ley *78, Peebles: Macbiehill, GR 36/18.51. D. J. McCosh, 1974, herb. D.J.M., det. P. D. Sell. §58/1/168. HtTERACIUM DOVRENSE Fr. *107, E. Sutherland: Creag na h-Uidhe, GR 29/38.37. A. G. Kenneth & A. McG. Stirling, 1973, herb. C. West, det. C.W. PLANT RECORDS 79 558/1/180. HtERACIUM PRENANTHOIDES Vill. *78, Peebles: by Gameshope burn, Talla, GR — 36/13.19. D. J. McCosh, 1972, herb. D.J.M., det. P. D. Sell. 558/1/196. HiERACIUM SPARSIFOLIUM Lindeb. *78, Peebles: Tweedsmuir, GR 36/09.24. D. J. McCosh, 1974, herb. D.J.M., det. P. D. Sell. 558/1/210. HiERACIUM STRICTIFORME (Zahn) Roffey *78, Peebles: by Gameshope burn, Talla, GR 36/13.19. D. J. McCosh, 1972, herb. D.J.M., det. P. D. Sell. 558/2/3a. PILOSELLA FLAGELLARIS (Willd.) Sell & West subsp. FLAGELLARIS *78, Peebles: Manorfoot, Peebles, GR 36/22.39. D. J. McCosh, 1971, herb. D.J.M., det. P. D. Sell. +558/2/6b. PILOSELLA AURANTIACA (L.) C. H. & F. W. Schultz subsp. BRUNNEOCROCEA (Pugsl.) Sell & West *78, Peebles: Tweedsmuir, GR 36/09.24. D. J. McCosh, 1972, herb. D.J.M., det. P. D. Sell. Ist definite record. 559/8. CREPIS PALUDOSA (L.) Moench *H14, Laois: + mile W. of Maganey, IGR $71.84. Riverside scrub. F. H. Perring & D. A. Webb, 1974, field record. 560/3. TARAXACUM LACISTOPHYLLUM (Dahlst.) Raunk. *99, Dunbarton. 560/6. TARAXACUM SILESIACUM Dahlst. ex Hagl. *1, W. Cornwall. 560/16. TARAXACUM FULVIFORME Dahlst. *99, Dunbarton. 560/17. TARAXACUM OXONIENSE Dahlst. *85, Fife. 560/29. TARAXACUM PLATYGLOSSUM Raunk. *85, Fife. 560/36. TARAXACUM FAEROENSE (Dahlst.) Dahlst. *H39, Antrim. 560/37. TARAXACUM SPECTABILE Dahlst. *99, Dunbarton. 560/42. TARAXACUM EURYPHYLLUM (Dahlst.) M.P.Chr. *1, W. Cornwall *99 Dunbarton. 560/44. TARAXACUM PRAESTANS H.Lindb,f. *1, W. Cornwall. 560/48. TARAXACUM DRUCEI Dahlst. *107, E. Sutherland. 560/50. TARAXACUM NAEVOSUM Dahlst. *95, Moray. 560/51. TARAXACUM LAETIFRONS Dahlst. *61, S.E. Yorks. 560/58. TARAXACUM STRICTOPHYLLUM Dahlst. *96, Easterness. 560/61. TARAXACUM NORDSTEDTII Dahlst. *85, Fife *94 Banff. 560/67. TARAXACUM SUBCYANOLEPIS M.P.Chr. *1, W. Cornwall *85, Fife *99, Dun- barton. 560/69. TARAXACUM SELLANDII Dahlst. *56, Noits. " 560/70. TARAXACUM ANCISTROLOBUM Dahlst. *1, W. Cornwall *96b, Nairn *99, Dun- arton. 560/71. TARAXACUM SUBLACINIOSUM Dahlst. & H.Lindb.f. *1, W. Cornwall *95, Moray. 7560/74. TARAXACUM PROCERUM Hag]. *1, W. Cornwall *56, Notts. $60/75. TARAXACUM PANNUCIUM Dahist. *1, W. Cornwall *106, E. Ross. 7560/76. TARAXACUM LINGUATUM Dahlst. ex M.P.Chr. & Wiinst. *85, Fife *95, Moray. 560/78. TARAXACUM ALATUM H.Lindb,f. *1, W. Cornwall *85, Fife. 560/79. TARAXACUM LINGULATUM Markl. *1, W. Cornwall *56, Notts. 80 PLANT RECORDS +560/80. TARAXACUM AEQUISECTUM M.P.Chr. *99, Dunbarton. 560/81. TARAXACUM CROCEIFLORUM Dahlst. *1, W. Cornwall *85, Fife *99, Dun- barton. 560/83. TARAXACUM EXPALLIDIFORME Dahlst. *1, W. Cornwall. 560/84. TARAXACUM eRe Ekman ex Raunk. *1, W. Cornwall. 560/94. TARAXACUM EKMANII Dahlst. *94, Banff *95, Moray. 560/95. TARAXACUM PORRECTIDENS Dahlst. *H39, Antrim. +560/97. TARAXACUM COPIDOPHYLLUM Dahlst. *61, S.E. Yorks. +560/98. TARAXACUM PECTINATIFORME H.Lindb.f. *95, Moray. 560/99. TARAXACUM AUROSULUM H.Lindb.f. *56, Notts. *61, S.E. Yorks. 560/100. TARAXACUM XANTHOSTIGMA H.Lindb.f. *56, Notts. *H39, Antrim. 560/103. TARAXACUM CORDATUM Palmer. *1, W. Cornwall *85, Fife *99, Dunbarton. 560/104. TARAXACUM ADSIMILE Dahlst. *85, Fife. 560/105. TARAXACUM LONGISQUAMEUM H.Lindb.f. *H39, Antrim. 560/106. TARAXACUM DAHLSTEDTH H.Lindb.f. *1, W. Cornwall *83, Edinburgh *85, Fife. 560/107. TARAXACUM DUPLIDENS H.Lindb.f. *1, W. Cornwall *99, Dunbarton. 560/113. TARAXACUM BRACTEATUM Dahlst. *99, Dunbarton *H39, Antrim. 560/114. TARAXACUM HAMATUM Raunk. *61, S.E. Yorks *85, Kinross. 560/115. TARAXACUM HAMATIFORME Dahlst. *94, Banff *H39, Antrim. 560/116. TARAXACUM MARKLUNDII Palmer. *1, W. Cornwall. 560/117. TARAXACUM LATISECTUM H.Lindb.f. *H39, Antrim. 560/119. TARAXACUM MACULATUM Jordan *1, W. Cornwall *§9, Dunbarton *H39, Antrim. 560/121. TARAXACUM RAUNKIAERII Wiinst. *85, Kinross *94, Banff *99, Dunbarton. 560/122. TARAXACUM HEMIPOLYODON Dahist. *95, Moray. 560/123. TARAXACUM OBSCURATUM Dahist. *56, Notts. 560/125. TARAXACUM POLYODON Dahlst. *1, W. Cornwall *92, S. Aberdeen. 4605/2. JUNCUS TENUIS Willd. *61, S.E. Yorks.: Hull docks, GR 54/10.28. E. Chicken, 1969, herb. E.C. Site now built over. 625/4. EpIPACTIS LEPTOCHILA (Godfery) Godfery *67, S. Northumberland: by R. South Tyne near Williamston, GR 35/6.5. B. M. Cunliffe & M. Rawes, 1973, field record, 1974 specimen herb. G. A. Swan. (Watsonia, 11:1 (1976)). 625/6. EpIpACTIS PHYLLANTHES G.E.Sm. *67, S. Northumberland: Settlingstones, GR 35/8.6. Lead mine. G. A. & M. Swan, 1974, herb. G.A.S. (Watsonia, 11:4 (1976)). PLANT RECORDS 81 628/1. LisTERA OVATA (L.) R.Br. 78, Peebles: Wakefield, West Linton, GR 36/13.53. D. J. McCosh, 1974, field record. 1st post-1930 record. 643/2 x 636/1. DACTYLORHIZA MACULATA (L.) Sod x GYMNADENIA CONOPSEA (L.) R.Br. aa W. Cornwall: Crousa Downs, The Lizard, GR 10/76.19. Wet heath. B. Garratt, 1972, field record, det. V. Summerhayes. +646/1. ACORUS CALAMUS L. *68, Cheviot: by R. Tweed, # mile S. of Norham, GR 36/89.46. G. A. & M. Swan, 1974, field record. 650/1. LEMNA POLYRHIZA L. 1*67, S. Northumberland: Saltwick plantation, near Morpeth, GR 45/17.82. B. & I. D. Wallace, 1967, 1974 specimen herb. G. A. Swan. 652/3. SPARGANIUM ANGUSTIFOLIUM Michx. *66, Durham: Bollihope Moor, GR 35/98.34. M. Burnip & G. G. Graham, 1974, DHM, conf. BM. 656/6. ELEOCHARIS UNIGLUMIS (Link) Schult. *45, Pembroke: Landshipping Ferry, GR 22/01.11. Upper salt-marsh. T. A. W. Davis, 1974, NMW. Ist localized record. (Nature Wales, 14: 202 (1975)). 663/62. CAREX DIvISA Huds. *1, W. Cornwall: Harlyn Bay, near Padstow, GR 10/87.75. Riverside field. L. J. Margetts, 1972, field record, det. R. W. David. 663/66. CAREX MURICATA L. subsp. LEERSI Aschers. & Graebn. *61, S.E. Yorks: Lily Dale, Millington, GR 44/83.53. Woodland ride. E. Chicken, 1965 herb. F. E. Crackles, det. R. W. David. 663/71. CAREX REMOTA L. *78, Peebles: Dawyck, GR 36/16.34. D. J. McCosh, 1972, herb. D.J.M. 663/72. CAREX CURTA Gooden. 1, W. Cornwall: Budock Bottoms, GR 10/77.31. L. J. Margetts, 1962, field record. Ist post-1930 record. 669/2. GLYCERIA PLICATA Fr. 78, Peebles: Candyburn, Skirling, GR 36/06.40. D. J. McCosh, 1974, herb. D.J.M. Ist post-1930 record. 669/3. GLYCERIA DECLINATA Bréb. *84, Linlithgow: Cockleroy, GR 26/98.74. I. R. Bonner, 1974, field record. 670/1 x 7671/2. FESTUCA PRATENSIS Huds. x LOLIUM MULTIFLORUM Lam. *66, Durham: Newsham, GR 45/38.11. Field-margin. M. Burnip, 1974, herb. G. G. Graham, det. T. G. Tutin. 4671/2 x 1. LOLIUM MULTIFLORUM Lam. x L. PERENNE L. *83, Edinburgh: Blackford quarry, GR 36/2.7. O. M. Stewart, 1974, E, det. C. E. Hubbard. +676/14. PoA PALUSTRIS L. 33, E. Gloucs.: Old Severn, Upper Lode, Tewkesbury, GR 32/88.33. S. C. Holland & Lady Ricketts, 1974, field record, det. C. E. Hubbard. Ist post-1930 record. 677/1. CATABROSA AQUATICA (L.) Beauv. 67, S. Northumberland: by Bothal Burn, near Pegswood, GR 45/23.87. C. A. & M. Swan, 1974, field record. Ist record for over 100 years. 683/13 x 10. BRomMuUS LEPIDUS Holmberg x B. MOLLIS L. *33, E. Gloucs.: Ashchurch army depot, near Tewkesbury, GR 32/93.34. C. W. Bannister, 1974, field record, det. C. E. Hubbard. 685/5 x 3. AGROPYRON JUNCEIFORME (A. & D. Love) A. & D. L6ve x A. REPENS (L.) Beauv. *109, Caithness: Stain, Keiss, GR 39/34.60. C.S.S.F. field meeting, 1972, herb. A. Copping. 701/4. AGROSTIS GIGANTEA Roth *109, Caithness: Thurso railway station, GR 39/11.67. C.S.S.F. field meeting, 1972, field record. F 82 PLANT RECORDS +701/8. AGROSTIS SEMIVERTICILLATA (Forsk.) C.Chr. 1, W. Cornwall: Falmouth docks, GR 10/81.32. L. J. Margetts, 1972, field record, det. C. E. Hubbard. Ist post-1930 record. 705/1. GASTRIDIUM VENTRICOSUM (Gouan) Schinz & Thell. 1, W. Cornwail: Trelissick, Fal estuary, GR 10/83.39. Old wall. B. Garratt, 1973, field recoid. Ist post-1930 record. . 707/1. PHLEUM BERTOLONIT DC. *78, Peebles: Whitehaugh, GR 36/26.39. C. Morrison, 1972, field record. 708/3 x 2. ALOPECURUS GENICULATUS L. x A. PRATENSIS L. #33, E. Gloucs.: Lower Lode, Tewkesbury, GR 32/88.31. S. C. Holland & Lady Ricketts, 1974, field record, det. C. E. Hubbard. Watsonia, 11, 83-88 (1976). 83 Book Reviews The Wild Flowers of Guernsey. David McClintock. Pp. 288. Collins, London. 1975. Price £4:-75. Guernsey is a small, geologically uniform island of 63 square kilometres, smaller than a grid square in the Azlas. Its soils are fertile, and its forests were completely cleared by Norman times, while few of its marshes remain undrained; it has no nature reserves. It has been densely populated for at least a thousand years, and currently supports 51,500 people, so that its population density is nearly three times that of Belgium or Holland. The local people are skilled and efficient horticulturists, who have radically changed their pattern of land use in the last hundred and fifty years, so that about ten per cent of the island is now under glass. Given these facts, we might expect the flora to be a poor one, with few surviving native species. In fact, David McClintock has been able to list 1,340 plant species which have been found on the island, compared with 684 listed by Babington in 1839 and 957 by Marquand in 1901. The new Flora combines immense botanical and historical erudition without ever becoming too technical: it is designed to be read as well as consulted. It is a very critical work, both in rejecting the unusually large number of spurious records and in the care with which correct names have been cited (though purists may disapprove of the retention of the initial capital for some specific epithets). It gives much information on local variations, so important in island floras, for example in Spergula arvensis, Oxalis latifolia and Salvia verbenaca (now lumped with S. marquandii). It includes all aliens, a policy which I approve, since this is a region where naturalization takes place rapidly. It gives excellent brief notes on status, frequency, history, horticultural merit, occurrence on other Channel Islands, local uses and names: English, Latin and local names are ali indexed together. Inevitably this Flora will be compared with Lousley’s Flora of the Isles of Scilly (1971), and the comparison reveals several omissions in the Guernsey work. The lack of almost all background information on geology, soils, physical features and climate is compensated by listing references to these topics in an appendix. More important, no attempt is made to interpret the remarkable facts presented concerning the occurrences and absences of native and introduced plants. No mention is made of the fact that the more interesting natives fall into several different categories with regard to their European distribution—Matthews’ ‘Mediterranean’, ‘Oceanic’ and ‘Conti- nental Southern’ groupings. Comparative tables of the numbers of naturalized species from different parts of the world might have analytic and predictive value. A number of the introduced species recorded in this work are not mentioned in Flora Europaea. This Flora gives no keys, e.g. for the following genera, which contain taxa on Guernsey which are not covered by standard British Floras: Agropyron, Asplenium, Calystegia, Centaurea, Mentha, Narcissus, Olearia, Pilosella, Rosa, Solanum and the hardy bamboos. Simple keys, or more hints on identification of the un- familiar species, would greatly increase the value of the work to visiting botanists. The thumbnail sketches of rarities in the text only give an idea of the plants, but their ‘jizz’, to use David McClintock’s own word, is more easily learnt from clear descriptions or magnified drawings of critical parts. The book is stoutly bound, the paper is of rather poor quality, and the number of misprints is small (e.g. amygdaloides mis-spelt on page 143). One small printing defect concerns the spacing of authors’ names and Guernsey names after the Latin name of each species. The uninitiated may be led into thinking that the islanders refer to Rosy Bindweed as Brummitt and Wavy Bitter-cress as Withering. Despite these minor defects, this book is a worthy addition to the long list of British local Floras. It is essential reading for all visiting botanists, and deserves a wide sale among Islanders and holiday visitors. H. J. M. BOWEN Kormofyternes Taxonomi. K. Larsen, with illustrations by B. Johnsen. Pp. 223 including 108 black and white plates of line drawings. Akademisk Forlag, K@benhavn. 1973. Price not stated. 84 BOOK REVIEWS This modest, but valuable, paper-back publication provides a survey of the Cormophyta, and is primarily intended for students. This term, which is not very familiar to British botanists, embraces the bryophytes, pteridophytes and flowering-plants. Concise family descriptions are provided, in a sequence following Takhtajan’s system, along with brief notes on economic plants and other points of interest. Numerous excellent plates by Bent Johnsen illustrate the diagnostic features of flowers and fruit, through habit drawings, floral details and floral diagrams. In common with the reviewer, most British botanists will lack the knowledge of the Danish language which would permit them to make free use of the text. However, the first-class illustrations speak clearly and in a thoroughly international tongue, and these in themselves are sufficient to commend the work to British readers. In a concise work for students, some restriction of coverage is both appropriate and essential. Naturally in this case, the selection favours families well represented in N.W. Europe, supplemented by some others, especially those in which major economic species are included. Nevertheless, the choice sometimes seems rather capricious, as when the Apocynaceae and Asclepiadaceae are allowed two lines between them, and the Verbenaceae fares little better. In contrast, the Bromeliaceae, Myrtaceae and Commelinaceae are allotted 18, 13 and 10 lines respectively. The reviewer understands that an English edition is under consideration and believes that there is a real need for a simple, concise review of the higher plants, a role that this book is well fitted to fill for amateur botanists as well as students. The outstandingly good illustrations are both highly informative and aesthetically attractive, and merit an English text, which would make them available to a much wider circle of users. J.F.M. CANNON Bibliographia Botanica Cechoslovaca 1952-1957. V. Skalicky & V. Holubova-Jechova. Pp. 649 in 2 vols. (pp. 1-308 in vol. 1; pp. 309-649 in vol. 2). Botanicky ustav CSAV, Pruhonice u Prahy. 1973. Price not stated. This long-delayed publication is a 5-year résumé, in compact, cyclostyled format, of the botanical literature of Czechoslovakia. All fields of botany are covered, and the arrangement of the 8,318 entries (which include the full titles of all the papers cited) is like that in the similar publication for 1965-1966 reviewed in Watsonia, 9: 161 (1972). Many additional subjects are cited by means of cross-references at the end, and there are very useful indexes to geographical regions, plant names, and authors of articles. Libraries will find this a valuable reference-work, and more con- venient than the annual bibliographies already available. A. O. CHATER How to Know Western Australian Wildflowers. A Key to the Flora of the Temperate Regions of Western Australia. Parts I-III. W. E. Blackall & B. J. Grieve. Pp. cxxiii+ 595, with 2 maps and numerous text illustrations. University of Western Australia Press. U.K. Distributors: Inter- national Book Distributors Ltd, Hemel Hempstead. 1975. Price £6-60. Previously issued between 1954 and 1965 in three separate parts, this unrevised reprint in a single volume is the outcome of ‘academic and public demand’. It is not a full Flora with comprehensive species descriptions but has the form of hand-written illustrated keys reproduced from printings of the original manuscripts. In addition there are explanatory notes on ‘How to use this book’ (for those without formal botanical training), a glossary, and separate indices to the three con- stituent parts. Despite its limitations and present incompleteness—twelve families (Solanaceae to Compositae) are still to come, in a later, promised volume—it is easy to see why there is the demand in (West) Australia for the publication of these keys in one handy volume. Australia, with many thousands of plant species, still has no comprehensive Flora other than Bentham’s Flora Australiensis {published between 1863 and 1878) and but few modern regional Floras. This volume must therefore be a welcome addition. The European botanist, not directly concerned with the Australian flora, will probably be more interested in the methods and techniques of presentation used. There is a general key to BOOK REVIEWS 85 families, which also enables one to refer a plant to the twelve families of the future volume. The dichotomous keys to genera and species form the body of the book; they incorporate numerous drawings as an integral part of the keys. The precision given by the ‘visual description’ and the elimination of all non-essential information produces a clarity and apparent simplicity of procedure. The facsimile handwriting is quite acceptable where entries are not crowded on the page; but many drawings are far from successful, especially in Part I where they are much reduced in size. There are many improvements in Parts II and III. The indices contain additional data: season of flowering and synonymy (Part III), distribution (Parts II and III), authorities for the Latin names (which are used throughout) (Parts II and III), but not in that for Part I, which fills over half the book. There are oddities in the sequence of families, by and large Englerian: Myrtaceae has been added to Part I and Goodeniaceae to Part II. There are some discrepancies in the treatment of the three parts which may be misleading: the change of scale of the drawings is signalled in the explanatory notes. Not being familiar with Western Australian species I cannot properly judge the efficacy of the keys; but, using specimens from New South Wales, I found the Key to families reasonably easy to use and had moderate success with appropriate genera and species. But is it really necessary to observe ‘fruit without vittae, compressed laterally’ (the drawing of which is unrecognizeable) before Conium can be keyed out? Of course there are snags; without corroborative descriptions a faulty identification may go unrecognized. But this method, pioneered (?) by Bonnier many years ago, has a lot to commend it, especially when it results in a single light-weight serviceable volume covering, in this case, 90% of the families represented in temperate Western Australia. It should be considered by anyone contemplating the writing of a working handbook to a comparably extensive flora. This is surely an invaluable handbook; it is excellently produced, sturdily bound in flexible covers and printed on glossy hard-wearing paper. And it costs only £6-60. A. P. CONOLLY Author and classified catalogues of the Royal Botanical Gardens, Kew. George Prior Associated Publishers Ltd, London. 1974. Author catalogue, 5 Vols. £175. Classified catalogue, 4 Vols. £142. Prior to May 1840, when the control of the Royal gardens at Kew passed from the Crown to the Commissioners of Woods and Forests, and Sir William J. Hooker was appointed as its first Director, there was no library or herbarium at Kew. The dried plant collections sent home by Kew collectors such as Caley, Good and Masson as well as many of their manuscripts were at the time in the possession of Sir Joseph Banks, and are now in the Department of Botany, British Museum (Natural History). No scientific establishment, however, can function efficiently without a comprehensive library, and soon after Sir William Hooker became Director he set about the formation of one at Kew. The founder collections were the libraries of Dr W. A. Bromfield (presented in 1852), George Bentham (presented in 1854) and Sir William Hooker (purchased in 1867). Today the library of the Royal Botanic Gardens, Kew, is one of the finest botanical libraries in the world, covering taxonomy, economic botany, plant geography, cytology, anatomy, physiology, and biochemistry. Adequate staffing in recent years has enabled them to maintain an excellent author catalogue and also to classify their entire stock and so provide a subject catalogue. The classification schedules used are not those of one particular well-known scheme, but a mixture of Bentham & Hooker for flowering plants, Kew’s own geographical schedule for Floras and an adaptation of the Dewey Decimal Classification for the remainder. It is important to note that the author catalogue includes pamphlets and a major part of the library’s very extensive collection of reprints. Important articles in obscure journals can often be found in a reprint collection and Kew are to be congratulated on having catalogued their vast collection and so being able to include them in their author catalogue. These volumes are a major contribution to botanical and horticultural bibliography. Printing directly from card catalogues is far from ideal but, alas, gone are the days of type-set-printed catalogues. P. I. EDWARDS 86 BOOK REVIEWS Flora of Iraq, Volume 3. C. C. Townsend & Evan Guest (editors). Pp. vii+ 662, with 111 plates, 1 coloured frontispiece and 6 distribution maps. Ministry of Agriculture, Republic of Iraq, Baghdad. 1974. Price £3-75. Thd third volume of this Flora (see Watsonia, 6: 390-1 (1968) for a review of Volumes 1 & 2) is devoted to the Leguminales. The Caesalpiniaceae and Mimosaceae are very sparsely represented in Iraq, mainly by introduced species; the Papilionaceae, however, with some 40 native genera and 300 species, form a significant part of the country’s flora. This family is also of paramount economic importance, since its members provide essential foodstuffs (pulses, forage and fodder crops) and many other products: gums and resins, tanning agents, fibres, vegetable oils, dyes, perfumes, medical substances and poisons and timber, as well as ornamental plants. The Flora of Iraq is unusual in placing a strong emphasis on the uses of plants, and this makes it of interest to a wide range of readers. In the present volume, for example, there are three whole pages concerned with manna and two with gum tragacanth. Volume 3 is written by the editors, working as a team: Mr Townsend is the taxonomist, while Mr Guest provides the economic and distributional notes and the Iraqi names of plants and plant products. The tribes of the Papilionaceae are delimited according to Hutchinson (1964), and yet, surprisingly, they are arranged in the order of Bentham & Hooker (1865). As was explained in Volume 2, the Flora of Iraq is modelled on Hutchinson’s (1959) treatment of the angiosperms and so it is odd to find this return to B. & H., especially as the Turkish legumes have so recently been revised and arranged in Hutchinson’s order. The latter is surely more natural; for example, it places the tribes Vicieae—Ononideae—Trifolieae in sequence, whereas Bentham & Hooker removed the Vicieae far from the Trifolieae, which were linked instead to the Genisteae. The present volume is notable for its careful, accurate and detailed species descriptions and for the fine accompanying illustrations. The format is clear and attractive and there are very few printing errors. It also continues a feature which other Floras would do well to emulate: there is a series of addenda comprising a gazeteer, a history of botanical exploration in Iraq, a glossary, indices to abbreviations, signs and symbols, author’s names and abbreviations for periodicals and other publications, and a selected bibliography. This part is not unwieldy, and is a great help to the working taxonomist. It would be useful if the frontispiece map of Volume 1 were reproduced in future volumes, so that the regions referred to in the distributional paragraphs— MAM, FPF, DLJ, etc.—could be more easily visualized. We look forward to the publication of the remaining five volumes of this admirable Flora. REFERENCES BENTHAM, G. & Hooker, J. D. (1865). Genera plantarum, 1 (2): 434-600. London. HUTCHINSON, J. (1959). The families of flowering plants, 2nd ed., 1. Oxford. HUTCHINSON, J. (1964). The genera of flowering plants, 1. Oxford. | F. K. KupicHa The Rural Tradition. W. J. Keith. Pp. xi + 310. Harvester Press, Brighton. 1975. Price £5-45. Here are 13 chapters of literary appreciation on works of 11 ‘non-fiction prose writers of the English Countryside’. They range from Walton to Williamson, and the author, Professor of English in the University of Toronto, presupposes a familiarity with their writings. His com- mentaries will no doubt be appreciated by afficionados, but on actual plants, or animals, there is just nothing. D. MCCLINTOCK Photographing Nature—Trees. Heather Angel. Pp. 96, with 24 colour and 38 monochrome photographs, Fountain Press, Argus Books, Kings Langley. 1975. Price £1-50. BOOK REVIEWS 87 In this pocket-size book, one of a series in which flowers and fungi are to be included, the author returns to a subject well-treated in her Nature Photography, Its Art and Techniques, Fountain Press, 1972. Here, one of our leading wild-life photographers has produced a practical guide to tree photography in field and studio. A mass of hints and techniques are provided and those who desire to obtain better photographic records will find the book invaluable. A basic ground- work of photography is assumed but a useful photographic glossary is included together with an equipment list, bibliography and index. Used as a recording instrument, the modern camera with its apparent simplicity has tended to produce far too many mediocre photographs. Those of us who take a camera into the field should not be satisfied in obtaining a mere record; with such a guide to technique we can only improve. Future titles are awaited with interest. G. A. MATTHEWS Flowering Plants. Evolution above the Species Level. G. Ledyard Stebbins. Pp. xviii+ 400, with 63 text figures. Edward Arnold, London. 1975. Price £13-50. When he investigates the course of evolution of supraspecific taxa and the factors involved therein, the student is denied recourse to experiment. When, as is often the case in flowering plants, he is also denied effective fossil evidence on which to make decisions, how should be proceed? At this point many workers give up the effort, muttering ‘It is all speculation’ or words to that effect. G. Ledyard Stebbins, who has already made so many original contributions to the study of various aspects of evolution, has not given up. Instead, in this fascinating and important work he gives detailed consideration to the main source of data that is available, namely evolutionary trends. Indeed, the titles of the two parts of the book speak for themselves: ‘Factors that determine evolutionary trends’ and ‘Trends of angiosperm phylogeny’. Stebbins begins by discussing certain principles of evolution, from which he proceeds to derive his own evolutionary philosophy. He believes that evolutionary trends have resulted from a succession of complex interactions between organisms and their environment, by means of natural selection; and that, in any organism, there exist several different gene combinations that might adapt it to any changed environment, the particular adaptive combination that does become established being dependent largely on the nature of the gene pool already present. In other words, recurrent natural selection results in an evolutionary trend, the direction of which depends on the state of the organism when the selective force begins to act. Having shown how trends can and do occur in any character, the author goes on to discuss how they can be recognized. In the second part, in which trends of various types in the angio- sperms are considered, he throws new and fascinating light on such familiar problems as the evolution of the flower and the origin and development of the monocotyledons. One could expatiate on so many topics that are discussed, agreeing with some of the views expressed, disagreeing to various extents with others; but the fact remains that this synthesis of Stebbins’ mature ideas on evolutionary processes and their results is a ‘must’ for anyone at all interested in evolution, despite the high cost of the book. Similarly, one could list some errors (one or two factual, a fair number typographical) that appear in the text; but these seem of relatively little importance. The format, however, makes the book less pleasing to read than it might have been, particularly the two unjustified columns per page and the reproductions of some illustrations at too small a scale. Nevertheless, this is a book to buy, to read and to keep at hand. N. K. B. RoBsON The Physical Biology of Plant Cell Walls. R. D. Preston. Pp. xiv+491. Chapman & Hall, London. 1974. Price £12-00. The appearance of a textbook of almost 500 pages on the subject of plant cell-walls and written by Professor Preston is a noteworthy event. The inevitable comparison with the late P. A. Roelofsen’s masterpiece The Plant Cell Wall (1959) is made by Preston himself in his Preface, 88 BOOK REVIEWS where he pleads his case for the restriction in subject matter that has been necessitated by the vast amount of literature which has accrued since 1959. In fact this book, which claims to deal with aspects of the plant cell-wall covered by the term ‘physical biology’, is primarily an exposition of those topics which have interested the author during nearly 50 years of research (Preston’s Ph.D. thesis, completed in 1931, was entitled The organisation of the cell wall applied to the elucidation of problems associated with growth). The first third of the book (chapters 2-6) deals with introductory physico-chemical topics, in particular chemical bonding and structure, X-ray diffraction, and polarizing and electron microscopy. The rest of the book deals with cell-wall structure (chapters 7-10), visco-elastic properties of cell-walis (chapter 11), and cell-wall growth and biosynthesis (chapters 12 and 13). As examples of the author’s self-imposed restrictions one may cite his lack of mention of scanning electron microscopy, and his confinement to algae and phanerogams. The strength of this book, however, lies in the authoritative manner in which the topics covered are reviewed and discussed by one who has been intimately concerned with the development and application of many of the relevant techniques. The reader with even a slight knowledge of the subject matter will not be in the least surprised at the relative emphasis placed by the author on the various aspects of his subject, nor will he be disappointed by it. With this slight reservation, one can safely predict that this treatise will swiftly become a standard textbook. C. A. STACE Précis de botanique (enseignement supérieur). I. Protocaryotes et thallophytes eucaryotes. R. Gorenflot. Pp. 184, with 108 figures. Doin, Paris. 1975. Price Fr. 98-00. This is an original and remarkable text. It gives first a good account of the protocaryotes, though not treating the bacteria in great detail. It continues with eucaryotes in three large chapters of equal length. These deal first with the morphology and anatomy of the thallus, secondly with reproduction, and thirdly with thallophytes in the environment and the biosphere. In each of these three chapters, topics are treated in a perfectly general way, cutting across the systematic categories. For example, vegetative reproduction, types of life-history, cell structure, symbiosis, are all considered synthetically, and not as separate algal and fungal fractions. The book ends with a short chapter on the main lines of classification of the leading algal and fungal groups. It is copiously and excellently illustrated, with many original drawings and diagrams and some good electron micrographs. It is packed with information and is well written and printed. The level is about that of a second year course in a British university. Expensive, but highly recommended. D. H. VALENTINE Watsonia, 11, 89-92 (1976). 89 Obituaries CATHERINE MURIEL ROB (1906-1975) Kit Rob died on 6th February, 1975 as the result of a grave illness of some months’ duration. She died at Catton Hall, near Thirsk, where she was born on 21st February, 1906. Educated at home by a governess, she later carried heavy domestic responsibilities for very many years, ultimately caring first for her sick mother, who died in 1959, and then for two elderly aunts, the last one dying in her hundredth year in 1974. Once a part-time dispenser and then a lecturer, she was latterly employed full-time as a member of the Yorkshire Rent Assessment Panel. During the war she served as a cook at Catterick Military Hospital. A noted, highly experienced amateur field-botanist, Kit was also a great character and a wonderful friend to many. With a sound working knowledge of the British flora, she was the unrivalled authority on the distribution of vascular plants in North Yorkshire for at least 36 years. Self-educated botanically, she was justly proud of being elected a Fellow of the Linnean Society in 1947 in recognition of her services to Yorkshire botany. Permanent testimony to her knowledge and continual effort ‘in the field’ is to be found in her numerous contributions to The Naturalist, particularly the valuable accounts of the plants of specified North Yorkshire localities in excursion circulars and reports from 1935-69. Her greatest contribution to botanical knowledge is to be found in the Atlas of the British flora. She was responsible for collecting records for the whole North Yorkshire area (v.c. 62 and 65), a tremendous task, and yet this is one of the best-covered areas due mainly to her own great personal enthusiasm and effort. Her account of the plants of the moors and dales in the Forestry Commission Guide North Yorkshire forests will continue to be of great help and interest to the visitor to the area. Her great knowledge of folk-lore and English names was invaluable in the preparation of the recently published English names of wild flowers (Dony, Perring & Rob, 1974). Kit had an important private herbarium which contained 5,000 specimens in 1959. Kit’s interest in wild plants had begun early in life. She joined the Wild Flower Society at the age of 17 and it was to her activities in this Society, together with the personal help and encourage- ment of Tom and Gertrude Foggitt, that she owed her wide basic knowledge of plant identification. Her active membership of four societies in particular was of great mutual benefit to the societies concerned. Always an outstanding member of the Wild Flower Society, Kit soon worked her way up to the heights of ‘Valhalla’, becoming secretary of its lower section in 1930 and succeeding Mrs Foggitt as secretary of the upper section in 1949, a position she held until 1971. A succession of Presidents have been grateful for her help and advice always readily given. One of the longest- serving members in the B.S.B.I., Kit joined the Botanical Society and Exchange Club of the British Isles in 1932, soon became a regular exchange member and was promptly made local secretary for North Yorkshire. It was a tribute both to her knowledge of British plants and to her ability to handle people tactfully that she was appointed distributor in 1946. When exchange was abandoned, referees and recorders were appointed in addition to local secretaries and Kit became all three for North Yorkshire until two of these offices ceased to exist; she was still recorder at the time of her death, as well as referee for popular names. She served on the Council of the B.S.B.I. as representative of the North-East Region from 1950-3, 1956-60 and from 1967-71, and was Vice-President from 1960-4. She joined the Yorkshire Naturalists’ Union in 1934 and became one of its most prominent and highly respected members. She held important offices including that of Hon. General Secretary from 1958-62, Hon. Excursions Secretary, 1963, and Divisional Secretary from 1939-63. She was plant recorder for North Yorkshire from 1938-71 and was responsible for the publication of plant records for all Yorkshire from 1957-65. She was elected President of the Union in 1969 being only the third woman to be so honoured. Always interested in conservation, she took a keen interest in the Yorkshire Naturalists’ Trust from the time of its foundation in 1946. She was Hon. Secretary of the Trust from 1952-7, when it owned 90 OBITUARY only two nature reserves, and she laid the foundations for the much wider influence that it gained subsequently. Since 1957, she continued to serve the Trust in many ways: as a council member from time to time, as chairman of two reserves and as a member of the north-west area group committee. She had also been a member of the council of the Yorkshire Philosophical Society and was its secretary from 1954-9. Both botanizing and meeting botanists were enjoyed to the full. Kit organized excursions for the B.S.B.I. and the W.F.S. and annually for the Y.N.U. over a long period of time, all in her own inimitable style. She was in great demand as a lecturer and gave generously of her time to various Natural History Societies, often travelling long distances, and in recent years she con- ducted adult education lectures, which were well attended. Completely at ease in front of a microphone, Kit took part in radio programmes concerned with the northern country-side, and the invitation to take part in the television series “The North-Country Naturalist’ was recognition greatly appreciated. What Kit Rob was able to do for Yorkshire botany and conservation owes as much to her personal qualities as to her botanical knowledge and even her many other interests played their part. Her own warmth of personality and genuine interest in others meant that she made friends with people in all walks of life, including land-owners and others of influence. Widely known and respected, her name was household, not only in Yorkshire but far beyond its boundaries. There was great satisfaction in honest endeavour and achievement. Not only did Kit become an eminent amateur botanist, but also in recent years a highly successful breeder of dogs which won prizes at shows all over the country, one of her Cardigan corgis winning a best-of-breed award at Cruft’s. The amount of knowledge which Kit Rob disseminated during 40 years or so is inestimable, and this was particularly valuable in the case of the more critical species and aliens. She is remembered by botanists all over the British Isles for her patient, kindly and valuable help to them, especially as beginners. The effect of her influence in stimulating botanical field-work in the whole of Yorkshire is incalculable. Personally I owe her much; soon after I took up field botany seriously we became regular correspondents and there were frequent invitations to stay at Catton Hall to examine her fine herbarium or to use her extensive library. Numerous botanists, both experts and humble beginners, have enjoyed similar generous hospitality. On those special occasions when one needed the support of a staunch friend Kit was somehow reassuringly there. She also did much to enrich the life of her friends by bringing into contact those who could be mutually helpful. Arrangements have been made for her herbarium to be sent to the offices of The Yorkshire Naturalists’ Trust Ltd., 20 Castlegate, York. A measure of the high regard and great affection with which she was held was demonstrated at the funeral service when the local parish church at Skipton Bridge was full to overflowing on a very foggy day. The B.S.B.I. was represented by David McClintock and others. Kit Rob will be greatly missed. We were privileged to know such a unique character; her spirit of service and friendship and her great sense of humour will be remembered. I am indebted to Dr J. G. Dony; also to Charles Rob, Mrs V. Schwerdt of the Wild Flower Society and Clifford Smith of the Yorkshire Naturalists’ Trust for assistance in the preparation of this appreciation. F. E. CRACKLES VICTOR SAMUEL SUMMERHAYES (1897-1974) With the death of Victor Summerhayes on 27th December 1974, at the age of 77, the B.S.B.I. has lost a most valuable member and I have lost a personal friend of long standing. When I first went to Kew in 1928 I soon got to know Victor, who by that time was well established on the permanent staff in the Herbarium, where he was in charge of the systematic Orchidaceae section as well as the geographical Australasia and Polynesia section, both of which he was expected to.run single-handed! In the following year, as general herbarium dogsbody, I was given the task of naming the extensive collection of plants from Fiji made by W. F..N. Greenwood, and so came OBITUARY 91 under Victor’s personal tuition, when I was greatly impressed not only by his thoroughness and attention to detail, but by his great patience in initiating a tiro into the finer points of taxonomy and nomenclature. I soon found common ground with Victor in our mutual interest in ecology and it was not long before I was invited to join the team of volunteers who were working on an ecological survey of Richmond Park under W. B. Turrill. Victor was keeping most of the records of the survey, and I used to spend many enjoyable evenings in his home in West Park Avenue, Kew, helping by marking the distributions of plant species on to record maps, while Victor wrote up the notes. Here, in Victor’s home, I soon got to know his wife, Dolly and their two boys, John and Gerald. I have very happy memories of those evenings, when the work was followed by interesting conversations and discussions and an excellent light supper provided by Dolly. I was not to work for long on Fiji, for early in 1930 I was told I would be seconded to the Colonial Office for attachment in May to the Aerial Survey of Northern Rhodesia. Victor was interested to learn about the orchids I came across in Africa, very few in fact as the expedition was held during the dry season. The following year I exchanged my motor cycle for a B.S.A. 3-wheeler, and I was able to take Victor into the country to study orchids. One of the most memorable of our many outings was one to Dorset when, following an early start, we met Sir Maurice Abbot-Anderson at 10 o’clock at Wareham for a conducted tour of various marsh orchid sites, ending up at Toller Porcorum. After a late tea at Beaminster we drove 135 miles back to deposit a very weary Victor at his home. Victor was particularly interested in hybrid swarms of Dactylorhiza species and I learnt from him the technique for studying such populations, a technique I found most useful when I came across hybrid swarms of orchids and other plants in Africa. Dolly and Victor spent much of the Second World War at Kew, when many of his colleagues were evacuated with sections of the Kew herbarium collections to Oxford or Gloucestershire. They luckily escaped injury when one of the first V2 rockets fell on the nearby Chrysler Works and shattered all the windows and brought down all the ceilings but one of their home, for the kitchen ceiling had been boarded up following an earlier bomb. After the War, Victor started his intensive field work in preparation for his Wild orchids in Britain published in the New Naturalist series (Summerhayes, 1951). This book is packed with original observations obtained by Victor personally during this period. Trained largely as an ecologist, he paid particular attention to the nature of the habitats. He visited as many as possible of the species in the field, making notes of the plants commonly associated with each. He obtained willing help from a number of B.S.B.I. members, and became closely associated with the late Donald Young in his detailed study of Epipactis. When the B.S.B.I. launched its mapping scheme, Victor Summerhayes joined in, and he and I spent several holidays recording in various parts of Britain, including Wales and southern Scotland. He had a very sound knowledge of the British flora, and his ecological training had enabled him to identify many plants, including a large number of grasses, from sterile material. The highlights of Victor’s early career and of his ecological work have been well covered by Brenan (1975). I would, however, like to emphasise the tremendous amount of background work which resulted in his paper on the effect of voles on vegetation (Summerhayes, 1941). For several consecutive years he took leave in the summer and travelled by road with his friend, A. D. Middleton of the Bureau of Animal Population, Oxford, to Newcastleton in Roxburghshire and Corris in Merioneth, where he carried out, meticulously, the sampling of the vegetation in areas of upland grassland, often under most unpleasant weather conditions. Victor Summerhayes had a delightful sense of humour, and he saw the funny side of many situations. One summer we were recording for the B.S.B.I. in a remote square in, I think, Roxburghshire, when we came across a field full of arable weeds. We had not been in it long when the farmer approached to find out who we were and what we were doing. Victor, with a surreptitious wink, put the fear of God into him by telling him we were on the staff of the Ministry of Agriculture, and so we were! The farmer was clearly worried, but did not feel he could turn us off, so we went on recording satisfactorily, whilst he went away expecting he would receive a rocket for having a field in such a neglected state. Victor thoroughly enjoyed an incident such as this. When, early in the fifties, a large part of the only British site for Orchis simia in Oxfordshire was destroyed by ploughing, Victor was one of the volunteer conservationists who visited the 92 OBITUARY remaining part of the site one Sunday each winter to keep it clear of scrub. He took part in this annual activity organized jointly by botanists at Kew and Oxford until 1964, when he retired and went to live in Devon. SORE I am refraining from dealing, here, with Victor Summerhayes’ outstanding work on the African Orchidaceae. This is well covered by Peter F. Hunt in Orchid Review (1975). I would, however, like to mention his tremendous foresight and tenacity in starting and building up a collection of orchids and other fleshy or zygomorphic flowers and of berried fruits preserved in spirit solution. For years it was a question of putting into and getting nothing out of the collection and, with no assistant, undaunted, Victor did much of the initial labelling of the bottles and specimens himself. Time and again he resisted pressure to discontinue the collection. When I went out to Northern Rhodesia in 1937 he asked me to collect specimens in spirit, and I believe my collection from Mwinilunga was the first major Kew collection to be accompanied by material in spirit. This has now become normal collecting procedure, resulting in the Kew spirit collection being far ahead of any other such collection in the world, and in families like Orchidaceae, Asclepiadaceae and genera such as Utricularia the chances are that the plant one needs to study will be represented. Victor was a small man with slight physique, but mentally always very much alive. He had a wonderful habit of gesticulating when, in conversation, he was describing the features of a particular orchid flower which he imagined before him enormously magnified! His clenched fist would be the rostellum whilst with his other hand he would outline the relative position of the pollinia and other essential parts. Another of his characteristics was his use of the word ‘svecies’ instead of ‘species’, a rather delightful pronunciation the reason for the use of which I never discovered. His wonderful botanical hat, which rivalled mine for dilapidation, had many amusing vicissitudes. More than once it parted from its owner when speeding in Donald Young’s open MG, and on one occasion it was abandoned in a pub at Carmarthen when we were staying at Haverfordwest, necessitating a return journey to Carmarthen to collect it next day. I remember one night, as I was leaving their home, Dolly commented on the disgraceful state of Victor’s hat on the hat-stand in the hall, only to dissolve into a state of acute embarrassment when I claimed the offending object! B.S.B.I. members who consulted him always found him helpful, and he had many friends among its members. His loss will leave a gap in British botanical circles that will be hard to fill. REFERENCES BRENAN, J. P. M. (1975). Victor Summerhayes: Obituary. Bull. Brit. Ecol. Soc., 6(2): 5-6. Hunt, P. F. (1975). The late Mr V. S. Summerhayes, O.B.G. Orchid Rev., 83: 77-78. SUMMERHAYES, V. S. (1941). The effect of voles (Microtus agrestis) on vegetation. Journ. Ecol., 29: 14. SUMMERHAYES, V. S. (1951). Wild orchids in Britain. London. E. MILNE-REDHEAD Watsonia, 11, 93-95 (1976). 93 Reports ANNUAL GENERAL MEETING, MAY 3lst, 1975 The Annual General Meeting of the Society was held in the lecture room of the Loreburn Primary School, Dumfries, Scotland, on Saturday, 31st May, 1975 at 12 noon. 40 members were present, with Dr S. M. Walters (retiring President) in the Chair. The minutes of the last Annual General Meeting, as published in Watsonia, 10: 321-322 (1975), were adopted unanimously. REPORT OF COUNCIL The adoption of the Report for the year 1974, as circulated to members, was proposed by Mr A. C. Jermy, seconded by Dr F. H. Perring and carried unanimously. TREASURER’S REPORT AND ACCOUNTS Mr J. E. Lousley, in supporting the Treasurer, expressed the hope that the Society would not be rushed into precipitous action in effecting economies. He felt that the Accounts showed no cause for undue despondency. The Treasurer hoped that Mr Lousley would be proved right, and the Secretary reported that the decision on any significant change in our publications had been postponed until Spring 1976 when the saving in costs through changes in format could be determined. The Treasurer reported that one Warburg Memorial Fund Award of £75 had been returned, and that a number of items on the 1974 expenditure account were non-recurring. The adoption of the Treasurer’s Report, together with the Accounts for the year 1974, was then proposed by Mr J. E. Lousley, seconded by Mrs O. M. Stewart, and carried unanimously. INCREASE OF MEMBERSHIP SUBSCRIPTIONS The Treasurer proposed an increase in subscriptions from Ist January, 1976, as circulated to members: Ordinary £5 (from £3) Family 75p (from S5Op) Junior £2 (from £1) Subscriber £5 (from £3) This was seconded by Miss M. Dickinson and, with some reluctance, was approved by the meeting and carried unanimously. ELECTION OF PRESIDENT Dr S. M. Walters from the Chair welcomed the nomination by Council of Mr E. L. Swann. This was seconded by Mr R. J. Pankhurst and unanimously carried. Mr Swann then took the Chair, expressing thanks to Dr Walters on behalf of the Society for his wise counsel! and active furtherance of the Society’s aims during his Presidency. ELECTION OF OFFICERS The re-election of Mrs H. R. H. Vaughan, Dr W. T. Stearn and the election of Dr C. T. Prime as Vice-Presidents was proposed by Miss M. McC. Webster, seconded by Miss E. R. T. Conacher and carried unanimously. The following officers were then proposed and elected without dissent: Mrs M. Briggs Honorary General Secretary, proposed Miss E. Young, seconded Mrs A. H. Sommerville; Mr M. Walpole Honorary Treasurer, proposed Dr S. M. Walters, seconded Mr B. W. Ribbons; Dr G. Halliday, Dr N. K. B. Robson and Dr C. A. Stace Honorary Editors, proposed Mr E. F. Greenwood, seconded Dr R. W. M. Corner; and the following officers en bloc: Dr J. T. Williams Honorary Meetings Secretary, Mrs G. Beckett Honorary Field Secretary, Mrs R. Hamilton Honorary Membership Secretary, proposed Dr A. J. Richards, seconded Miss E. J. Rich. Dr C. T. Prime and Dr J. T. Williams were welcomed as new officers, and warm thanks were expressed to all officers for the great deal of work and time given to the Society’s 94 REPORTS affairs. Particularly thanked was Mr E. F. Greenwood who retired after 9 years as an Honorary Editor. ELECTION OF NEW MEMBERS OF COUNCIL Nominations had been received for Professor D. H. Valentine, Mr G. Messenger, Mr B. S. Brookes and Mrs Y. Heslop-Harrison to fill the four vacancies on Council. These members were unanimously elected and their order of precedence, as given, was determined by ballot. ELECTION OF HONORARY MEMBER Council had recommended the election of Mrs J. G. Dony, a member of the Society since 1948 and Honorary Membership Secretary from 1964 to 1974. The President, proposing from the Chair, said this would be the first time a botanist and his wife were both Honorary Members of the Society. The nomination was seconded by Mr R. Mackechnie and carried unanimously and with pleasure by the meeting. The nomination had not been a surprise to Mrs Dony (Dr and Mrs Dony proof-read the Annual Report) and a letter of thanks if elected, sent by Mrs Dony, was read. ELECTION OF HONORARY AUDITORS Council’s recommendation for the re-election of Messrs Thornton Baker & Co. was proposed by Mr M. Walpole, seconded by Mr J. E. Lousley and carried unanimously. The Treasurer pointed out that auditing our accounts represented a loss of business to the auditors, and it was agreed that the Society was most grateful to them for the continuance of their valuable service. ANY OTHER BUSINESS Miss M. McC. Webster raised the problem of botanical communications in northern Scotland. Mr R. Mackechnie commented on the thin spread of membership in that area, and Mrs A. H. Sommerville said that the C.S.S.F. had done marvellous work in co-ordination. Some B.S.B.I. members thought that there was a lack of representation and an undertaking was given to refer this to the B.S.B.I. Co-ordinating Committee for consideration. The Director of Education for Dumfriesshire was thanked for permission to use the school; the C.S.S.F., in particular Miss E. R. T. Conacher, C.S.S.F. Honorary Meetings Secretary, and the local botanists and friends who had helped organise the meeting were appreciatively applauded. The meeting closed at 13.05. M. BRIGGS COMMITTEE FOR THE STUDY OF THE SCOTTISH FLORA TWENTIETH ANNUAL REPORT TO 31ST DECEMBER, 1974 The Committee met twice in Perth and arranged an Exhibition Meeting at the University of Glasgow (by kind permission of the Regius Professor of Botany). 11 field meetings were included in the Scottish programme for the year. The Exhibition Meeting was held on Saturday, 2nd November, 1974 and about 100 people were present. The following exhibits were shown: posters on plant conservation and Society activities (B.S.B.I.); Ruppia cirrhosa and other aquatic plants from Uist (Mrs J. Clark); photo- graphs of places visited in Easterness during the C.S.S.F. survey (Miss E. R. T. Conacher); some sedges and grasses of Ireland (A. Copping); Glen Affric, 1974 (R. W. M. Corner); six species new to East Ross (Miss U. K. Duncan); mapping in 1974 of Groenlandia densa, Littorella lacustris, Myosurus minimus and Orchis morio (Miss L. Farrell); Ranunculus flammula? subsp. scoticus, other records and maps from the Glen Strathfarrar field meeting, 1974 (D. Kingston); New Zealand Epilobium spp. in Scotland, Gunnera in the British Isles and Polygonum? molle in Britain (D. McClintock); progress in and publications concerned with the C.S.S.F. Inverness-shire Survey (B. W. Ribbons); a Wardian case: an aid to plant study (Miss A. Rutherford); flower paintings and records from Kirkcudbright (Mrs O. M. Stewart); variation in frond morphology in Polypodium australe, and Rubus scoticus, a recently described bramble of the Clyde area (A. McG. Stirling); Hebridean and Icelandic plants (E. C. Wallace); the Eildon Hills: a vegetation study (Scottish REPORTS 95 Wildlife Trust, Tweed Valley Branch). After tea, Dr S. M. Walters, Director of the University Botanic Garden, Cambridge, and President of the B.S.B.I., spoke on priorities in botanical conservation. Following a buffet supper in the University of Glasgow Refectory, colour trans- parencies of C.S.S.F. field meetings held in 1974 and of the meeting in Lapland in 1973 were shown by Miss E. R. T. Conacher, R. W. M. Corner, D. Kingston, R. Mackechnie, J. D. S. Martin, B. W. Ribbons, Mrs A. H. Sommerville and R. E. Thomas. Of the 11 field meetings in the programme, two were cancelled (Strathdearn and north-western Lochaber); one (to Beinn Laoigh) was arranged on 29 June by the Alpine Botanical Secretary of the B.S.E. (for report see B.S.E. News no. 14); and the other meetings were held at Brodick, Isle of Arran, Clyde Isles (21-24 June with the Andersonian Naturalists of Glasgow) led by Mrs A. H. Sommerville (11), in Mamore Forest, Westerness (28 June—1 July) led by A. A. P. Slack (5), at Fort William, Westerness (13-20 July) led by A. A. P. Slack (8), in Glen Affric Forest, Easterness (20-27 July) led by R. W. M. Corner (4), at Tomatin, Easterness (20-27 July) led by Miss E. R. T. Conacher (3), at Kindrogan Field Centre, East Perth (24-31 July) led by A. C. Jermy (11), in Glen Strathfarrar, Easterness (27 July-3 August) led by Miss E. R. T. Conacher (7), and at Fort Augustus (3-10 August) led by H. A. McAllister (5). The attendance figures are given in brackets. For reports of these meetings see Watsonia, 10: 456-462 (1975) and B.S.E. News nos 15 and 16. The eight elected members appointed Mr G. H. Ballantyne, Dr U. K. Duncan, Mr A. A. P. Slack and Mr A. McG. Stirling as members of the committee for the calendar year 1975. Mr R. Mackechnie was appointed Chairman, Mr B. W. Ribbons Honorary Secretary, Mrs A. H. Sommerville Honorary Treasurer, Miss E. R. T. Conacher Honorary Meetings Secretary, Mr A. McG. Stirling Honorary Field Meetings Secretary and Mr R. Mackechnie a member of the B.S.B.I. Conservation and Records Committees for 1975. Mr Stirling resigned as Honorary Field Meetings Secretary in October and in December Mr B. S. Brookes was appointed to this office. The Chairman was represented at B.S.B.I. Council on 19 February and 29 October by Mr Ribbons and on 4 May by Miss Conacher. He was represented by Mr Brookes at the B.S.B.I. Conservation Committee on 26 September and by Mr Ribbons at the B.S.B.I. Records Com- mittee on 25 January and 30 August. The Organizing Committee for the Inverness-shire Survey met twice in 1974, in February and September; all five members attended the earlier meeting and four were present in September. Number 3 of the Inverness-shire Newsletter was issued in February. The progress of the survey was illustrated by an exhibit prepared jointly by the recorder for Easterness and the Honorary Secretary of the C.S.S.F., and mounted at the C.S.S.F. Exhibition meeting in Glasgow early in November, and again at the B.S.B.I. Exhibition meeting in London later in the same month. A working party was appointed to arrange for the publication of the results of the Survey. The list of recorders for the Scottish vice-counties was kept under review and recommendations were made to the B.S.B.I.; no entries for the Red Data Book were received for checking during the year; a leaflet briefly outlining the aims and functions of the Committee was printed; arrange- ments were made for the forthcoming Annual General Meeting of the B.S.B.I. in Dumfries. Various administrative matters were discussed with the officers and councils of the parent societies. The accounts showed a deficit of £37.90. This has been refunded in equal sums by the two parent societies who also made grants to cover the cost of printing the leaflet on the aims and functions of the Committee. The Committee wishes to express thanks to Mr A. McG. Stirling for serving as Honorary Field Meetings Secretary 1971-74, to the officers of the parent societies, to the leaders of the field meetings and the exhibitors at the Exhibition Meeting and to other members and friends who helped the Committee during the year. B. W. RIBBONS FLORA OF A CHANGING BRITAIN Edited by F. H. Perring The Report of the 1969 Conference which reviewed factors responsible for changes in the British flora, particularly since the end of the Second World War, and endeavoured to predict other changes which might take place between the present time and the end of the century. Topics covered included the effects on the flora of climatic changes, atmos- pheric and other pollution, changes in transport systems, etc. PLANTS WILD AND CULTIVATED Edited by P. S. Green An account of the Conference held by the Royal Horticultural Society and the Botanical Society of the British Isles in September 1972. All the papers read at this Conference on horticulture and field botany are included. Demy 8vo., 231 pages, with 8 half-tone plates, and figures in the text. THE CONSERVATION OF THE BRITISH FLORA Edited by E. Milne-Redhead The Report of an important and most successful Conference held by the Botanical Society of the British Isles at Durham in April 1963. This report includes most of the papers read at the Conference, together with a verbatim account of the discussions that followed. Demy 8vo., 80 pages. Bound in buckram Details from E. W. CLASSEY LTD, PARK ROAD, FARINGDON, BERKSHIRE, SN7 7DR INSTRUCTIONS TO CONTRIBUTORS PAPERS and short notes concerning the systematics and distribution of British and European vascular plants as well as topics of a more general character and wider appeal are invited from both members and non- members of the Society. Careful attention to these notes before a paper is submitted can save the author and editors much work, and do much to speed publication. Papers must be submitted in final, fully-corrected form. They should be typewritten, with wide margins, double-spaced, on one side of the paper only. If possible, two copies should be sent, of which one must bea top copy. Authors should keep a fully-corrected carbon copy of their typescripts for reference. Write clearly and concisely, rigorously excluding extraneous matter. It sometimes helps the clarity of an argument to place extensive experimental or tabular data in appendices, rather than in the main text, Technical terms are of value only as aids to clarity, precision and conciseness of expression. Take care that they are used consistently, and defined where necessary. Ill-defined or general terms often need definition in a particular context, and can often with advantage be replaced by something more precise, as by the ‘deme’ terminology of Gilmour and Gregor in genecology. Make certain that the main conclusions of the paper are clearly displayed to the reader. Avoid complicated hierarchies of headings, and check carefully the consistency of those that you use; a table of contents, setting out the full hierarchy of headings with the MS. page numbers, is often helpful even if it is not to be printed with the paper. Avoid footnotes as far as possible, and keep cross references by page number to a minimum. Tables, unless very small, should be typed on separate sheets and attached at the end of the typescript. They should be kept within a reasonable size, and as simple in structure as possible. Keys should be in one of the generally accepted forms; Flora of the British Isles by Clapham, Tutin and Warburg (2nd ed., Cambridge, 1962) and Flora Europaea (Cambridge, 1964-72) provide suitable patterns. Names of genera and species should be underlined, but any other typographical indications should be inserted lightly in pencil. Names of vascular plants should normally follow the List of British Vascular Plants by J. E. Dandy (British Museum (NH) and BSBI, London, 1958) and changes published in Watsonia, 7: 157-178 (1969) or Flora Europaea (Cambridge, 1964-72) and may then be cited without authorities. Other- wise authors’ names must be cited, at least on the first occasion where they appear in the text and followed by the ‘Dandy’ or Flora Europaea name in parenthesis. Authors must follow strictly the International Code of Botanical Nomenclature (Utrecht, 1972). Synonyms should be cited in chronological order; misapplied or illegitimate names should be placed in square brackets. Examples will be found in recent parts of Watsonia. Except for citations of the place of publication of plant names, which may be given in abbreviated form in the text, full references should be listed in alphabetical order of authors’ names at the end of the paper; the form used for references in a recent part of Watsonia should be followed carefully. Names of periodicals should be abbreviated as in the World List of Scientific Periodicals (4th ed., London, 1963-1965). References to herbaria should follow the abbreviations given in British Herbaria (BSBI, London, 1958) and Index Herbariorum Part 1 (Sth ed., IAPT, Utrecht, 1964). Foreign language references should, where necessary, be transliterated into the roman alphabet according to a recognized convention; for Cyrillic script (Russian, etc.) the Royal Society (BS 2979: 1959) or UNESCO schemes are acceptable. Unless authors expressly indicate that they cannot get access to the necessary literature, the editors will assume that they have checked the correctness of all titles, abbreviations, transliterations and references. Papers should begin with a short abstract, in the form of a piece of connected prose conveying briefly the content of the paper, and drawing attention to new information, new names and taxa, and the main conclusions. Line-drawings should be boldly drawn in Indian ink on Bristol board, smooth surface white cartridge paper of good quality, or suitable draughtsman’s tracing materials, and should normaily be suitable for reproduction at about one-half to two-thirds (linear) their original size. Very large originals should be avoided. Scale is best indicated by a rule on the drawing itself. lf a magnification is quoted this should be checked carefully against the final block. Graphs can be reproduced from originals on graph paper with feint blue ruling, but all lines to appear on the finished block must be inserted in Indian ink. Lettering on line-drawings and graphs should be inserted lightly in pencil, and will be finished in uniform style. The following standard symbols are available for graphs, etc.: A, &, O,O, 0, +, x, ©. Avoid other symbols as far as possible. If an illustration includes plant names or place names, these should be typed clearly on a separate sheet of paper. Photographs can be accepted only where essential. They must be of first-rate technical quality, of good but not excessive contrast, and of a size and character suitable for the necessary reduction. Remember that fine detail may be lost on even the best half-tone blocks. Legends to text figures and plates should be typed on a separate sheet of paper and attached at the end of the typescript. If in doubt about the citation of names or references, or the presentation of illustrations or tabular matter, contributors are advised to consult the editors before submitting their typescripts. Further useful information and advice will be found in General Notes on the Preparation of Scientific Papers (2nd ed., Royal Society, London, 25p.). Proofs. Authors will normally receive a galley proof. Particular care should be given to the thorough checking of references and tables. It should be assumed that only actual errors of typography and fact can us ae in proof; as far as possible any words of phrases deleted should be replaced by others of equal ength. Twenty-five offprints are given free to authors of papers. Further copies may be obtained at the Society’s current price, and must be ordered when the proofs are returned. The Society as a body takes no responsibility for views expressed by authors of papers. Papers and short notes should be sent to Dr G. Halliday, Dept. of Biological Sciences, The University, Bailrigg, Lancaster. Books for review should be sent to Dr N. K. B. Robson, Dept. of Botany, British Museum (Natural History), Cromwell Road, London, SW7 5BD. Plant records should be sent to the appropriate vice-county recorders.