Pod ws wade 9 hs a a ee ved wk Se ah NI Na EL ae nee, pw re poh fie ge hy AONB Dec ae eer er HH TOONS OR INV ewed PD AA oa Thad OL mA ea ® Cb as tet Se ee Spe teeiy Vhe tbat g Ne MOIS od ab aN wey VPN aN een, te wey WA CPO TL Dee S 3 yan nv Bereeny WAVE Nay hay Ce en phates Ce ere a See ey a SERN me aan es Ty davay bavy mh ee REN Wyn bey VAs aay est Pec ree VM oe ub at we whet Ragen VM OT AC yay wos vas “y ee LO cae 0 Ly “ se Mate a 8 sae Wey Neath ati Bele dha VON 6 Word ig a hanes Belg Haaay ot gb Yaany wiv Se PVN heey HM mls NA by es 180 mb" eae ery Wo al gh m1 MN hy NUM N NOH at Whew ey Ah Athy whe NS Yh tits Ce) OMENS Ay VAR eh MM AS UN BO Thom ed gn Ok Peel ab ab ie je visas Chehab nb oh tiey Ved Metity Me wy PW pth ih ahah bod eth aby ie nD Pere aga om ah ide dvd hes gt WAb aN ey ite Veh ac ant Oto SEAR wh Om Mei ha’ Peer ae A Se ee reer a os eran ELTON Te ner Wee ee PME MIEN swt eat Pow ere pita gst roe dhe pt Ph PA ibe So dyin Scar Oa ny P ee aay ee Pee iit Ser aunayy nv COUN re oh th REL eA ai, OA Teer as eee 8 hey ee ere MD WAND A baal 4 3 P'Ve Wn gthastt Ree) HOM Gale’ ere arn too er Cree a ayy AD elle hall uy wht at sh “ eh eer othe “teal Me A i i M ahiy Sramet WS ater tliat athe ONC hor eat Ld em Pd act AE ON lh MVM aate vil, AN, ba ine Rin 0 Pt lig MM Ate ah beta turmetids ha Mahon whi Bi wed ino tb ch Seah P Wty YW Mot Rn yt BAN a 8 A aWall w Machel wee Bm ali adh Se Me MaldeM beth tan thy: CN nihil UM Th ane Nit Med Ree hth Mn 6 talon Sat att BEN WALA MAM a Me diet Fumes BNA ahr bie oth Or a er] Be Sr Ler es “ wh Ati de ‘ © 1 a a Beate SAM Me Awd wd Maite sla aM ibe Madd steph ‘ a We ee Mh He baal Vath hg olathe at wha eh Da wilt eS BL, ath Me bo hi MiNi din s'llanht HM os 6M OM aMaivalbeatiainnrde Mala vaate NT AO BL AAR A IN sc ah deren rf ow SO tha ‘ See er ee Mee NM atlas +8 gion Monee ALA gle Sta ry Went ale vay RO 1 Vet amenities din! wad, SAM ad ah asa ely ou al day’h VPS alt ait uhh Wo 4 dibid nl Aya ya ‘at edit Su ath AM IMM Mel Mieke sos Pes rl hap Meh Boag uw eH aL Ne HM allel va Cie ag MPa vw erat “ ao non Teer) ars ae Nee ae Vinee or FMA ata eth Hig Win tn vow i 32 iii 105 172 621 OF THE GRAY HERBARIUM Digitized by the Internet Archive In 2014 https://archive.org/details/watsonia18bota WATSONIA THE JOURNAL OF THE BOTANICAL SOCIETY OF THE BRITISH ISLES VOLUME 18 EDITED BY J.R. AKEROYD, J. R. EDMONDSON, R. J. GORNALL C. D. PRESTON anp B. S. RUSHTON 1990-91 PRINTED FOR THE SOCIETY BY Eaton Press Limited, Wallasey, Merseyside PUBLISHED AND SOLD BY THE BOTANICAL SOCIETY OF THE BRITISH ISLES, C/O DEPARTMENT OF BOTANY, BRITISH MUSEUM (NATURAL HISTORY), LONDON SW7 5BD v ee ace 13028 TA" ety ¥ AT 68 3H ; eq Jet Haris it MHT F "a % é — ’ * A é : » ¥ | = » jie / -T Sy aie a fie a | oi 7 pe Bi SMUL0Y as *y s , i (iu) 7 $] ‘ re" a4 Ni‘ Ls mt o% YOR: HA. aa _ - ’ : ® : a * 7 - ; » a y a ws vY'l , ’ ‘ 4 ! 23g ra -_ : : i] é ‘ » v4 A ® | et oo Ad To i ~ zr STATOR R MUWTAGiEL Ue tit eeiguh oad YCRSVINU ORAVAAN on ye : hepeesh tA. 8 rat on te ; Buté -_ i _ oe oS ois > oo ooo DATES OF PUBLICATION Part 1, pp. 1-118, 26nd February 1990 Part 2, pp. 119-246, 21st August 1990 Part 3, pp. 247-332, 27th February 1991 Part 4, pp. 333-452, 20th August 1991 . 23; line 19 up. Tt. line’ 26. . 71, line 15 up. . 75, line 14 up. 76, line 23 up. 79, line 6. - Ohne Sup. 3/97 line 20. . 181, line 12. a al & hig 0 SAF Errata For: For: For: For: For: For: For For ALBONIS bella-donna viridus viridus viridus Hippurus : officinalis For: For: americanum Sinapsis : greyli read: read: read: read: read: read: read: read: read: read: ALBIONIS belladonna viridis viridis viridis Hippuris sagittifolia americanus Sinapis grayl v AUT 27 1 baas a! P - ‘’ « - +. » >t a] regA tei< 08 ea wie + 7) gaa gis C1 anil pra ees SS anlf fT .g- "ees ‘yu 21 onil ny Y wry qu &L oni} :20 . ae ott OF 0 snil mi CONTENTS MILNE-REDHEAD, E. The B.S.B.I. Black Popular survey, 1973-88 MCALLISTER, H. A. & RUTHERFORD, A. Hedera helix L. and H. hibernica (Kirchner) Bean (Araliaceae)) in the British Isles BINGGELI, P. Detection of protandry and protogyny in syeamor aks pseudoplatanus L.) from infructescences “pe t ALLEN, D. E. The Rubus flora of the Isle of Wight va KELLy, D. L. Cornus sericea L. in Ireland: an incipient weed of Geiltad Fotey, M J. Y. The current distribution and abundance of Orchis ustulata L. in southern England GosLer, A. G. Introgressive hybridization oe Crataegus monogyna Jacq. and C. ils eres DC. in the ae ae Thames sept England CRACKLES, F. E. ‘A leeion x ‘eon Lamotte nm. pipe eee: in Yorkshire, with special reference to its spread along railways GULLIVER, R. L. The rare plants of the Howardian Hills, North Yorkshire, 1794-1988 ... e:, a ry oh fy. 4. A, SHoRT NOTES Allen, D. E. Micromorphy in New Forest Rubi Brewis, A. Yellow Ivy Broomrape & - Dalby, D. H. A new combination in Cochlearia (nee Dawson, H. J. Chromosome numbers in two Limonium species Gilbert, O. L. Wild figs by the River Don, Sheffield ; Haliday, G. Crepis praemorsa (L.) Tausch, new to western Europe Horsman, F. The first record for Vaccinium uliginosum L. var. pubescens (Wormsk.) Hornem. from the British Isles Kent, D. H. The Shasta Daisy a Newton, A. Rubus stenopetalus Lef. & Muell. in Britain Preston, C. D. Potamogeton filiformis Pers. in Anglesey ... Sell, P. D. Some new combinations in the British flora tek Sell, P. D. The typfication and nomenclature of Erica bak pac Bab. Webb, C. J. New Zealand species of ee iene. naturalised in Britian and Ireland Book REVIEWS ... OBITUARIES REPORT Annual General Meeting, 6 May 1989 GraHaM, G. G. & PRIMAVESI, A. L. Notes on some Rosa taxa recorded as occurring in the British Isles Haworty, C. C. & RicHarps, A. J. The estou nae and revision of Dahlstedt’s species of Taraxacum Weber based on British or Irish plants HawortH, C. C. Six native species of Taraxacum new to the British Isles Wesster, S. D. Three natural hybrids in Ranunculus L. el Batra- chium (DC.) A. Gray SILVERSIDE, A. J. The nomenclature of some hybrids of the Scie salicifolia group naturalized in Britain ... oe Vv PAGES 81 81-82 83 82-84 84-85 85-87 87-88 89 89-90) 90-91 o2 92-93 93-95 97-109 111-114 115-117 119-124 125-130 131-138 139-146 147-151 INDEX TO WATSONIA VOLUME 18 Fo.ey, M. J. Y. An assessment of populations of Dactylorhiza traunstei- neri (Sauter) Soo in the British Isles and a comparison with others from Continental Europe Smmpson, D. A. Displacement of Eloded cdnallenSe whelik by ibd nuttallii (Planch.) H. St John in the British Isles CLARK, J. W. & Jermy, A. C. Additions to the flora of Mull and adjacent small islands Newton, A. & Porter, M. Five palit es —— Wales SHorT NOTES Bevan, J. Hieracium britannicum F. J. Hanb. in Wales Corner, R. W. M. Cardamine amara L.: its occurrence in montane bubitats in Britain David, R. W. The distribation ‘6 Gases aietopinguile Schamalchiee (C. paradoxa Willd.) in Great Britain and Ireland 3 Hofmann, H., Edmondson, J. R. & digi ia G. The Gunknes hexbar. ium of W. H. Youdale d Pearman, D. Alopecurus bulbosus Gansii in Doser Raybould, A. F., Gray, A. J., Lawrence, M. J. & Marshall, D. E The origin and taxonomy of Spartina xX neyrautii Foucaud Rechinger, K. H. Rumex acetosa L. subsp. ae Caneel Valdes: Bermejo & Castroviejo in Britain Silverside, A. J. A new hybrid binomial in Mout | es Stace, C. A. New names and combinations in the British flora ... PLANT RECORDS Book REVIEWS ... OBITUARIES oe ace ome cafe 4 Wess, D. A. BONG JEN sereceni 1990: Genera, hollow curves and boojums ny ae e - ; RumsEY, F. J. & Jury, S. L. An account ae Ofebenche in Britain ra Ireland Rumsey, F. J., SHEFFIELD, E. & HAuFLER, C. H. A re-assessment of Pteridium aquilinum (L.) Kuhn in Britain Hopson, D. D. The status of Populus nigra L. in the Rrepisbita of tretantl ETTLINGER, D. M. T. Two new varieties of British Dactylorhiza FLINK, K. E. & Hy_Mo, B. Two new species of Cotoneaster SHORT NOTES Al-Bermani, A.-K.K.A. & Stace, C. A. A new per of Festuca rubra L. Horsman, F. Abnormal aeeane in Platantita obibhah che (Ciister) Rei- chenb. and Gymnadenia conopsea (L.) R.Br. subsp. conopsea Ingram, G.I. C. & Dunster, C. W. A nonresupinate abnormality in Orchis purpurea Hudson ... Page, C. N. Herbarium sheets of the rare fern X Asplenophyllitis microdon (T. Moore) Alston in Edinburgh Rose, F. Stace, C. A. New names and combination of three hybrids A new subspecies of Gymnadenia conopsea (L.) R. Br. Book REVIEWS ... REPORT Annual General Meeting, 12 May 1990 Vi 153-172 173-177 179-187 189-198 199-200 200-201 201-204 204-206 206-207 207-209 209-210 210-212 212-213 214-228 229-237 238-246 247-255 257-295 297-301 303-305 307-309 311-313 315-316 316-317 317-319 319 319-320 320-321 323-329 331-332 INDEX TO WATSONIA VOLUME 18 Lack, A. J. & Diaz, A. The pollination of Arum maculatum L. - a historical review ae new observations SILVERSIDE, A. J. The identity of pm hous ree EK ad its nomencla- tural implications ... Cono.L.y, A. P. Polygonum lihiangense W. Smith: rejected as a natura- lized British species Dunn, A. J. Further observations on Size germanica L. RIHAN, J. R. & Gray, A. J. The distribution of hybrid Marram coe x Calammophila baltica (Fligge) Brand, in the British Isles Frost, L. C., Houston, L., Lovatr, C. M. & Becketr, A. Allium sphaerocephalon L. and introduced A. carinatum L., A. roseum L. and Nectaroscordum siculum rie pea on St Vincent’s Rocks, Avon Gorge, Bristol WILLIs, A. J., MARTIN, M. H. © Tuten. KB; Orchis purpurea Hudson in the Avon Gorge, Bristol HAZELDON, E., NArssitT, T. & RICHARDS, A. J. Differential Aauiddden efficiency within a hybrid swarm between Dactylorhiza purpurella (T. & T. A. Stephenson) So6 and D. fuchsii (Druce) Sod HorsMaNn, F. A new dactylorchid hybrid SHORT NOTES Akeroyd, J. R. Anthyllis vulneraria L. EP: eae si ee an alien kidney-vetch in Britain Bevan D. & Rich, T. C. G. Bevan’s Bittercress: the ay ies geet variant of Cardamine X fringsii Wirtgen also present in Britain Chatters, C. The status of Pulicaria vulgaris Gaertner in Britain in 1990 David, R. W. Carex X gaudiniana Guthnick —a rarity? ... Dietrich, W. The status of Oenothera cambrica Rostanski and O. novae- scotiae Gates (Onagraceae) Lang, D. C. A new variant of Ophrys apifera Fatdsori in Britain Meikle, R. D. Salix < seminigricans A. & G. Camus: a willow ae new to the British Isles Sell: PD. & Webster,.S. D. The Fatethott of Rico bachii Wirg. Stace, C. A. Three new grass combinations : Trist, P. J. O. Festuca arundinacea Schreb. var. strictior (Hack. )K. Richt Wentworth, J. E., Bailey, J. P. & Gornall, R. J. Contributions to a cytological catalogue of the British and Irish flora, 1. PLANT RECORDS Book REVIEWS ... OBITUARIES Vii 333-342 343-350 351-358 359-367 369-379 381-385 387-390 391-393 395-399 401-403 403-405 405—406 406-407 407-408 408-410 410-412 412-413 413 414-415 415-417 419-436 437-445 447-452 lad i. ats t Sis ey | w I aps? ih sullt} 2 ; nv & furs - a6 .1 ak ; bY 2193197 Sivi Dry ty 10 voted je ib ‘ot; lel davnksd be fran , MG. as J ae NZ | \s MATVNO r\ : Pao uborts at iti i (er Shae rua S irdtimnl ‘“ ) ‘- 7 Tale des - os ee ei8 Pot { chi inf b ’ U comraca at we | J ' eb Sg mM inbive vans rT * ith, iy a fo plc : ; ‘ , 4 sat wu gus ¢ sey 7 bP A rofigtoas he TB A a’ ¥ pvarilaay VN eines s ¢ | to f Li y “HANS hz Me VeV oS . i. > ao cs A .i tow a | : 7 b ~ + : 26 . Stine) looney Hi 10 Stehity oly ni are x Ya.) “) Ls ~_ AY TG bien OTs a ‘ oe ) nf ' its me) 3 et) 4 . ny : i Tatton eer . ' {yest ek BI A va +25 Ve itt hein veoh paar hoete “UE EN 6 an Roo syd batt beianoe sAdestdy teri toni: 2 aaa Pee Glace iet. istethatiog Sh Hiaiyp: Ike, ps as oie TM 4449 vob? bi 5 {SARK} ar mre Vi a oT ri beoelesdug 3 ee iy susan OG eC teown re Jeo i617 WwO tO cue. an vines idaihs Me art ae au iy | | Fart cS ' ON, Lat rat We os " . 4 ! 9 aU eo THAME HONEA i Pant : ia 7 f = ian] ALOT eS tisia bY ib its G >i) Peters cy An} igen wi no iF ait Alu I bin oe 79M bel ae 4 * . cath c e — - =< 4 i v ite ate « 4 7 ih 4 - On] =i ay MEITSR — — ; seg, a ’ ‘ 7 jan. ; ; 4 ne A : : Cort iL fl Avis: TE ie 5 at anh) cvagy 4). E540) aie) ea) janitor is Yarn 4 5 ; ne, . belies eal Meu\at * 0 wei ‘4 | Jooband .A0 q hoy a APytsben <2 yht esse, 9109 Sz airs he, a) ay Oo CME may, DoebONr t ah sil iN cok oad -G GSebnk Re ee stvnerly Liaw C205, a A3a8 ; jowy bt Uy Seared to sant aay Af 1 (iti Ay Reet rs } ' an * ta 4 pet) ebs..4 2 Page 2 i SREY wea) . vou Be WD dat aL SOESTER SB mange’ rt Heseadomeld aT CRE > ni ‘ ‘<¢ \ £ . ay ¢ 7 #2 ed » 4 9 oe oe ' — Yr aa wv a \- r 4 7 - ~ de : Ls « s L 4 = vs | -” J tt J Watsonia, 18, 7-15 (1990) a] Hedera helix L. and H. hibernica (Kirchner) Bean (Araliaceae) in the British Isles H. A. McALLISTER University of Liverpool Botanic Garden, Ness, Neston, Cheshire, L64 4AY and A. RUTHERFORD Moniaive, 19 South King Street, Helensburgh, Dunbartonshire, G84 7PU ABSTRACT Hedera helix L. sensu lato consists of diploids, H. helix L. sensu stricto, and tetraploids, H. hibernica (Kirchner) Bean, which have distinct geographical distributions and can be differentiated morphologically especially by their trichomes. INTRODUCTION The plant known as Irish Ivy, Hedera hibernica hort. or H. helix subsp. hibernica (Kirchner) McClintock, is widely used as a ground cover plant which frequently escapes from cultivation and may be found far from habitation. Its rapid growth, large and uniform leaves, hardiness, shade tolerance and lack of inclination to climb make it a landscape architect’s ideal subject for planting beneath trees and shrubs and for stabilising steep slopes. The origin of the material in cultivation has long been a matter for dispute (Lawrence & Schulze 1942) though many writers have stated that it is native in Ireland, and Mackay (1836) mentions receiving specimens from Ballybunion in Kerry. However, this ivy is rare in cultivation in Ireland and Andrews, who was the originator of the myth of ‘Irish’ ivy in Ireland (Mackay 1836), is known to have been unreliable over the labelling of his plants (M. J. P. Scannell, pers. comm.). By the second half of the last century, it was not regarded as native there (Hooker 1870; Colgan & Scully 1898). Whatever its origin, it is now very common throughout Britain (Rutherford 1979), America (Lawrence & Schulze 1942) and probably many other countries. In an attempt to discover more about the plant, A. R. originated the B.S.B.I. Network Research Project on Irish Ivy (1975-1979); see Rutherford (1979). Apart from the gross morphological characters of large, uniform, broadly lobed leaves, Irish Ivy can usually be distinguished from the common ivy by its smell. The odour of H. helix L. is very disagreeable to many people, but the scent of cut twigs or petioles of Irish Ivy is much more powerful, distinctly sweeter and more resinous. Only H. colchica K. Koch and H. algeriensis Hibb. are equally odoriferous. Jacobsen (1954) reported H. hibernica hort. to be tetraploid (2n=96) and H. helix to be diploid (2n=48). He used cultivated material of H. hibernica but believed it to represent a wild species. CYTOLOGICAL METHODS The method used was a modification of Dyer’s lacto-propionic orcein technique (Dyer 1963) in which the root tips were squashed in the undiluted saturated solution of orcein in 1:1 lactic acid:propionic acid. Rapidly growing root tips were placed in vials of a saturated solution of a- monobromonaphthalene in tap water for about 4 h at room temperature. The root tips were then H. A. MCALLISTER AND A. RUTHERFORD "Sa|s] YSHUg 94) Ul URog (JoUYyoITy) voIwsaqiy "YH (96=UZ) plojdenay (q) pue “] xyay vsapay] (Sp=UZ) plojdip (e) Jo syunod Jo uoNNquisiq ‘| AUNDIy HEDERA IN THE BRITISH ISLES 9 fixed in 1:3 methanoic (acetic) acid:ethanol (98%) for at least 24 h, before being hydrolysed in 1M hydrochloric acid at 60°C for about 5 minutes in a water bath, and then transferred to 70% ethanol for storage. A root tip was then examined under a dissecting microscope, placed in a drop of stain, macerated with needles, tapped and squashed. The squash preparations were examined for cells containing well separated chromosomes and, when such were found, the chromosomes were counted. With clear preparations, counts of precisely 2n=48 or 2n=96 were obtained, but on poorer preparations, where only approximate counts (45-48 or 90-96) could be made, the results were recorded as 2n=c.48 or 2n=c.96. CYTOLOGICAL RESULTS It was soon confirmed that the cultivated Irish Ivy, H. hibernica hort., and some other cultivars were tetraploid, as well as much of the wild material from all along the western seaboard of Britain and Ireland, western France and Spain. In the wild, tetraploids seem to occur to the exclusion of diploids in southern Ireland, the Isle of Man, western Wales, the West Country eastwards to the Isle of Wight and the New Forest, the Channel Isles, western France, northern Spain and probably most of that country except the Sierra Nevada region where diploids occur, and in the Algeciras—Gibraltar area and in Portugal, near Lisbon, where hexaploids (2n=144) have been found. Diploids and tetraploids were found together along the eastern boundary of the area of distribution of the tetraploids, but further east only diploids were found, apart from the occasional naturalised tetraploid H. hibernica hort. (Fig. 1). The diploid thus appears to be the native ivy north and east of the tetraploid’s area in Britain. Diploids have been counted from eastern France, Denmark, Switzerland, Italy, Yugoslavia, Turkey, the Crimea and the Caucasus. Thus the tetraploid seems to occur to the exclusion of the diploid on the Atlantic seaboard of Europe, from Gibraltar to southern Scotland, and the diploid to the exclusion of the tetraploid north and east of these areas, extending eastwards to the Crimea and the Caucasus. It would appear that the diploid only reaches the Atlantic in central and northern Scotland. The one notable exception to this is the isolated occurrence in Sicily of tetraploids which are morphologically like diploid H. helix. As the diploids and tetraploids have distinct, barely overlapping, geographical distributions, it may be said that they are behaving as distinct ‘biological’ species, with different climatic preferences, so the two cytotypes are probably in competition with one another where they meet. No hybrids (triploids, 2n=72) have yet been found despite many cytological determinations on material from areas where the two species meet. MORPHOLOGY As both the wild diploids and tetraploids have clearly all been previously referred to H. helix, it was considered that any character which might distinguish the two cytotypes was likely to be fairly cryptic. H. hibernica hort. was thought to be perhaps an extreme variant of the tetraploid, so it was carefully compared with a typical diploid H. helix. As the trichomes on the young leaves give useful diagnostic characters for distinguishing other species of Hedera, those on H. helix and H. hibernica hort. were examined in detail. These trichomes always consist of a short stalk bearing a central boss from which radiate a variable number of unicellular rays. In the diploid some of the rays of most of the trichomes stand up at an angle to the leaf surface giving a bristling appearance (Fig. 2a). By contrast, in H. hibernica hort. these rays almost always lie flat along the surface of the leaf, all in one plane (Fig. 2b). It was soon evident that all the tetraploids, excepting those from Sicily, had trichomes of the H. hibernica hort. type, while all the diploids had trichomes of the type found in the typical diploid examined. It therefore seemed clear that H. hibernica hort. was a distinctive clone or variety of a widespread tetraploid taxon, most specimens of which would previously have been classified as H. helix. Although very obvious in young leaves on rapidly growing shoots, the trichome character is by no 10 H. A. MCALLISTER AND A. RUTHERFORD : Ler} Feary A) Wey ~ 4 RE Tc > . “oa, Pe) a ah LA ag (0 ner AG ? ayaa « FiGURE 2. Drawings from stereoscan electron micrographs of the trichomes of (a) Hedera helix L., (b) H. hibernica (Kirchner) Bean. HEDERA IN THE BRITISH ISLES 11 means easy to use. Very young leaves of the diploid seem to be covered in fine grey-white velvet, the effect being created by the denseiy packed trichomes. On the tetraploid comparable young leaves seem much less hairy, since few of the trichomes stand out at an angle from the surface of the leaf. However, the best leaves to examine for determination of this character are slightly further back on the shoot, where the lamina expansion has separated the trichomes so that individual trichomes can be clearly distinguished. Observations in a good light with a X20 hand lens are usually adequate, and with good, young, vigorous material held up to the light the cytotypes can often be distinguished with the naked eye — but such observations should always be checked with a lens. It is best to examine trichomes on the underside of the leaves, but those on the upper surface and on young leaves may also be examined. Trichomes on the upper surface and trichomes with rays standing out from the surface tend to be lost first as these are most likely to be abraded, so surfaces presenting few trichomes may give unreliable determinations. The last sites to lose their trichomes are the interveinal areas between the main veins, where they adjoin the petiole, on the underside of the leaf. Trichomes on slightly older stems of both cytotypes tend to have all their rays appressed to the stem, and some trichomes on the veins of the tetraploid have rays standing out from the leaf surface, so trichomes on stems and veins are best disregarded. Furthermore, in winter, in exposed positions, on slow-growing or flowering shoots, the trichomes of the diploids with their projecting rays may become dislodged or be flattened on to the leaf surface, and such material may be unidentifiable unless taken into cultivation and fresh young leaves examined. It can be seen, therefore, that on the trichome character, the diploid is much more likely to be misidentified as the tetraploid than vice versa. Used with care, examination of the trichomes is a reliable means of distinguishing between the Common (diploid) and the Atlantic (tetraploid) Ivies, but other characteristics are also more or less correlated with the chromosome number (Table 1). The leaf shape is very variable in both. Many ‘races’ of each pass through many changes between the creeping, sterile phase and the flowering, mature phase. Some tetraploids scarcely alter, having almost flowering-type foliage at all stages, and a variant of this has long been known from the Channel Isles (Druce 1912). Both cytotypes usually produce the familiar ivy-shaped leaves at an early stage in the plant’s development. Later, before the mature phase is reached, both have, usually, a pedate-leaved phase in which the lobes are attenuate and the central much longer than the side lobes, and towards the flowering branches these lateral lobes diminish and entire leaves are produced. ECOLOGY In moist, sheltered sites the tetraploid is capable of dominating the woodland floor, roadside banks and cliffs in a manner of which the diploid seems incapable, even in the most favourable conditions. As early as 1872, Hibberd remarked on ivies with giant leaves in the Vale of Conwy and we have noted populations on Porkellis Moor, Cornwall, near Blackgang on the Isle of Wight, West Symonds Yat in Avon and even in woods by the mouth of the River Stinchar in Ayrshire, near the northern limit of the natural distribution of the tetraploid. The ability of the tetraploid to produce larger leaves on longer petioles, and to have longer internodes, apparently allows it to invade habitats unavailable to the other cytotype. Shingle banks just above high water level of spring tides are often colonised by the tetraploid, but the diploid has never been found in such habitats. The vigour of the tetraploid is clearly seen among the sea-cliffs at Kennedy’s Pass and at Pinbain Burn-foot, Ayrshire, where it is the dominant plant. Ivy grows to a considerable height up the cliffs, covers the seaside boulders and creeps through dry sunny roadside verges where it is not discouraged by being mown. Where not cut, this ivy competes successfully with Pteridium aquilinum, Dryopteris filix-mas, Mercurialis perennis, Filipendula ulmaria, Poa pratensis, Rumex crispus and Rubus latifolius, among other tall-growing species. It flourishes in two habitats at the former site which would probably have proved too extreme for the diploid, these being a dry, sunny exposure in very well-drained soil and a marshy ditch. In areas where both cytotypes occur, especially in Galloway in south-western Scotland (A. J. Silverside, pers. comm.), the tetraploid is mostly found in the milder, coastal regions while the diploid tends to be the ivy of the inland woods. 12 H. A. MCALLISTER AND A. RUTHERFORD TABLE 1. ACOMPARISON OF THE CHARACTERISTICS OF HEDERA HIBERNICA AND H. HELIX Character Growth Leaf size Pedate leaves Leaf texture Leaf colour Hyaline layer Leaf sinuses Leaf veins Trichomes Sap H. hibernica (Kirchner) Bean Fast and vigorous, often has long inter- nodes, up to 18 cm or more. New shoots may be thick and succulent. Leaves variable both at different phases and among different plants but overall considerably larger, up to 10 cm wide by 10 cm long in moist sheltered localities. The leaves at the pedate phase frequently the largest and almost equal in surface area to those at sub-fertile and fertile phases. The tip of the longer central lobe usually rounded. Leaves usually coriaceous, frequently fleshy-waxy. In cold the interveinal areas of leaves may turn pinkish or light bronze. The veins develop apple-green borders. Some forms with low anthocyanin content only have red margins or do not darken at all, rarely the leaf becomes red-maroon. The hyaline layer is more readily apparent and may appear thicker due to the up- turned margins. The sinuses are mostly deep and often strongly arched; leaves may be funnel- shaped. The veins at the sterile phases rarely raised on the leaf surface and seldom differ in colour from the ground, usually bordered in a paler shade or yellow- green. Trichomes appear sparse due to the rays lying parallel to the leaf surface; they are often tinted fawn or more rarely orange- brown. Sap odour strong, often pine-like and sweet. TAXONOMY H. helix L. Markedly less vigorous and _ usually slower; internodes often quite short. New growth often wiry. Leaves variable at different phases and among different plants, but generally markedly smaller at all phases. The leaves at the pedate phase mostly produced near the ground and on the whole plant almost always the smallest; the tips of the longest lobes acute. Leaves leathery, often of papery texture, but thin in coastal areas. In cold, particularly in frosty places, the leaves may turn purple-black all over; the ground colour darkens considerably. The hyaline layer is not readily visible, due to the down-turned margins. The sinuses may be deep but the blade is usually flat, apart from the margins. The veins at the sterile phases almost always raised on the leaf surface like fine wires, often silver-white, rarely edged with paler colour. Trichomes appear as grey-white pubes- cence on new tips, due to the rays project- ing at various angles and becoming enmeshed. Sap odour much weaker and usually rather acrid. The original aim of the investigation was to elucidate the status of H. hibernica hort. When tetraploids were found to be widespread in the wild, the question arose as to what was the relationship between the morphologically very distinct tetraploid, H. hibernica hort., and the wild tetraploids which previously would have been included within H. helix. The agreement in trichome characters between H. hibernica hort. and the wild tetraploids led us to believe that the cultivated plant might be a clone or group of clones of the wild tetraploid. However, the so-called Irish Ivy was unusual in its uniformly large leaf size and non-climbing habit, contrasting with the variable leaf size and normal climbing of most of the wild tetraploids. On the Isle of Wight wild tetraploids were discovered producing some leaves identical to those of the cultivated plant, and in Herefordshire at HEDERA IN THE BRITISH ISLES 13 Symonds Yat, colonies of wild ivies were found almost a match to H. hibernica hort. The Isle of Wight plants bear the Irish Ivy leaves on shoots in transition from the vegetative to the flowering (mature) state. Most of the wild tetraploids produce leaves of this type, but the Isle of Wight samples were most pronounced. It therefore seems that the cultivated Irish Ivy is a clone, or group of similar clones, stabilised in the semi-arboreal condition. It resembles the creeping/juvenile phase in having pliant stems bearing some of the specialised attaching roots, but these are less numerous than normal in juvenile ivies. The stems run along the ground, sometimes climbing walls and more rarely trees, and cuttings root very readily. The chief similarities to the arboreal phase are in the reduction of the ability to cling and thus to climb, due to fewer adhesive roots, and the broader leaves with shorter, wider lobes. When it does climb it can mature into the flowering condition with radially symmetrical shoots of brittle wood, totally lacking rootlets and having entire leaves. Seedlings are very like the parent, but do seem more ready to climb, and, like it, have rather larger and broader leaves than is typical for the younger stages of wild tetraploids. Similar observations have been made in Germany (G. Griiber, pers. comm.). H. hibernica hort. therefore probably consists of a clone or clones from the south of England. Horticulturalists have long been aware of slight variation in H. hibernica hort. Some clones are darker in the foliage or are not glossy, and another clone has more sharply lobed leaves than normal. In the main, however, it is uniform. Variegated sports occur, including ‘Variegata’, ‘Sulphurea’, ‘Marmorata’ and ‘Rona’. Also, seedlings may arise by self-fertilisation. The present situation is that there are diploids and tetraploids normally referred to H. helix, with a tetraploid cultivar being called H. hibernica hort. or H. helix subsp. hibernica, a name that has become current in horticulture, and that has recently been validated nomenclaturally (McClintock 1987). At what taxonomic level should they be recognised? As they differ in ploidy level, geographic distribution, distinctly in one morphological, albeit cryptic, character, and in other less obvious ways, we consider it desirable to distinguish them as separate species. We are aware that this decision may be controversial as many would prefer to recognise them as subspecies (cf. Lum & Maze 1989). If Hedera were an obscure genus known to few we would perhaps have concurred. However, it is very well-known to botanists and gardeners and there are numerous cultivars, so that the use of subspecies would lead to long clumsy horticultural epithets and possibly to the ignoring of distinction between the subspecies. The level of subspecies is well suited to situations where geographical or ecological variants exist at the same ploidy level, as with Betula pubescens Ehrh. (Walters 1964), or where cytotypes are not easily or always distinguishable, as in Asplenium trichomanes L. (Crabbe et al. 1964) or Galium palustre L. (Clapham 1962). However Ehrendorfer & Puff (1976) treat G. elongatum C. Presl as a separate species, despite the admission that intermediates occur with G. palustre L. We consider that the situation with British ivies is more like that found within the Dryopteris dilatata complex, in which northern diploids have been distinguished as D. expansa (C. Presl) Fras.-Jenk. & Jermy (D. assimilis S. Walker) and are differentiated from the tetraploid, D. dilatata (Hoffm.) A. Gray, primarily by a cryptic spore character, but also by a not very easily defined morphology. Linnaeus’ specimens consist of at least four samples of Hedera helix, three of which are too mature for certain determination and for this reason specimen 280.2 in LINN, which has a sterile shoot and typical diploid trichomes, is here selected as the lectotype. Thus H. helix may be retained for the diploids. The earliest name for the tetraploid is probably H. hibernica hortul. (De Candolle 1830), a nomen nudum as there is no description. The first mention with a description is by Kirchner in Petzold & Kirchner (1864) as “‘H. helix var. hibernica hort. Schottischer Epheu’’. In 1914 Bean raised this taxon to specific rank as H. hibernica (Kirchner) Bean. Thus H. hibernica is the name which now must be used for the tetraploids (McAllister 1981), the cultivated clone becoming H. hibernica (Kirchner) Bean cv. Hibernica. Most ivy cultivars are referable to H. helix, but some others, as well as cv. Hibernica, belong to H. hibernica, including ‘Digitata’, ‘Deltoidea’, ‘Variegata’, ‘Maculata’, ‘Albany’, ‘Hamilton’, and ‘Helford River’ (Lawrence & Schulze 1942; Rose 1980, 1983; Griiber 1983; Sulgrove 1981a & b). The semi-adult nature of H. hibernica hott. is paralleled in H. helix by a remarkable clone collected at South Glendale in the south-west of South Uist in the Outer Hebrides. This looks superficially like cv. Hibernica but has leaves from 6 cm wide by 5 cm long to 12-5 cm wide by 9 cm long; the texture is thinner than that of the Irish Ivy however, and the margins have the typical undulations of maturing juvenile leaves of H. helix. 14 H. A. MCALLISTER AND A. RUTHERFORD SYNONYMY The synonymy of most ivies in cultivation and their cultivars is given in Lawrence & Schulze (1942), but is summarised here with H. hibernica treated as a distinct species. 1. H. helix L., Sp. Pl. 202 (1753). Lectorype: sheet 280.2 (LINN), designated here. Common Ivy. 2. H. hibernica (Kirchner) Bean, Trees & shrubs hardy in the Br. Isl. 609 (1914). Atlantic Ivy. H. helix var. hibernica Kirchner in Petzold & Kirchner, Arb. Muscav. 419 (1864). H. hibernica Carr. in Rev. Hort. 71:163 (1890), nomen nudum. H. helix subsp. hibernica (Kirchner) D. McClintock, Suppl. wild fl. Guernsey 23 (1987). We propose the common name Atlantic Ivy for H. hibernica in the wild state and suggest restricting the use of Irish Ivy to the cultivar Hibernica. PHYTOGEOGRAPHY The distribution of H. hibernica, the Atlantic Ivy, is very similar to that of certain heaths, Erica cinerea having a slightly more extensive range, while E. vagans and E. mackaiana have progressively more limited ranges, and E. ciliaris extends further south into Morocco. It is interesting that although these heathers belong to totally different types of community from the ivy, all are evergreen and so subject to the same freezing and desiccation stresses in cold winters which presumably limit their eastward spread. They, together with the more widely distributed H. helix and Ilex aquifolium, are among the very few larger evergreen flowering plants native to northern Europe. The distribution of the two species in the British Isles shows H. helix with an almost continuous distribution and H. hibernica in somewhat separate areas on the western coasts. This pattern suggests that H. hibernica formerly had a more extensive, continuous range which has been reduced by H. helix invading from the south-east. In milder, wetter parts of the country it seems that H. hibernica can occupy the available habitats and hold off the advance of H. helix. In less mild areas the occasional exceptional cold spell may do sufficient damage to H. hibernica to leave niches free for colonisation by the more winter-hardy H. helix. H. hibernica occurs in the south-west of England, most of southern and western Wales and the northern Welsh coast. There is an isolated record for Silverdale in northern Lancashire and extensive isolated populations occur in the mild coastal parts of Galloway in south-western Scotland. Only H. hibernica has been found on the Isle of Man, and the distribution of the two species in Ireland might suggest that H. helix has only recently invaded from Scotland, from the Mull of Kintyre, and may be extending its range. The situation on the main island of Great Britain could be expected to have more or less stabilised for present climatic conditions, but it will be interesting to watch what happens following a succession of cold winters. ORIGIN OF H. HIBERNICA H. hibernica is here classified with H. helix and H. azorica Carriere (H. canariensis ‘Azorica’ hort.) because it has the same long, white-rayed trichomes. It differs from these diploid species in that its rays lie more or less appressed to the leaf surface, in a similar way to the rays of the other much larger grouping of ivy species, the orange-red, small-haired species of Hedera, found mostly to the south and east of the Mediterranean and Black Seas (McAllister 1981). There is the possibility therefore that the tetraploid H. hibernica could be an allotetraploid derivative of H. helix and a species from the second group. In North Africa, in the Atlas Mountains, there is an as yet undescribed diploid with large five-lobed ivy-shaped leaves and considerable vigour which seems an ideal candidate. H. hibernica resembles this species in its capacity to produce larger leaves than those of H. helix, in the greater average number of rays per trichome and the fawn to slightly orange colouration of the central boss of the trichomes of some populations, as well as the attitude of the HEDERA IN THE BRITISH ISLES I) rays, the sweeter sap odour found in the larger group of ivies and the more luxuriant growth. H. hibernica is thus intermediate in several characteristics between H. helix and the undescribed species from Morocco. ACKNOWLEDGMENTS The authors would like to thank the Librarians of the Royal Botanic Gardens, Kew and the Royal Botanic Garden, Edinburgh, for their assistance in researching references; K. Veltcamp for taking micro-photographs; C. D. Preston of the Biological Records Centre for help with records and making the maps; D. McClintock and H. Heine for originally encouraging the study of H. hibernica hort.; J. K. Hulme for providing accommodation for many ivies in the Botanic Gardens at Ness; C. E. Jarvis and J. Fiddian-Green for assistance with the choice of a lectotype; and all individuals who obtained Hedera samples for us. REFERENCES Bean, W. J. (1914). Trees and shrubs hardy in the British Isles 1: 609. London. CLAPHAM, A. R. (1962). Galium L., in CLAPHAM, A. R., TuTin, T. G. & WaRBuRG, E. F. Flora of the British Isles, 2nd ed., pp. 782-783. Cambridge. CoLGan, N. & ScuLLy, R. W. (1898). Contributions towards a Cybele Hibernica, 2nd ed., p. 160. Dublin. CrabsE, J. A., JERMy, A. C. & Lovis, J. D. (1964). Asplenium L., in TuTin, T. G. et al., eds. Flora Europaea 1: 14-17. Cambridge. De CANDOLLE, A. P. (1830). Hedera. Prodr. Syst. Nat. Reg. Veg. 4: 261. Paris. Druce, G. C. (1912). Hedera helix L. var. sarniensis Druce. Rep. botl Soc. Exch. Club Br. Isl. 3: 163. Dyer, A. F. (1963). The use of lacto-propionic orcein in rapid squash methods for chromosome preparations. Stain Technol. 38: 85-90. EHRENDORFER, F. & Purr, C. H. (1976). Galium sect. Aparinoides, in TutTin, T. G. etal., eds. Flora Europaea 4: 21. Cambridge. GRUBER, G. (1983). Merkmale Anbau und Verwendung Winterhalter Hedera Arten und Sorten, pp. 75-87. Geisenheim. HIBBERD, S. (1872). The Ivy: a monograph, p. 27. London. Hooker, J. D. (1870). Students’ Flora of the British Islands, p. 172. London. JACOBSEN, P. (1954). Chromosome numbers in the genus Hedera L. Hereditas 40: 252-254. LAWRENCE, G. H. M. & ScuuwZE, A. E. (1942). The cultivated Hederas. Gentes Herb. 3: 142-144. Lum, C. & MAzeE, J. (1989). A multivariate analysis of the trichomes of Hedera L. Watsonia 17: 409-418. MCcALLIsTER, H. A. (1981). New work on ivies. Int. Dendr. Soc. Year Book 1981: 106-109. Mackay, J. T. (1836). Flora Hibernica, p. 135. Dublin. McC uintock, D. (1987). Supplement to the wild flowers of Guernsey, p. 23. St. Peter Port. PETZOLD, E. & KIRCHNER, G. (1864). Arboretum Muscaviense. Berlin. Rose, P. Q. (1980). Ivies, pp. 114-118, 147-148. Poole. Rose, P. Q. (1983). ‘Hibernica’. The Ivy, J. Br. Ivy Soc. 6(1): 34-35. RUTHERFORD, A. (1979). The B.S.B.I. Irish Ivy survey. B.S.B.I. News 22: 8-9. SULGROVE, S. M. (1981a). The variegated Hibernicas. The Ivy Bull. 7(4): 29-34. SULGROVE, S. M. (1981b). Nomenclatural analysis of the variegated Hibernicas. The Ivy Bull. 7(4): 35-39. WALTERS, S. M. (1964). Betula L., in Tutin, T. G. et al., eds. Flora Europaea 1: 57-58. Cambridge. (Accepted May 1989) ~ bndirses i ° . ty 3: nite any eens oy i bm mehy a bow at Bi nen Ki hyoompmny of most vied inl dull gt ty seemed od here wa i, Ne. . | 5 3 f - actieiet L A sa pia. stab: : IN - ne os ; i ; tid A Vang hed ad, a digs i the’ y i Vf e ~ e i. i wit ¢ v pa rat j aid -, biay: Percy bleed pod it Ae ay mpi J Bey Ceres SEO ett Pies r S 4 re) 217; OM sry aff Sina fat Tt ee tea + ie rs te viie Sar aiien Te mGshy licen We ‘antes > pee} ie enobust) on wo oly ab aoivi-¢oemten 40} mE abomen noe gnihj rong) w a € u FAT ery rit As cay 3! Phe Ny we rich ATT hy WY : roti send Yorad f94 HE po o— ' f ) ae C uit ive , iA a. i 2e-ul cies tS \ 1% ee , a Be aoe HeTOMETeting ae PL . - (ie ; : 1 , s@ wn } hy ' tk “4 Q au. (AAs -" a ey ainda ece ot, vos , ve x ua . syst i Ms ai i 4 ’ ‘ é J . SV <3 ti) % ‘ ‘in ne fe HS ni be) a ig Bate he tes wn as ‘Wal St f } dene: itn? C1) Gia siaclqotty ral? ‘ ) 4 +¢é he a aA. 4 " watt | = alga - 7 4 p tk ye er t? } , Vie sin oA ahd 5] a _ 4 Vit aw tarts dis ; . - » Te y a "te t 49> fer a tated Ted: . 4 } sy th af 4% de, 5a ROMO DL USF ROWS iE 0 eck 7 . ‘ arnt aN parent Ett dy. Mi . 7) Atria Sf ity vay ~) LATA) ij J + . bd iM ie we ss wk \ si y RTT, RLU AR OCU) PT PLC reeesal eulo Y » ‘ a: \ 33 A at «4 Pei} haa ay A ri 7 % ce lee | | TOME yee a Maal yc [ptt SNe: , at Pie SEC TAT SV oe Lo OP . a - . a - : ~ 4% % t ’ See AP Sra ee }aCy Ws ‘3 | MoATORe are far Btitt pis * 7 we ~ noyneoalk Sone aR. > rma X Bag f % i Ohi Si t&) ay c » ’ : : ae b ba Ry nile d GO aye y nh ‘pH odlet 48 te Ae : : 7 ar Ee youre wi dad LS oT Ae) A ne bE 26 ofp )0 Whit wrk aX] Wither lta eo y ren oft felSel) .M 2 aaa \ iA panic! f orgy ob op etaylein bere aioe ary 1) ue © ol / aT . [ ; . i : Le a nvis j aie wiles \ fs € id lo i.e ree eo ae ag arrere | enomiens ALC C he rif ’ , - bt "1 nar & fects td “et plod specie ~~ i é a - ” 5 “at Seivtig i , ~y - ay m iy . ODE 4 « ‘ - ‘if i. wrtall-aiee) peciag! Meaieru, Mand seam aie : - vit vi Te re cy ott fs. ’ 4 a (yor ner cl cave “ schon ne a Wd prigcobe J ) “‘S ct SOMIW TF yy hvtseun au nen ae o) a Ue Watsonia, 18, 17-20 (1990) 17 Detection of protandry and protogyny in Sycamore (Acer pseudoplatanus L.) from infructescences P. BINGGELI Department of Biological & Biomedical Sciences, University of Ulster, Coleraine, Co. Derry, BT52 1SA, Northern Ireland ABSTRACT In Sycamore (Acer pseudoplatanus L.), trees may be protandrous when their inflorescences start with a sequence of male flowers followed by a sequence of female flowers, or protogynous when the reverse sequence occurs. The sex expression on an inflorescence may change more than once and, in Ireland, five modes of sex expression have been observed. When studying such heterodichogamous species it is essential to determine their sexual morphs. A method to sex Sycamore using infructescences only is described. Using a variety of characters such as fruit size, percentage parthenocarpy, infructescence length, size of terminal bud scar and position of the fruit on the first secondary axis, it is possible to distinguish between protandrous and protogynous individuals. INTRODUCTION In Sycamore, Acer pseudoplatanus L., all flowers are functionally unisexual and appear sequentially on a single inflorescence. When male anthesis takes place before the stigmas become receptive, the inflorescence is described as protandrous, and is protogynous when the reverse sequence occurs. Likewise individual trees may be either protandrous or protogynous. Such a sexual dimorphism is called heterodichogamy (Stout 1928). On a single inflorescence the sex of sequentially opening flowers may differ more than once in time, and de Jong (1976) described eleven different modes of sex expression within an inflorescence, of which five have been observed in the north of Ireland (Fig. 1). Protogynous individuals will produce inflorescences of Mode B and very rarely a few of Mode G. Protandrous individuals are far more variable as they have inflorescences of Mode C, D, or E, ora mixture of these. Male-flowering trees are described as protandrous because in some years some or even all their inflorescences have female flowers. Similarly protandrous individuals will exhibit large annual variation in the proportion of inflorescences of Mode C, D, and/or E (de Jong 1976). Scholz (1960) has reported the existence of female-flowering individuals. There is no evidence that trees will change their modes of sex expression with age. In heterodichogamous species certain characters, such as fruit production, percentage of fertilized fruits and the number of carpels per fruit, may vary between morphs (Binggeli, unpublished). In order to explain such variations it is then essential to know the sexual morph of the trees studied. Because sexing the flowers of trees in spring is rather time-consuming and/or impractical on tall specimens, a method using the morphological characters of the infructescence was developed. RELIABILITY OF METHOD The method for the identification of the sex expression of the inflorescences using infructescences was developed in 1983, and was tested with 95% success when comparing the flowering data of 240 trees obtained in spring and the determination of the sex expression using infructescences in the autumn. The test was repeated in 1984 with 100% success. 18 P. BINGGELI Flowering sequence Modes co} re} cS O11 mo ©) W G Figure 1. Modes of sex expression observed on Sycamore inflorescences in the north of Ireland. Flowering sequences are represented by male (’) and female (2) type flowers. A roman numeral subscript one (I) represents an initial flowering period, a roman numeral subscript two (II) a subsequent flowering of the same type. Modes according to de Jong (1976). DESCRIPTION Using fruiting material only, one can differentiate between protandrous and protogynous individu- als and also between infructescences of Mode B and Mode G of the latter group using the characters listed in Table 1 and as shown diagrammatically in Fig. 2. In Mode G the structure and position of the fruits of the first part of the stalk is similar to the normal protogynous infructescence (Mode B), but it is longer and it bears a few small parthenocarpic fruits at its end (Fig. 2C). It is, however, impossible to distinguish between infructescences from Mode C and D. The very few male- flowering individuals (Mode E; less than 1% in Ireland) will not be recognised with this method and only the shoot morphology will provide indications of their existence. In these, soon after flowering, TABLE 1. MORPHOLOGICAL DATA FROM INFRUCTESCENCES DIFFERENTIATING BETWEEN PROTANDROUS AND PROTOGYNOUS INDIVIDUALS, AND ALSO BETWEEN MODE B AND MODE G OF THE LATTER GROUP Protogynous Character Protandrous Mode B Mode G Fruit size (length of samara in cm) 2-5—3°5 3-0+-4-0 3-0-4-0 & (1-:5-2°5 terminal) Parthenocarpy* 30-90% 10-40% 10-40% Length of infructescence >10 cm <10 cm >10 cm Terminal scar (diameter in mm) 0-5-1-0 1-0-2-0 0-5 Usually diagonal Position of fruit on 1st secondary axis On a tertiary axis Terminal Terminal (2nd axis ends with a scar) *Parthenocarpy is assessed as the percentage of empty samaras. In the field they are recognized by their smaller size and also by their distinctly empty nutlet. HETERODICHOGAMY IN SYCAMORE 19 @ ©, FLOWERS Q oO, FLOWERS ¢ ©, FLOWER BUDS Oo OO FLOWER BUDS © Ox FLOWER BUDS Ficure 2. Architectural differences between protandrous and protogynous inflorescences and infructescences (see Table 1 for other diagnostic features) and shoot morphology after leaf and infructescence fall in Sycamore. A, Protogynous inflorescence (21; flowers (O) of Mode H are Oy in Mode B (O)). B, Protogynous infructescence, Mode B. C, Protogynous infructescence, Mode H. D, Protandrous inflorescence. E, Protan- drous infructescence. F, Vegetative shoot. G, Flowering shoot (Mode E). H, Fruiting shoot (Flowering Modes By, Cy Dede G): 20 P. BINGGELI the inflorescences will fall and the two growing terminal buds will be closely appressed (Fig. 2G). On the other hand in other modes of flowering, female flowers, even if unfertilized, will produce fruits, because of a high parthenocarpic tendency in maples. Such infructescences will remain on the trees most of the summer leading to two well separated terminal buds (Fig. 2H). It should be noted that some small flowering side shoots may not produce any buds. Cnly practice allows one to determine the sex expression of the individual with accuracy from infructescence material, and whilst the majority of the individuals examined fit easily into one or other of the two morphs, nevertheless some trees have features which do not always fit entirely the description given in Table 1 and Fig. 2. For instance some infructescences of Mode B do have a terminal fruit, but this is never the case for protandrous modes of flowering. The size of fruits and infructescences, and the number of fruits per infructescence given in Table 1 are applicable to Sycamores encountered in most of the British Isles and the Alps. However in areas with a very favourable climate (e.g. some parts of lowland Switzerland) measurements of fruit and infructescence size and the number of fruits per infructescence may be higher, and therefore the values listed in Table 1 may be misleading. ACKNOWLEDGMENT Brian Rushton’s comments on the manuscript and corrections were much appreciated. REFERENCES JonG, P. C. DE (1976). Flowering and sex expression in Acer L. Meded. LandbHoogesch. Wageningen 76: 1-201. ScHoLz, E. (1960). Bliitenmorphologische and biologische Untersuchungen bei Acer platanoides L. und Acer pseudoplatanus L. Ztichter 30: 11-16. Stout, A. B. (1928). Dichogamy in flowering plants. Bull. Torrey bot. Club 55: 141-153. (Accepted May 1989) Watsonia, 18, 21-31 (1990) yA The Rubus flora of the Isle of Wight D, E.. ALLEN Lesney Cottage, Middle Road, Winchester, Hants., SO22 5EJ ABSTRACT Fifty-nine named microspecies of brambles (Rubus fruticosus L. agg.) are reliably recorded from the Isle of Wight (v.c. 10). Four of these are of alien status. Separated from Hampshire by a strait nowhere wider than 8 km, the island nonetheless diverges markedly from that county in its Rubus flora. Five species with British Isles ranges that are strongly western are known in Wight but not in Hampshire; six other species occur in the island relatively more plentifully. More strikingly, Wight lacks three of Hampshire’s commonest and most generally distributed species, a fourth having arrived in its sole locality apparently only recently. Conspicuous absentees are also to be found in the groups of species the island shares respectively with the Bournemouth and Southampton districts. A slighter affinity with Sussex is seemingly the product of a late infiltration. INTRODUCTION The Isle of Wight (v.c. 10) can be expected to be peculiarly rewarding to the student of the Rubus fruticosus L. aggregate. Its 381 km? are patently a broken-off fragment of Hampshire (of which up to 1890 it formed part administratively) and in that county this group attains probably its greatest diversity in the British Isles, on account of its southerly position and the overlap there of the rich Rubus florulas of the south-east and the south-west of England. How far the more widespread brambles of Hampshire are represented in Wight, and vice versa, thus constitutes a valuable test of the extent to which a comparatively narrow stretch of water (nowhere wider than 8 km) acts as a barrier to dispersal in this genus. This water barrier is known to have been in place since early in the Flandrian, for marine deposits on both sides of the Solent, dated on the basis of the pollen content of overlying peats, indicate that the present coastline was attained at least by 7,000—8 000 B.P. (Devoy 1987). Apart from a perhaps rather limited number of Rubus species that were present at the time of the severance, therefore, and the four which give evidence of having been introduced recently through human agency, the majority of the brambles to be found in the Isle of Wight today must either have had their origin within the island, as the products of local crosses, or been brought in over the centuries by berry-eating birds (not necessarily, of course, only from England). The period since the severance is too short for any significant degree of endemism to have developed in the island’s ‘macro’ flora and fauna. Nor are the ‘macro’ flora and fauna sufficiently different in their composition from those of Hampshire to afford much scope for biogeographical comparison. [t is on to microspecies, consequently, that one is forced to fall back to find ample indications of divergence; and Rubus far surpasses all other lowland groups of these in the richness of its diversity combined on the whole with freedom from any suspicion of introduction through human agency. Comparative study is rendered far from straightforward, however, by major ways in which the two counties differ physically, most notably in their geology — apart, that is, from the obvious facts that Wight is an island, only one-tenth as large and crucially that much further south. Hampshire, broadly speaking, is a belt of chalk sandwiched between two belts of gravel and thus for the most part hospitable to this mainly calcifuge group of plants. By contrast, fully half of Wight consists of clay and chalk and only the Lower Greensand which occupies the southern third is out-and-out ‘Rubus country’. And even that smaller amount of potentially suitable terrain is deceptive, for most of the natural woodland the island formerly possessed has been lost and the few considerable expanses of acid ground which were still surviving until the late 19th century have either greatly 22 D. E. ALLEN diminished or disappeared altogether. Away from the south-eastern corner, productive localities tend now to be isolated ‘oases’; and there would be fewer even of those were it not for the fact that several of the chalk downs have a cap of gravel, and harbour as a consequence plateau heaths which in certain cases are quite extensive and sufficiently out of the way to have escaped elimination by modern farming pressures. HISTORY OF RUBUS INVESTIGATION IN WIGHT Fortuitously, Wight was the scene of one of the very earliest investigations of Rubus in Britain. This was in the 1840s, at the hands of Thomas Bell Salter, a physician who settled in Ryde in 1839 and brought with him an interest in the group already kindled on the Dorset heaths around his boyhood home at Poole (Salter 1839). Unfortunately, in common with all other specialists in the genus at that period, he interpreted the species far too broadly and his published records (Salter 1845, 1848, 1856) are consequently of all too limited use. On the other hand numerous specimens collected by him survive, mainly in BM and E, with a few also in CGE, SLBI and W. Borrer’s herbarium (K). Most of these have proved to be redeterminable, though regrettably they do not include some of the entities which he over-optimistically identified with species figured and described by Weihe & Nees (1822- 27). C. C. Babington held Salter’s work in high regard and twice paid him visits, on the second occasion, in September 1846, collecting with him in the field (Babington 1897). He was particularly impressed by an eglandular bramble with large white flowers, which Salter sent him from a wood near Shanklin, and named it R. salteri in his honour. That action was in one way unfortunate, for the species appears to be endemic to just this one locality; but on the other hand the existence of this taxon served through the years to lure subsequent students of the genus to the island (though some of them, like W. C. Barton in 1932 and N. Douglas Simpson in 1938, seem to have paid no more than a fleeting visit for the sole purpose of collecting this one species). Between Salter’s time and the present only three other workers sampled the group in Wight at all widely. Two of these, J. G. Baker on a short visit in 1868 (records published in More 1871), and F. Townsend during the 1870s, while compiling the first edition of his Flora of the then single county (Townsend 1883), were Babingtonians taxonomically and in too many cases it is impossible to be sure to which present-day species their names relate. For, unlike Salter’s, hardly any of their specimens seem to have survived — none of the 50 or so variants which Baker claimed to have seen (including “‘R. fuscoater” and “‘R. rosaceus’’, with localities tantalisingly cited) and only two of Townsend’s (now in SLBI). The third worker, however, was the unrivalled authority of the post- Babington era, W. Moyle Rogers, who came over from his home in Bournemouth for a week in 1901 and was guided to a goodly range of the likeliest habitats by the island’s then leading resident botanist, F. Stratton. The 14 taxa added as a result (Rogers 1902) were the first determined in line with subsequent interpretation of the genus and therefore the first whose identities are for the most part unambiguous. Unfortunately, though, no voucher specimens appear to have been kept of some of the newly-recorded species (two of which have not been rediscovered). After Rogers the island’s brambles were effectively ignored until 1974. In that year I made the first of a succession of visits, several of a week in duration, in the course of which I have explored most of the more promising ground systematically. I am indebted to A. Newton, who paid a week’s visit himself in 1977, for much assistance with determinations throughout this period. My voucher material (including for all new vice-county records) has been deposited in RHMC and in certain cases in BM as well. SPECIES RECORDED The order and nomenclature in the systematic list that follows are those of Edees & Newton (1988). Unless otherwise indicated, habitats are as stated in that work. A number in brackets after a specific name is the total of 10-km grid squares in which the species has been recorded (maximum 10). An asterisk (*) denotes an addition made to the vice-county list in the period 1974-88, including redeterminations of older material, and + a species introduced to the island exclusively by human agency (‘alien’). RUBUS IN THE ISLE OF WIGHT 23 [R. DIVARICATUS MUELLER . The Wilderness, Rogers. Near Lynn, Salter. Both records need confirmation. | R. FISSUS LINDLEY (1) Apparently extinct. Hyde, near Shanklin, Rogers. A species that is still quite plentiful in parts of the New Forest. R. NESSENSIS W. HALL (4) Rare. Still in small quantity in America/Apse Castle Woods, where Salter (BM,E) knew it in abundance; also in Youngwoods Copse, in two places in its detached fragment to the west. The Wilderness, Rogers. Freshwater Marsh; Parkhurst Forest, Townsend. R. PLICATUS WEIHE & NEES (5) Less widespread than formerly, but still relatively plentiful on moist acid alluvium in The Wilderness and at the southern end of Munsley Moor; also on dry heathy ground on Bleak Down, Staplers Heath, Bonchurch Down and (a single bush) in Parkhurst Forest. R. SULCATUS VEST (1) Apparently extinct. Alverstone Copse, 1847, A. J. Hambrough, as R. suberectus (CGE, det. B. A. Miles). Near Alverstone, ‘“‘non rare in dumetis humidis’’, 1838, W. A. Bromfield, as ?R. plicatus (E). The localities are presumably one and the same. R. VIGOROSUS MUELLER & WIRTG. (5) Formerly widespread, now rare. Golden Hill Park, Freshwater, a large colony (with Viola lactea Sm.); Headon Warren, one bush, 1981, old pasture at south-eastern corner of Walter’s Copse, Newtown; hedge-top at cross-lanes north of Bleak Down (40/514.832); also until c.1978 (when eliminated by ploughing) plentiful among gorse outside Mount Farm Copse, Ningwood. Some of its former localities — Easton Marsh, A. H. Wolley-Dod; Colwell Heath, Rogers (SLBI) — have been altered out of recognition, but it is unclear why it seems to have disappeared from three others from which Rogers recorded it: Niton Down (“in great quantity’, 1883, CGE), Bleak Down and Parkhurst Forest. Further early records are: between Freshwater and Yarmouth, 1889, 7. R. A. Briggs; King’s Quay Copse, near Osborne, 1869, Stratton (OXF). *R. ALBONIS W. C. R. WATSON (3) Locally plentiful round Newport: St. George’s Down gravel pits, abundant; East Standen Wood, common; Great Briddlesford Wood; Staplers Copse, common; Parkhurst Forest, three bushes. Some of the old records of R. macrophyllus Weihe & Nees agg. may refer to this species. It may also have been the R. schlechtendalii recorded by Rogers from Marvel Copse, near Newport. *R. BOULAYI (SUDRE) W. C. R. WATSON (1) Rare. Parkhurst Forest, in some quantity in old oakwood in one area at northern end. *R. ERRABUNDUS W. C. R. WATSON (2) Rare. Parkhurst Forest, locally in some quantity; Bohemia grounds; St. George’s Down gravel pits, one bush. In all cases not the typical variant (which occurs in one or two places in Hampshire) but the micromorph that is general throughout the New Forest. [R. LENTIGINOSUS LEES Parkhurst Forest, “‘apparently this’’, Rogers. I cannot trace any specimen and this is an unlikely locality for this alder carr species, which was formerly interpreted more broadly. | R. LEUCANDRIFORMIS EDEES & NEWTON (7) Frequent across the northern half of the island (being tolerant of clay), but almost absent from the south. Particularly plentiful in Whitefield Wood, Brock’s Copse, Thorness Wood and Newbarn Copse, Calbourne. Salter’s records and specimens (BM,K) show that this, rather than R. lindleianus, was what he mainly understood by his R. nitidus. 24 D. E. ALLEN R. LINDLEIANUS LEES (8) Frequent. Particularly plentiful in The Wilderness, Parkhurst Forest and Sticelett Copse. Salter knew this mainly under the name of R. carpinifolius: there are specimens of his so determined from three localities (BM,CGE,E) and an 1847 one of Hambrough’s from a fourth (CGE). R. MOLLISSIMUS ROGERS (1) Possibly extinct. Bleak Down, in plenty by The Wilderness, Rogers & Stratton (BM). Also recorded by Rogers from Parkhurst Forest, but abundant and generally distributed there is a similar bramble which I believe to be a distinct, unnamed species (‘H252’). I have seen this in many parts of the island and in several places in the far south of Hampshire as well. It is common in the Caricetum paniculatae community in The Wilderness. *R. OXYANCHUS SUDRE (2) Very local: confined to West Wight. Mount Farm Copse, Ningwood, in abundance; Cranmore Wood, a patch; Norton copse, Yarmouth, a patch. Townsend recorded near Shorwell R. macrophyllus var. glabratus, the name under which he knew this species well in the Bournemouth district, but no specimen has survived and the determination seems unlikely to be correct. *R. PURBECKENSIS BARTON & RIDDELSD. (3) Rare. Staplers Copse, plentiful in an open, once-boggy strip by Blacklands Farm; Parkhurst Forest, two clumps in open heathy area north of Tuckers Gate. Near The Hermitage, St. Catherine’s Down 1916, C. Bucknall, as ?R. hirtifolius (BRIST). R. PYRAMIDALIS KALTENB. (2) Rare. Borthwood Copse, occasional; Bohemia grounds, one patch. Near Kingston, 1879, Townsend, det. Rogers. The micromorph (var. parvifolius Frid. & Gelert), the commoner variant in the New Forest, was collected by Miss E. S. Todd in 1926 from a “‘sandy common between Lake and Horringford”’ (BM, SDN) —- which sounds like Apse Heath. By a slip of the pen Miss Todd wrote ‘‘between Ventnor...” on the SDN label, which was transmuted into “‘below Ventnor’ — an impossible location — when the record was published by Rayner (1929). R. SALTERI BAB. (1) Rare. Still in some quantity in Apse Castle/America Woods, where Salter originally collected it in 1845, taking it for a tall form of R. plicatus (CGE,SLBI). Watson (1958) adds ‘‘and the downs to the south’’, but I have not succeeded in finding it there; his assertion that it is “frequent in the south-east of the island”’ is certainly much too sweeping. A clump of a putative hybrid with R. vestitus was found in Apse Castle Wood by Barton in 1932 (Barton 3788-3795, 3798 (BM)). The “‘R. salteri”’ recorded by Baker from several places in mid-western Wight was clearly R. leucandriformis. *R. ALTIARCUATUS BARTON & RIDDELSD. (3) Very local. Blackpan Common, Sandown; Borthwood Copse and hedgebanks to the west, frequent; north end of Apse Heath Copse, one bush; Eastview gravel pits, abundant, and a colony on nearby Bleak Down; Head Down; Shorwell to Berry Hill, frequent along ridge. This may have been the R. hirtifolius collected in Apse Castle Wood by Briggs and Rogers pre-1882 and later determined by Rogers and Focke as a form of R. leucandrus. R. AMPLIFICATUS LEES (3) Occasional. Quarr Wood; Firestone Copse; America Wood; Burnt House lane and copse; Bohemia grounds, frequent around gorse-heath; top of Bonchurch Down. Salter (1856) described it as frequent in woods and thickets, a claim supported by specimens of his from Shore Copse, Fishbourne (BM,CGE,E,SLBI), where in his day it was ‘‘abundant”’, and from Brays Hill (BM). *R. BOUDICCAE A. L. BULL & EDEES (4) Locally common in the south-east on the Lower Greensand, especially between Newchurch and Sandown, trickling north to St. George’s Down and reappearing in quantity way to the west in the Mottistone-Brighstone district. Usually in dry oakwoods or heathy scrub, it is frequent in alder- RUBUS IN THE ISLE OF WIGHT 25 willow carr on Blackpan Common. There is a specimen from Sandown in BM collected by F. A. Rogers in 1894 (det. Rogers as R. argentatus). R. CARDIOPHYLLUS LEF. & MUELLER (9) Local; common in some localities, rare or absent in many others. Tolerant of the clay. Widely recorded and collected (mainly as R. cordifolius) by Salter. *R. CORNUBIENSIS (ROGERS EX RIDDELSD.) RILSTONE (1) Very rare. Apse Heath Copse and adjoining America Wood, two or three scattered bushes only, 1974 and 1979. Not otherwise known outside Cornwall and Devon. *R. CURVISPINOSUS EDEES & NEWTON (3) Rare. Golden Hill Park, Freshwater, a large colony; Cortheath Firs, on margin of former heath (40/ 442.901), recurring southwards to Guyers Heath. *R. DUMNONIENSIS BAB. (3) Rare. St. George’s Down gravel pits, occasional; hedgebanks west of Borthwood Copse; St. Catherine’s Down, two clumps on crest of ridge. Two old records from West Wight — Freshwater, Rogers; Headon Warren, above Alum Bay, Stratton, det. Rogers — must be treated with reserve in the absence of specimens, as Rogers interpreted this species over-broadly. R. NEMORALIS MUELLER (5) Occasional: widely but rather thinly scattered and unexpectedly scarce in very suitable localities (just a single bush in The Wilderness, for instance). R. POLYANTHEMUS LINDEB. (7) Local: common in the south-east on the Lower Greensand, but absent from most of the western half of the island. At Shanklin it grows on dry exposed sea-cliffs (as also at Bournemouth). *R. PROLONGATUS BOULAY & LETENDRE EX CORBIERE (3) Very local: mainly in the district east of Yarmouth. Mount Farm Copse, Ningwood Common and Cranmore Common, in plenty and becoming abundant near the coast; St. George’s Down gravel pits, local; Staplers Copse, one bush. R. SUBINERMOIDES DRUCE (6) Common in many of the copses along the northern clay belt, also in Youngwoods Copse, Alverstone, in the south-east; but rare at best in the remainder of the island. R. SPRENGELII WEIHE (6) Local. Almost confined to the south-east, where it is still quite widespread; plentiful in at least two heathy woods (Borthwood Copse; Bohemia grounds) and occurs as well on the gravel caps of the higher downs. *R. ARMIPOTENS W. C. BARTON EX NEWTON (4) Very local. Abundant in conifer plantation, Mottistone Common; Head Down, Niton, one bush; Mount Farm Copse, Ningwood; Wootton Bridge, in hedges. This highly disjunct distribution lends support to my impression that this is a recent immigrant: an abundant species of south-eastern England, it is patently adventive in several of the few places in which it occurs in the western half of Hampshire. A. Newton, however, recalls seeing it in several places on his 1977 visit in which it had a natural look and believes it may be of more ancient status. *+R. HYLOPHILUS RIP. EX GENEV. (1) Very rare. A small patch in Firestone Copse, Newton. Apparently introduced with conifers. *R. LAMBURNENSIS RILSTONE (5) A variable bramble, common - in places abundant — in Wight and the adjacent parts of south-west 26 D. E. ALLEN Hampshire is regarded by Edees & Newton (1988) as conspecific with this Cornubian species. It was collected twice by Salter (BM) but not distinguished by him from R. vestitus. It is evidently one parent of a distinctive member of Section Corylifolii (““H375”’) that is widespread in the island. *+R. PROCERUS MUELLER EX BOULAY ‘HIMALAYAN GIANT (7) A fast-increasing escape, already widespread — and even common in Shanklin. R. ULMIFOLIUS SCHOTT (10) Very common. [R. WINTERI MUELLER EX FOCKE A bramble collected by E. S. Marshall in 1891 south-west of Shanklin towards Ventnor (BM,CGE) was misdetermined as this by Focke. It is either an unnamed local form or a recent R. ulmifolius hybrid. | R. ADSCITUS GENEV. (2) Rare. Except for a colony in Barton Wood, above Osborne Bay, confined to West Wight and mainly along its coastal margin. Originally found by Briggs in 1889 on the long-vanished Colwell Heath (BM) and at Totland Bay, it still boasts colonies on the cliff north of Colwell Chine and, more surprisingly, among chalk scrub along the inland side of Tennyson Down, where the high oceanicity presumably outweighs the unfavourable geology. *R. SURREJANUS BARTON & RIDDELSD. (1) Very rare. A large colony of the pink-flowered var. wealdensis Barton & Riddelsd., otherwise known only in W. Sussex (v.c. 13), in Cook’s Copse, north of Newbridge. It has the appearance of a comparatively recent arrival. R. VESTITUS WEIHE (5) Locally abundant (as Baker noted), but mainly confined to the clay and chalk. The red-flowered variant seems rare: I have it on record only from Quarr Wood, Ryde. R. MUCRONATIFORMIS (SUDRE) W. C. R. WATSON (3) Very local. Abundant in three old oakwood fragments south of Newport: Marvel Copse (a Rogers find, the only locality known up to 1976), copse by Standen House and wooded patch at eastern end of St. George’s Down. Otherwise scarce and extremely scattered: Parkhurst Forest, one bush; Beech Copse, Godshill Park; Calloways Farm lane, near Bohemia; top of St. Boniface Down; Mount Farm Copse, Ningwood; Headon Warren. *R. AEQUALIDENS NEWTON (2) Rare. In great abundance on the plateau heath capping St Boniface Down (the highest point on the island) — whence presumably have come recent-looking clumps in Parkhurst Forest, in two places towards the northern end of the main east drive, first noted in 1982. *R. HANTONENSIS D. E. ALLEN (3) Very local. Apse Heath Copse, abundant in northern half in holly wood; St Catherine’s Down, in two places; Head Down, Niton, in great profusion even on sunny hedge-tops. R. LEIGHTONII LEES-EX LEIGHTON (6) Frequent. Thinly scattered over the eastern two-thirds of the island, but locally plentiful in several of the woods, on clay and gravel alike. Unexpectedly, also common along the crest of the ridge from Shorwell to Berry Hill, with R. altiarcuatus. Salter knew it only around Ryde, as R. radula var. hystrix (BM,CGE,E). [R. MELANODERMIS FOCKE Listed by Edees & Newton (1988) for v.c. 10 in error.] RUBUS IN THE ISLE OF WIGHT oH | *R. MICANS GODRON (3) Local. In most of the clay copses from the Medina valley to Ryde, in several of them plentifully. There is an 1846 Hambrough specimen from Havenstreet in E (as R. apiculatus). This species seems to be indigenous in Wight, whereas in Hampshire it has everywhere a ruderal distribution and an adventive look. *R. MOYLEI BARTON & RIDDELSD. (1) Very rare. In small quantity in Quarr Wood, Ryde. Too abundant on the opposite side of the Solent for it to be likely that this is other than a recent colonist, even though there is a specimen in MANCH (det. Baker, as R. macrophyllus) collected on the coast by Quarr Abbey by C. Bailey in October 1888. *R. CINEROSUS ROGERS (1) Rare. Thorness Wood, common at eastern end (on clay) — whence must have come a solitary clump in Parkhurst Forest near Noke Gate. R. DENTATIFOLIUS (BRIGGS) W. C. R. WATSON (5) Locally common. Plentiful in woods and on heaths along the south-eastern coastal belt from Sandown to St. Catherine’s Down, and up to 6 km inland as far as Godshill and Alverstone; also in the north, in clay copses adjacent to Wooton Creek and in Parkhurst Forest (where it is locally common). It is sad that the new epithet of vectensis which Watson bestowed on this species in 1937 has proved to be superfluous, for it was certainly very apt. Watson chose as his lectotype one of several specimens collected by Salter, who, as his determinations (BM,E,K) show, was nevertheless badly confused by this variable species. *R. EFFRENATUS NEWTON (3) Very local. In abundance in two widely-separated localities: Blackpan Common, near Sandown, and Head Down, Niton, chiefly among bracken in the open but spilling into the oak hanger, with an outlying patch 1-5 km to the west by The Hermitage. When first collected in 1982, this was taken for a local cross between R. sprengelii and some glandular-aciculate bramble. Its identity with this species of north-western Wales, hitherto supposedly endemic there, was only realised and confirmed six years later. *R. LEYANUS ROGERS (2) Very local. Apart from a clump on the top of St Boniface Down above Ventnor, apparently confined to the Brighstone area: abundant in Moortown marshes and copses; Gaggershill Lane, a patch; conifer plantations on Mottistone Common and (Newton, 1977, in quantity) Brighstone Down. Probably a recent immigrant: Townsend worked the Moortown locality in the 1870s and recorded nothing resembling this species. R. BLOXAMII (BAB.) LEES (2) Very local: apart from two areas of Parkhurst Forest (where Townsend knew it), confined to West Wight, where it occurs in some quantity in all suitable habitats and abundantly in at least three copses. R. ECHINATUS LINDLEY (8) Local (rather than ‘‘frequent’’, as Salter described it). Mainly in the northern half of the island, as it tolerates clay and chalk, but, as in Hampshire, very patchy and plentiful only in one or two localities: Whitefield Wood, Guyers Heath and — according to Townsend — downs above Idlecombe. R. FLEXUOSUS MUELLER & LEF. (5) So well-marked a species that the failure of Salter, Baker and even Townsend to record it may mean that it has greatly increased, perhaps as the result of extensive felling. Certainly it is now widespread and abundant in several woods (Marvel Copse, Newport; Youngwoods Copse; Thorness Wood; America Wood; Bunkers Copse, Rookley) and, more often than in Hampshire, plentiful even in 28 D. E. ALLEN two or three places on heathland sheltered by bracken or gorse. It is conspicuously absent from the high-level heaths and I have seen but a single bush in all of West Wight. R. INSECTIFOLIUS LEF. & MUELLER (2) Rare. Bunkers Copse, Rookley, locally in plenty; St. George’s Down gravel pits, one bush; Parkhurst Forest (where Rogers saw it), in two areas but scarce. Possibly the two records in Flora Vectensis for R. fuscus Weihe also belong here: Bohemia grounds, Hambrough; heathy common about Lynn Farm, Salter. *R. LARGIFICUS W. C. R. WATSON (2) Rare. Godshill Park, in abundance; Borthwood Copse, one large patch; Mottistone, in two places. In all cases the Wealden variant, not the pinkish-flowered variant of Hampshire. *+R, RUDIS WEIHE (1) Abundant in the Rectory shrubberies, Old Shanklin, spilling over into nearby gardens. An obviously accidental introduction, but interesting in showing that this common North Downs species would have thrived had it only chanced to have arrived by natural means. [R. RUFESCENS LEF. & MUELLER Pan, Newport, Rogers. Though common in central and south-eastern England, this becomes a rare species in most of Hampshire and, as Rogers mistook poor material of R. leightonii for it in at least two cases, the record is best treated with reserve in the absence of the specimen. | *R. ANGUSTICUSPIS SUDRE (1) Very rare. In quantity along the southern margin of Whitefield Wood, between Ryde and Brading. A species otherwise of south-eastern Wales and the southern Welsh Marches. +R. DASYPHYLLUS (ROGERS) E. S. MARSHALL (1) Apparently extinct. St. George’s Down, 1870, Stratton (OXF) — presumably an adventive associated with the gravel workings. A fast-spreading colonist from further north in England, this species is unknown as yet in southern Hampshire. R. PHAEOCARPUS W. C. R. WATSON (3) Very local. In quantity in certain copses on either side of Wootton Creek (the site of the only pre- 1970s record, by Rogers), outstandingly so in Blanket Copse; also on the south coast, scattered over Blackgang Chine and a patch on Rew Down, Ventnor. Perhaps a recent immigrant: this is a species of south-eastern England which is noticeably spreading southwards in Hampshire. *R. RILSTONEI BARTON & RIDDELSD. (1) Very rare. Southern end of Beech Copse, Godshill Park, a large colony mainly growing in shade. When discovered in 1977, this was the first record of this species away from the far south-western part of Britain; but it has since been found at the south-eastern corner of the New Forest. R. CONJUNGENS (BAB.) ROGERS (4) Local. Salter knew this as R. wahlbergii, finding it chiefly in the north-east. A specimen of his from Bembridge constitutes the lectotype of Babington’s variety. R. NEMOROSUS HAYNE & WILLD. (8) Relatively more widespread and plentiful than in Hampshire, where it is mainly confined to the New Forest fringes. Locally common in wooded districts on the clay, but also in quantity in places off the clay (Moortown marshes, Brighstone; about Bohemia). R. PRUINOSUS ARRH. (4) Frequent. In both chalk scrub and acid woods on the Lower Greensand. RUBUS IN THE ISLE OF WIGHT 29 R. TUBERCULATUS BAB. (9) Common in the coastal areas (e.g. all over West Wight), thinning out inland. INNOMINATI As in other counties, numerous bramble bushes are to be met with that cannot be matched with any as yet named species. The majority of these are doubtless the highly localized products of crosses more or less in situ, but several variants occur in quantity over a wide area and some of these may eventually be rated worthy of description. The more noteworthy of the latter are:— ‘H257 A very distinct, densely glandular bramble occurring in great profusion in many of the clay copses between Ryde and Newport and in the south-eastern portion of Parkhurst Forest; also in the far south, in very small quantity, in the roughs on Ventnor golf course and on a wood-margin by Godshill Park House. A shade-lover, it is the Isle of Wight counterpart of Hampshire’s R. glareosus Rogers. It was collected on the margin of Combley Great Wood (‘‘near Guildford Farm’’) by Salter and Babington in 1845 (CGE,E,K), originally as R. rosaceus and later as R. lejeunei. There is also a specimen of Hambrough’s (woods, Haven Street, 1846) in E labelled R. pallidus. ‘H863’ Misdetermined on different occasions as R. cissburiensis Barton & Riddelsd. and R. babingtonii Salter (both of which are Hampshire but not apparently Isle of Wight species), this may be a cross between these two species in origin. It completely and abruptly replaces ‘H257’ in various clay copses on the western side of the Medina valley, recurring in profusion in other substantial pieces of woodland on gravel further west, spilling over here and there into Parkhurst Forest and reappearing, further west still, again in abundance, in Walter’s Copse, Newtown. This range is more or less coterminous with the royal hunting park that existed there in medieval times. I have collected the same bramble from a clay copse in central Hampshire, near Steventon (SU/528.483). “H165’ Abundant in Parkhurst Forest but tightly confined to it. There is a specimen in SLBI collected there by Townsend in 1879 labelled R. pallidus (subsequently referred by Watson to R. thyrsiflorus). This is identical with a bramble long known in parts of West Sussex which has been variously named R. fuscus subsp. obscurus (by Rogers) and R. erraticus (by Watson). Another unnamed species (‘H440’), widespread in the New Forest and long known as R. leucostachys var. angustifolius Rogers, occurs in one area of Parkhurst Forest. Townsend collected it there too, according to Rogers, who recorded it himself from Bleak Down. DISCUSSION The total of 59 named species accepted above as reliably recorded from the island is not particularly impressive when set against Hampshire’s 128 species. Even the far more isolated Isle of Man, well to the north in a region of altogether lower Rubus diversity, has 34 (Allen 1986). Concealed within the bare figure, however, are some very striking peculiarities. For a start, Wight has six species that are unknown in Hampshire. One of these is the narrow endemic R. salteri, privileged with a name by a mere historical accident, the other five all strongly western species: R. aequalidens, R. angusticuspis, R. cornubiensis, R. dumnoniensis and R. effrenatus. A sixth species, R. micans, might reasonably be added to this list if I am right in my belief that it is not indigenous in Hampshire. R. angusticuspis and R. cornubiensis, otherwise not known nearer than Gloucestershire and Devon respectively, have the appearance of relatively recent arrivals, but the others have a more established look (though R. aequalidens seems to be in the process of producing distant daughter colonies from its large population above Ventnor). Extensive heaths on high ground just inland of the English Channel form a natural halting-place for birds on 30 D- ES ALEEN passage, and may account for the island having two such unlikely species as R. aequalidens and R. effrenatus, the former with its headquarters in Pembroke (v.c. 45), the latter, more extraordinarily, known elsewhere only in north-western Wales. It is the list of absentees, however, that contains the greater number of surprises. Three of the commonest and most generally distributed Hampshire brambles, R. cissburiensis, R. glareosus and R. milesii Newton, are completely missing from its offshore neighbour and a fourth, R. moylei, has achieved its solitary foothold probably only recently. There is no apparent reason why all of these should not occur equally plentifully in Wight. Also absent inexplicably are such other widespread Hampshire brambles as R. bercheriensis (Druce ex Rogers) Rogers, R. bertramii G. Braun, R. euryanthemus W. C. R. Watson, R. platyacanthus Mueller & Lef. and R. rubritinctus W. C. R. Watson. The lack of the last of these is especially remarkable, for it is not only a western species but frequent round most of Southampton Water; the fact that it is not in Wight would seem to imply that it has colonized the coastal zone opposite only comparatively lately. The island’s climate may not indeed be as amenable to western species as one is tempted to assume from the presence of, and in some cases frequency of, those particular ones that do occur. R. adscitus certainly has the appearance of having only just managed to cling on, restricted till recently to the very extremities, in marked contrast to its wide distribution in Hampshire, while R. questieri Lef. & Mueller and R. longithyrsiger Lees ex Focke, both locally abundant in the district facing Cowes, are missing from Wight altogether. Nor are the various brambles special to the Southampton region, several of them as yet unnamed, as well represented across the Solent as one would expect from the profusion in which they occur on the adjacent mainland. The unnamed bramble ‘H107’ (in several places in south-eastern Wight and abundant in Youngwoods Copse, Alverstone) and R. hantonensis are the only two that have been found. Instead of being just an extension of Hampshire, the Isle of Wight is, rather, an amalgam of Dorset and Sussex in the overall character of its Rubus flora. As if symbolically, it is with those two other neighbours, indeed, that it shares its most widespread non-Hampshire species, R. dumnonien- sis. The Sussex element, admittedly, is small and consists merely of R. largificus, R. surrejanus var. wealdensis and the abundant ‘H16S’ of Parkhurst Forest, all of which have a recent look about them in their Isle of Wight localities; and it is conspicuously without such expectable species as R. decussatus W. C. Barton ex Newton (which is present, for example, on Hayling Island) and R. naldretti (J. W. White) W. C. R. Watson. In so far as it can be said to exist, therefore, it is probably no more than the outcome of haphazard, late infiltration. By contrast, the element connecting the island with Dorset — or rather, with the area round Bournemouth ~ is not only larger, but includes three species which have every appearance of being very long-established: R. boulayi, R. mucronatiformis and R. purbeckensis. Looking more recent, and mainly concentrated in West Wight, are R. bloxamii, R. curvispinosus, the micromorph of R. errabundus and R. oxyanchus. Again, though, there is a glaring absentee in the shape of R. melanodermis Focke, an absence all the more surprising in view of the frequency of that species on the nearest coast of France, in the district round Cherbourg. R. subintegribasis Druce ought to be present too, and perhaps also R. pullifolius W. C. R. Watson — though for the latter Milford-on-Sea appears to be the eastern limit in the coastal corridor opposite, which perhaps indicates a limited ability to spread. Finally, there is a group of species which, while occurring in Wight as well as Hampshire, are relatively more plentiful in the former than in the latter. These are R. albionis, R. amplificatus, R. boudiccae, R. dentatifolius, R. lamburnensis and R. vigorosus. Three of these are western species and in their case the difference may perhaps be attributable to the greater oceanicity of the island’s climate. R. boudiccae, however, is otherwise known only in the Midlands and East Anglia, so some other factor must presumably be at work to have made Wight so favourable to its spread, perhaps high summer temperatures. ACKNOWLEDGMENTS I should like to thank A. Newton for commenting on a draft of this paper in the light of his very extensive acquaintance with the group throughout the British Isles. I am also indebted to B. RUBUS IN THE ISLE OF WIGHT 31 Shepard for guidance on topographical points and for assistance in other ways with this project over the years. REFERENCES ALLEN, D. E. (1986). Flora of the Isle of Man. Douglas. B[aBincton], A. M., ed. (1897). Memorials, journal and botanical correspondence of Charles Cardale Babington, pp. 128, 138, 439. Cambridge. Devoy, R. J. (1987). The estuary of the Western Yar, Isle of Wight: sea-level changes in the Solent region, in Barer, K. E., ed. Wessex and the Isle of Wight: field guide, pp. 115-122. Cambridge. Epess, E. S. & Newron, A. (1988). Brambles of the British Isles. London. More, A. G. (1871). A supplement to the ‘Flora Vectensis’. J. Bot., Lond. 9: 72-76, 135-145, 167-172, 202-211 (Rubus, pp. 140-141). Rayner, J. H. (1929). A supplement to Frederick Townsend’s Flora of Hampshire and the Isle of Wight. Southampton. Rocers, W. M. (1902). On the distribution of Rubi in Great Britain. J. Bot., Lond. 40: 150-157. SALTER, T. B. (1839). Appendix, illustrative of the botany of Poole and its neighbourhood, in SYDENHAM, J. The history of the town and county of Poole, pp. 467-491. Poole. SALTER, T. B. (1845). Remarks on some forms of Rubus. Ann. Mag. nat. Hist. 16: 361-372. SALTER, T. B. (1848). Rubus (pp. 286-287), in BROMFIELD, W. A. Notes and occasional observations on some of the rarer British plants growing wild in Hampshire. Phytologist 3: 205—213, 269-291, etc. SALTER, T. B. (1856). Rubus, in BROOMFIELD, W. A. Flora Vectensis. London. TOWNSEND, F. (1883). Flora of Hampshire, including the Isle of Wight. London. Watson, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge. WEIHE, K. E. A. & NEES VON ESENBECK, G. C. D. (1822-27). Rubi Germanici. Elberfeld. (Accepted August 1989) rl ar Nace + a Ronald wwii at: wees tho tay his ti New tase He “nay plaka: nk, i ; . bavir? 5 epee 6 a sar b ‘a ie ate re , re sy ay Shas P 5 1%’! Goes has ALY * Hany ae AN ib ant ‘ + Oe ia v re saci To a8 ie a Aah Mee ee bbb Tac iat yr re 1ai4 4 yz" "% = a air wh? G5 et Lie ‘rye bahae lA. Aas! ap Au > 4 ‘ h Hy ys , rs AUR) dag 4d rt - . i) AMD wa. 3 TOA aa A aes as end OCR a i _ ate 4 : a | nth Gad’ SiG ar ee ? oF a : 4 yt) TAL eye tes \ id ‘Ast Ropmune tt , cree CR HER VRSERSET) 1 Ooo ¥D aA ti Waray Hore Ag if synibonben rete why bh. igsieeeurt ype Me Pua ahd + CGUSINTS £ yiy of R Tea ‘ i Ve. 14) : + a; aS Re rk cn ts tes vat jook ake ; Mn micionsiy wittheot sun enpedialhe Jepe are epesent. Tor horned Me ie ie ges: a fir #4 ¢ Can Se maideee exist, ranean 5 aii i ' j . ' j ; , OF ol pe’ any a de, J st, = © : i = d “ Wy, rf ; i. a . t ha on . ' ex as 2 o Wei a oe? pe Te aA, x. lbrlernas Ye - j ‘ ity Tot V8 ‘ nr; ; wires i) rere ’ bey » wat > * he hi is -) j Al yt i pte is elt, ‘din —_— 4 mT eYaur Sve ci air : raw wher ode \ ror re rane eek woroan weet } ave ees t ave . | oF Lae : ew ey 1. Gi! uy Ome we, Wien || nd ‘ ) :" pe if ; i nt i | =i Racking ae noe a pate bn ne ght x ‘ aay / me ie Fr | eee s ad | even _ Al um . f a y : : i ; i wo t . "i 1 RAL ; i Watsonia, 18, 33-36 (1990) 33 Cornus sericea L. in Ireland: an incipient weed of wetlands D.. KELLY School of Botany, Trinity College, Dublin 2, Ireland ABSTRACT Cornus sericea L. (Cornaceae) has been widely planted in wet habitats in Ireland as an ornamental shrub. It spreads through layering, and has become more or less naturalized at many sites. A list of Irish records is presented. C. sericea poses a threat to natural vegetation, especially to wetland woods. INTRODUCTION Cornus sericea L., Red Osier Dogwood, a shrub from eastern North America, has also been known as C. stolonifera Michx., Thelycrania sericea (L.) Dandy and Thelycrania stolonifera (Michx.) Pojark. It differs from the native C. sanguinea L. in its white fruits, its brighter, usually blood-red twigs, conspicuous in winter, and its ‘suckering’ habit. C. sericea also differs in being adapted to habitats subject to waterlogging, where it has been widely planted as an ornamental. (It is often planted in company with Salix alba L. var. vitellina (L.) Stokes, Golden Willow, another ornamental whose twigs provide colour during the winter.) C. alba L., from northern Asia, is very similar to C. sericea but is ‘non-suckering’ and the stone of the fruit is differently shaped. C. alba is reported as an escape in Britain (Moore 1987), but not in Ireland. MODE OF PROPAGATION AND DEGREE OF NATURALIZATION Cornus sericea is described by Cook (1968) as ‘cultivated for ornament and locally naturalized”’ in Austria, Britain, Finland and Switzerland. It is recorded in several local Floras as naturalized in England and Scotland from Kent (Philp 1982) to Angus (Ingram & Noltie 1981). In Wales, it is recorded from seven counties as an established alien of woodland and scrub (Ellis 1983). It was not included in the standard Irish Flora until the most recent edition (Webb 1977), where it is mentioned as ‘‘naturalized in a few places by rivers or in marshy ground”’. Scannell & Synnott (1987) record it from six Irish vice-counties, on the basis of my site records, all included below. Site records have already been published from West Galway (v.c. H16) (Webb & Scannell 1983) and Limerick (v.c. H8) (Reynolds 1988). For the compiler of records this species is generally a borderline case: it is rarely quite wild, but it is commonly no longer to be regarded as cultivated. At most sites the species clearly owes its presence to deliberate planting, and the distances that it has spread are limited. The proportion of flowers that set seed is normally low in Ireland, and I have searched in vain for seedlings at many localities. The increase of the species is very largely by lateral spread, by rooting from decumbent branches. The decumbent branches may grow horizontally at soil level for a few decimetres before turning up to form leafy shoots, which could be mistaken for juveniles derived from seed. This unspecialized form of vegetative reproduction seems best described as ‘layering’; the plant is described as “suckering” by Moore (1987) and as “‘stoloniferous” by Cook (1968). However termed, the process means that a single bush can expand to form an extensive thicket. Vegetative propagation over longer distances may have taken place by means of detached pieces of shoot. A natural means of dispersal would be through the action of rivers in spate. The single bush on Foynes Island in the Shannon estuary (v.c. H8; Reynolds 1988) is the most convincing case for 34 D. L. KELLY natural dispersal. The site is not near houses, and there is no evidence of the species having been planted anywhere on the island. The site is within the reach of high tides, and it seems quite possible that a piece of the plant was washed up here, having been torn from the Shannon’s banks somewhere upstream (e.g. from the station in Limerick city, v.c. H9). However, I have not found any cases of detached fragments in the process of becoming established, and I conclude that this must be, at best, a rare event. ANNOTATED LIST OF IRISH RECORDS The following is.a list of all sites of naturalization or incipient naturalization of Cornus sericea in Ireland known to me; the records are mine unless otherwise stated. V.c. H2, N. Kerry: Ross Castle (ruin), $.W. of Killarney, GR V/95.88. Margins of wet woodland close to Lough Leane, spreading by layering, 1972-88, TCD; north of Reen Point, S.W. of Killarney, wet woodland, GR V/94.89, 1973-74. V.c. H8, Co. Limerick: Cappercullin Glen (southern end), Glenstal Abbey (formerly Glenstal Castle), Moroe, between Cappamore and Newport, GR R/73.56, dominant and spreading in wet bottom of narrow glen in old demesne, 1973-86. N.E. of Battery Point, Foynes Island, Shannon estuary, GR R/24.52, single bush about 2-5 m high and 4 m diameter, with many decumbent rooting branches, growing within a few metres of the top of the stony shore just above high water mark, surrounded by large Salix cinerea subsp. oleifolia and Rubus fruticosus agg., 1985-88, S. Reynolds (pers. comm.) and Reynolds (1988). Naturalized by the lakes in Curragh Chase demesne, between Askeaton and Adare, GR R/4.4, August 1985, S. Reynolds (pers. comm.) and Reynolds (1988). Adare, right (east) bank of R. Maigue, GR R/468.466, single small stand beside the bridge, at the foot of a garden, 1986-88. V.c. H9, Co. Clare: Limerick city between Sarsfield Bridge and the Treaty Stone, bank of R. Shannon, GR R/57.57, medium-sized colony, “suckering freely”, April 1985, P. S. Wyse Jackson (pers. comm.). V.c. H11, Co. Kilkenny: Dysertmore, S.W. of Inistioge, left (eastern) bank of R. Nore, GR S/65.33, single medium-sized colony in low-lying river fringe dominated by Salix spp., September 1988. S.W. of New Ross, roadside, GR S/69.26, three small patches in hedgerow, up to 3.5 m high, layering, September 1988. V.c. H12, Co. Wexford: R. Barrow near Mount Garret Bridge, N. of New Ross, naturalized on the left (eastern) bank, GR S/72.30 — “‘I have known it there for many years” (S. Reynolds, pers. comm., 1988). V.c. H14, Laois: Lowlands Wood, Abbeyleix estate, left (eastern) bank of R. Nore, GR S/42.82, locally dominant as an understorey species in alluvial woodland, forming thickets up to 4 m high under canopy of Quercus robur L. (less waterlogged sites) and Salix cinerea subsp. oleifolia (more waterlogged sites), 1972-89, TCD; Grantstown (Granston) Lough, between Rathdowney and Durrow, GR S/31.79 and S/31.80; former demesne of Grantstown Manor, wet woodland with many planted species; scattered beds of C. sericea fringing lake at northern and south-western ends, also on small island (crannog?) in the middle of the lake, May 1986. V.c. H16, W. Galway: R. Corrib, western bank, on the N.W. margin of Galway city, large thicket — “It is steadily spreading among native vegetation by suckering, and may by now be regarded as naturalized” (Webb & Scannell 1983). V.c. H19, Co. Kildare: Donadea Castle (ruin), Staplestown, ornamental lake, GR N/833.329, small stand on marshy lake-margin, ‘“‘spreading by suckering’. 1985, P. S. Wyse Jackson (pers. comm.); 1987, D. L. K. & M. Sheehy Skeffington. Rye Water, Carton demesne, N.E. of Maynooth, wetlands by river and by tributary stream forming boundary with Co. Meath, GR N/ 95.38, forming dense thickets 2.5—5 m high, spreading by layering; includes an area with shallow braided streams between mud banks, with Caltha palustris L. and abundant Ranunculus ficaria L. growing under C. sericea, March 1987. V.c. H21, Co. Dublin: 1 km W. of Lucan, left (northern) bank of R. Liffey, GR O/024.352, two small stands c. 10 m apart, the lower branches subject to immersion, backed by deciduous woodland, 1987-89. R. Liffey, both banks, between Lucan and Palmerstown, GR O/05.35, O/ 05.36, O/07.35, O/08.35, O/09.35, occasional small stands at river margin, the lower branches CORNUS SERICEA IN IRELAND dD subject to immersion, the stands spreading by layering, M. Norton & S. Reynolds (pers. comm.), D. L. K. & M. Sheehy Skeffington, 1988-89. Santry estate, Santry, GR O/408.165, in swampy woodland, growing rampant at the edge of a stream, 1982-86, P. S. Wyse Jackson (pers. comm. ). Banks of R. Tolka near Ashtown, GR O/11.37, O/10.37, naturalized along river bank, 1988. M. Norton (pers. comm.) V.c. H22, Meath: Boyne Canal between Slane and Drogheda, close to the entrance to Townley Hall, GR O/035.755, April 1983, P. S. Wyse Jackson (pers. comm.). Daltonstown House, N. of Balrath (between Slane and Ashbourne), estate woods, GR N/99.66, forming dense thickets in damp cutover deciduous woodland, 1985-88. Rye Water, wetlands by river and tributary stream, Carton demesne (see entry under v.c. H19). V.c. H23, Westmeath: Clonhugh Woodland, Ballynafid, N. of Lough Owel, GR N/408.604, forming extensive thickets in swampy ground in semi-open woodland, spreading by layering, 1985-86, D. L. K.'& S. F. Iremonger: V.c. H25, Co. Roscommon: Drummans Island (actually a promontory), Lough Key Forest Park (formerly demesne of Rockingham House), southern shore of Lough Key, GR G/84.04, wooded lake-margin, spreading by layering, 1973-86. V.c. H28, Co. Sligo: near Waterglade Garden, Aghamore, near south-western corner of Lough Gill, GR G/71.32, several bushes by roadside, October 1986. Near Hollybrook House, W. of Lough Arrow, GR G/77.12, small stand in hedge by main road, October 1986. V.c. H29, Co. Leitrim: south of Clooncoe Lough, near Rinn Lough, GR N/11.91, in wetland wood, spreading from road margin and forming thickets up to 3 m high, May 1984. Near Aghavas, N.E. of Mohill, GR H/1.0 or H/2.0, roadside, spreading by layering, August 1984, D. L. K., D. A. Webb & N. McGough. V.c. H33, Fermanagh: Crom Castle demesne, S.W. of Derrymacrow Lough, W. of Newtownbutler, GR H/36.24, small stands by side road in conifer plantation, up to 3 m high, spreading by layering (also forming scattered shrubberies in wetland below Crom Castle, clearly planted), June 1987. V.c. H36, Tyrone: Curran Lough, N.W. of Benburb, GR H/50.54, locally plentiful, 1988, J. S. Faulkner (pers. comm.). Augher Lake, GR H/560.538, 1988, I. McNeill (pers. comm). V.c. 37, Co. Armagh: Caledon House estate by R. Blackwater, W. of Killylea, GR H/7.4, locally plentiful, 1988, J. S. Faulkner (pers. comm.). V.c. H39, Co. Antrim: Rea’s Wood, fringing Antrim Bay at north-eastern corner of Lough Neagh, GR J/14.85, forming dense thickets up to 4.5 m high in the midst of damp deciduous woodland, spreading by layering (also planted at nearby big house — formerly Skeffington Lodge, now Deerpark Hotel), September 1986, D. L. K., R. Forbes & S. Beesley. V.c. H40, Co. Londonderry: between Kilrea and Garvagh, several patches and single bushes along a country road, GR C/895.134, C/887.135. ““The largest and oldest is a stretch of about 50 metres starting from C895134 in the east’’; this originated from “‘a couple of clusters of say 4 or 5 bushes on the road verge” planted in the 1930s. The bushes do not seem to be spreading, February 1989, D. S. Lambert (pers. comm.). These records are of the yellow-twigged variety, C. sericea var. flaviramea (Spaeth) Rehd., ‘Golden Twig Dogwood’. ECOLOGY AND AGGRESSIVENESS Cornus sericea has been widely planted beside ornamental waters in parks and demesnes; it is clearly becoming naturalized at scattered localities throughout Ireland. Records are confined to lowland sites on more or less eutrophic soils, subject to varying degrees of waterlogging. A typical soil profile was investigated beneath a thicket of C. sericea in the Abbeyleix estate (v.c. H14, Laois; GR S/ 419.816), on river alluvium under a canopy of Salix cinerea subsp. oleifolia. The soil surface was covered by a thin layer of leaf litter and was almost devoid of herbs and bryophytes. The soil was a gley with a wet humus-rich horizon, making a gradual transition at 4-8 cm to a very wet, clayey horizon, grey-brown with orange mottling. The top 5 cm had a loss on ignition of 62% and a pH of 6.8 (1:1 water:fresh soil). Analyses gave total N = 6100 yg ml~! (22,600 ug g~'), C/N quotient = 15.6, total P = 200 ug ml‘ (750 wg"), available K = 22 yg ml“! (83 wg g_') and available Ca = 2000 ug ml~! (7450 ug g~'). For the conservationist, the species is already of considerable significance. It is an aggressive 36 D. L. KELLY invader of natural and semi-natural wetland habitats, both open and wooded. Its extensive, dense thickets have no equivalent in our natural vegetation. The pre-existing herbaceous flora is almost totally suppressed, so species-rich communities are converted to species-poor ones. The most serious case known to me is in the alluvial woodland by the R. Nore in the Abbeyleix estate. A substantial portion of this historically important wood (Kelly & Fuller 1988) is now rendered impenetrable by dense thickets of C. sericea. The problem is reminiscent of the invasion by Rhododendron ponticum L. in woodlands on acid soils (Cross 1982), but the lack of seeding in C. sericea means that its control should be relatively straightforward. Ireland’s wetland woods are diverse and full of interest; they are, however, sparsely distributed, small in area and subject to various other threats to their survival. Control of C. sericea should therefore be a matter of serious concern. ACKNOWLEDGMENTS I am grateful to Ms S. Reynolds, Dr P. S. Wyse Jackson, Ms M. Norton, Ms D. S. Lambert, Dr J. Faulkner and Mr I. McNeill for field records; to the Soils Division of An Foras Taluntais/Teagasc (The Agricultural Institute) for soil analyses; and to Miss M. J. P. Scannell, Mrs M. Briggs, Mr P. Hackney and Professor D. A. Webb for helpful information and comments. REFERENCES Cook, C. D. K. (1968). Cornus L., in Tutin, T. G. et al., eds. Flora Europaea 2: 313-314. Cambridge. Cross, J. R. (1982). The invasion and impact of Rhododendron ponticum in native Irish vegetation. Roy. Dublin Soc., J. Life Sci. 3: 209-220. Exuis, R. G. (1983). Flowering plants of Wales. Cardiff. INGRAM, R. & NortigE, H. J. (1981). The flora of Angus. Dundee. KELLy, D. L. & FULLER, S. (1988). Ancient woodland in Central Ireland: does it exist?, in SALBITANO, F., ed. Human influences on forest ecosystem development in Europe, pp. 336-339. Bologna. Moore, D. M. (1987). Cornaceae, in CLAPHAM, A. R., Tutin, T. G., & Moore, D. M. Flora of the British Isles, 3rd ed. Cambridge. PHILP, E. G. (1982). Atlas of the Kent flora. Kent. REYNOLDS, S. (1988). Plant records from Co. Limerick (H§8). Ir. Nat. J. 22: 533-534. SCANNELL, M. J. P. & SyNnnott, D. M. (1987). Census catalogue of the flora of Ireland, 2nd ed. Dublin. Wess, D. A. (1977). An Irish Flora, 6th ed. Dundalk. Wess, D. A. & SCANNELL, M. J. P. (1983). Flora of Connemara and the Burren. Cambridge. (Accepted March 1989) Watsonia, 18, 37-48 (1990) 37 The current distribution and abundance of Orchis ustulata L. in southern England M. J. Y. FOLEY 87 Ribchester Road, Clayton-le-Dale, Blackburn, Lancs., BB] 9HT ABSTRACT Orchis ustulata L. (Orchidaceae) was formerly widespread over much of the calcareous grassland of southern England. Three hundred and seventeen previously recorded sites in 152 separate 10-km squares have been identified, and a survey of the present status of the species at these shows that it has become extinct at many of them, and that there are now only 60 colonies (in thirty one 10-km squares) which definitely survive, only nine of which usually exceed 200 flowering plants. The past and present distribution of O. ustulata is illustrated, and some aspects of its ecological preferences and causes of decline discussed. Current colony strengths are indicated and all records traced are presented. INTRODUCTION In a previous survey (Foley 1987), the current distribution and abundance of Orchis ustulata L. in northern England was reviewed and the causes of its decline discussed. A similar survey of sites in southern England has now been completed and the results are reported below. O. ustulata is widely distributed throughout central Europe and reaches its northern limit in the island of Gotland, southern Sweden (Ekstan et al. 1984), and the Leningrad region of the USSR at 60° N (Meusel, Jager & Weinert 1965). It is most frequently found in the British Isles on the old, undisturbed chalk and limestone grasslands of southern and south-eastern England at which a light grazing regime has been maintained, and which have not been subjected to agricultural ‘improve- ment’. The species is unrecorded in Ireland and Scotland, and the only record from Wales is almost certainly an error. The area covered by this survey includes the whole of southern England, namely vice-counties 1— 34, 36-40, 53, 55 and 55b, and completes the author’s survey of the known distribution of O. ustulata within Britain (Foley 1987). The following account is based upon field-work carried out by the author and colleagues, and investigations into old records. DISTRIBUTION Three hundred and seventeen recorded sites have been identified in southern England, of which 186 are definitely extinct, a further 24 are possibly so, and 47 are of unknown status but likely to be either small or extinct. These last records are from Wiltshire (v.cc. 7-8) and E. Sussex (v.c. 14), where it has been difficult to trace precisely all of the many records in old Floras and other sources, some of which may duplicate known sites. The past and present distribution of O. ustulata is shown in Figure 1. Out of a total of 152 recorded 10-km squares, this species is definitely present in only 31. The progressive extinction in other squares can be seen in Table 1. The great majority of sites were, and are, on calcareous ground — principally the chalk downs, but also on limestone grasslands. In the past, the highest density of sites was in Wiltshire, Hampshire and Sussex, but good colonies were also known in Dorset, Wight, Kent, Hertfordshire, Berkshire, Cambridgeshire, Bedfordshire, and Gloucestershire. Even today, the main stronghold of O. ustulata is still the chalk downland of N. and S. Wilts. M. J. Y. FOLEY 38 -souenbs Wy-(OT Ul panod “g¢s¢ “SS “ES ‘Or-9E ‘pE-T “99°A ‘puv[suq UJOYINOS Ul YIDjNISN $1494 9 S v € #0861 ONIAIAYNS 7714S * OS ANaISSOd yO LONILX3 XX jo uonnquisip juosoid pue ysed oy] *[ TAN ORCHIS USTULATA IN SOUTHERN ENGLAND 39 TABLE 1. TIME-SCALE OF THE PROGRESSIVE EXTINCTION OF O. USTULATA FROM THE 152 RECORDED 10-KM SQUARES OF SOUTHERN ENGLAND Total number Extinction Date Pre- of 10-km square Period:— Unknown 1900 1900-29 1930-59 1960+ Extant records Number of 25 55 13 14 14 31 152 10-km squares (v.cc. 7-8) and E. Sussex (v.c. 14), where the plant survives, often in very strong colonies, on some of those fragments of grazed downland which have escaped modern agricultural upgrading. Many of these are nature reserves or areas of special conservation interest relating to orchids and to chalk grassland plants in general. Unfortunately, and in contrast with many sites in northern England, where steep slopes or outcropping rock has prevented ploughing, most southern sites can be destroyed relatively easily in this way. In southern England, the western limit of O. ustulata was formerly near Wembury, S. Devon (v.c. 3), but the presence of this colony has apparently not been confirmed since 1932. It may be that the small Fontmell Down colony, Dorset (v.c. 9), now represents the western limit of extant sites, which then rapidly increase in number eastwards, with Wiltshire (especially v.c. 8) possessing the largest populations in Britain. ECOLOGY O. ustulata is most frequently to be found in forb-rich, moderately grazed calcareous grassland which has been physically undisturbed in the past, and untreated by artificial fertilisers or herbicidal/ pesticidal sprays. Fairly constant associates include Poterium sanguisorba, Anthyllis vulneraria, Lotus corniculatus, Primula veris, Rhinanthus minor, Polygala spp., and Gentianella spp.. The orchids, Orchis morio, O. mascula, Platanthera bifolia, Gymnadenia conopsea, and Dactylorhiza fuchsii are frequent associates also, and less often Coeloglossum viride and other plants such as Blackstonia perfoliata and Campanula glomerata. The usual habitat is on gentle to steeply sloping calcareous ground. This can be of almost any aspect, but is most frequently warm, south-westerly to south-easterly-facing on rather steep-sided hollows, banks, and dry valleys in the chalk downs. Ancient earthworks, by virtue of having remained undisturbed for many hundreds of years, are often a favoured habitat and frequently harbour sizeable colonies. There are, however, very large areas of suitable grassland containing all the species normally associated with O. ustulata, but from which the plant is absent, and even some of the largest colonies are quite localized in extent. Only the three strongest located in this survey, each having several thousand flowering plants in a good season, have continuous populations covering an area greater than 1 ha, and many good sites are much smaller, or if not, the plants within them are thinly and sporadically distributed. What is now probably the finest remaining British colony, consisting of an estimated 3000—S000 flowering plants, is in S. Wilts. (v.c. 8), and lies on gently undulating, multi-aspect downland forming a large continuous population with associated outliers. The site has a well-documented history of traditional grazing and agricultural usage reaching back possibly two millenia and is now fortunately fully protected as a nature reserve. A second very large colony, also in v.c. 8, and estimated at 2000-3000 plants, lies on the very steep west-facing slope of a dry valley and is quite different to the former in topography, and possesses probably the highest concentration of plants (1000+ per ha) of any of these very large sites. Since the decline of the rabbit population through myxomatosis, the level and timing of managed grazing appears to be critical to the long-term survival of O. ustulata. Light grazing, preferably by Sheep, from early Spring to the end of April, with a further spell in late July through until September, plus occasional dressings of natural organic manure, seems to be the favoured pattern at sites with the largest populations. This time-scale leaves ample opportunity for flowering and seeding. Cattle grazing, unless very light, may destroy the orchid by trampling or cause damage to 40 M: J..Vi FOLEY friable turf, thereby allowing the establishment of intrusive colonising plants, and is probably best avoided. Some sites are currently very overgrazed, e.g. Yarnbury Castle (v.c. 8), where small stunted plants, often only 3-5 cm in height, are found. In contrast it has been found (Foley 1987) that the plant can survive in quite rank, ungrazed localities, where tall specimens of 20-25 cm are normal, although survival of such colonies may be for a limited time only, and at least one such colony in northern England appears to have a diminishing number of plants. In southern England O. ustulata has a peak flowering period from late May to early June, after which the plants rapidly senesce, especially if seed has not been set. However, a late-flowering variant flowering in early- to mid-July occurs at several sites forming colonies often quite separate from the earlier-flowering variant, although for example at Willingdon, E. Sussex (v.c. 14), both variants may grow together. Most records of this late-flowering variant are from Wiltshire, Hampshire, and Sussex, where it occurs in relatively close proximity to the earlier-flowering variant, but such colonies seem to comprise comparatively few plants, usually less than 100. Later-flowering variants have been recorded from continental Europe, but are unknown in northern England. Morphologically, there are no obvious differences between the two variants, although late- flowering plants sometimes appear to have darker sepals. In N. Wilts. (v.c. 7), one colony of late- flowering plants lies within 1 km of one of the early-flowering ones, both in a typical habitat for O. ustulata and with similar associates. Both also form discrete colonies, and such a difference in flowering period is difficult to ascribe; rather surprisingly, in this case, these late-flowering plants are somewhat smaller than the nearby normal variant. It is possible that heavy Spring grazing over a period of many years, might lead to the development of a late-flowering ecotype, but whatever the reason, it requires further study. White-flowered variants of O. ustulata, referable to var. albiflora Thielens, have been recorded in the past from several sites in southern England (see Records), but none were encountered in the course of the present survey, although recently reported from two sites in northern England (Foley 1987). CURRENT STATUS AND CAUSES OF DECLINE The current status of O. ustulata at each site is denoted below (Records) by the letters: A = typically 1-10 flowering plants; B = 11-25; C = 26-50; D = 51-200; E = 201-1000; F = 1000+; PX = possibly extinct; X = extinct; U = unknown status, but likely to be either small colonies or extinct. Most extant sites and many other recent and less recent ones have been visited by the author and colleagues within the last few years, and estimates of colony size are usually based on observations made over several seasons. Comments relating to the above scale of estimates and to presumptions of extinction have been made previously (Foley 1987) and are equally applicable to the present survey. Of the 42 vice-counties in the survey area, O. ustulata has in the past been recorded from 35, at some 317 separate sites. The present survey has shown the species to be now extinct, or probably so, in 21 vice-counties (Table 2); in a further eight, the number of sites is now reduced to less than 20% of those recorded in the past. In a further three the position is slightly healthier, but even here TABLE 2. CURRENT STATUS OF O. USTULATA IN SOUTHERN ENGLAND BY VICE-COUNTY (SEE RECORDS) Surviving at present in less than 20% of Presumed recorded sites in Modest populations Main Not recorded extinct Vics survive populations L Ane lds ate Onl Oal7.019, 9, 10,313 5,205,22,,30; 11, 12,15: 7, 8, 14. 215, 30. 2A, L3¢245 Ldip sph oe LB, Libs Ly Sh, 22, 54. 30, 21, 39, AU, 5), J90. ORCHIS USTULATA IN SOUTHERN ENGLAND 41 TABLE 3. ABUNDANCE OF O. USTULATA FOR EACH VICE-COUNTY OF SOUTHERN ENGLAND Vice-county Colony status (see code) 2S —- — - _ Nu beer of Number Name A B C D E F a PX x colonies 1 W. Cornwall No record 2 E. Cornwall No record 3 S. Devon 1 3 4 4 N. Devon No record 5 S. Somerset No record 6 N. Somerset 7] 7. 7 N. Wilts. 3 2D 2 1 2 7 1 18 8 S. Wilts. 6 1 3 3 36 1 1 51 9 Dorset Ds 1 9 is 10 Wight 1 1 10 12 11 S. Hants. 1 1 1 4 7 12 N. Hants. 5 2 9 16 13 W. Sussex 1 5 6 14 E. Sussex 5 5 2 1 1 9 WD 25 15 E. Kent 6 1 8 15 16 W. Kent 3 3 17 Surrey Z 8 10 18 S. Essex No record 19 N. Essex 2 2 20 Herts. 1 1 9 11 24 Middlesex 1 1 22 Berks. 1 7 9 ib 7/ 23 Oxon 1 1 2 24 Bucks. 5) 5 25 E. Suffolk 1 1 26 W. Suffolk 5 5 27 E. Norfolk No record 28 W. Norfolk 1 1 29 Cambs. a2 12 30 Beds. 1 9 10 31 Hunts. 1 1 32 Northants. 4 4 33 E. Gloucs. 1 1 19 P25 34 W. Gloucs. 8 8 36 Herefs. 9) 2 37, Worcs. 3) 5 38 Warks. No record 39 Staffs. 2 7), 40 Salop 6 6 3)5) S. Lincs. 1 9 10 25) Leics. 2. 4 55b Rutland 3 3 Total 30 11 6 4 5 4 47 24 186 Sil7, populations are generally small, and an examination of Table 3, in which current colony strengths in each vice-county are displayed, shows that S. Hants (v.c. 11) has only two small populations remaining, N. Hants (v.c. 12) has seven modest ones, and E. Kent (v.c. 15) has seven small ones. Apart from the total extinctions indicated above, other rates of decline have been dramatic — Gloucestershire (E. Gloucs., v.c. 33, and W. Gloucs., v.c. 34) had a total of at least 29 separate populations in the past, but only one colony was definitely found to survive during the course of this survey, and other vice-counties with many previously recorded populations show the same trend, e.g. Dorset, v.c. 9, (two remaining from twelve), Berks., v.c. 22, (one from 17), Wight, v.c. 10, (one from twelve), Herts., v.c. 20, (one from eleven), and Beds., v.c. 30, (one from ten). The stronghold 42 M.'J. Y° FOLEY TABLE 4.ABUNDANCE OF O. USTULATA IN THOSE 10-KM SQUARES OF SOUTHERN ENGLAND IN WHICH THE SPECIES SURVIVES 10-km square Number of colonies Colony status (see code) 31/8.1 31/9.5 40/4.8 41/0.1 41/0.4 41/0.6 41/1.2 41/1.6 41/2.2 41/2.4 41/2.7 41/2.8 41/3.3 41/3.5 41/3.6 41/4.5 41/5.5 41/5.8 42/1.3 43/9.4 50/5.9 2A 51/3.0 51/4.0 51/4.1 51/5.0 51/8.6 32/13 52/3.3 61/0.4 61/2.4 n> iui Le 9 | m> om lope) E>. “ & ce 0 ,A,A,A,B,B,E re eR TD RRR RR RRR RR RB RP Re ND eR hh BRR Rh WR Re PPP UPPrPrNOIDPSPADPS PPS PPP PP PUPPrnNPaMy > le QO ,A,A,A,B s Total 31 of distribution is now undoubtedly S. Wilts (v.c. 8), E. Sussex (v.c. 14), and to a lesser extent, N. Wilts (v.c. 7), all of which have some excellent thriving colonies, and on some of the extensive downland especially in S. Wilts., other relatively small colonies may yet await discovery. However, it can be seen that of the 60 extant colonies confirmed during this survey, there are now only nine which in a good year contain typically 200+ flowering plants, and only four of these exceed 1000. A breakdown of current colony size by 10-km square is given in Table 4. Causes of decline appear to be more frequently due to agricultural improvement than to other direct physical causes, and to some extent this contrasts with the situation in northern England where other pressures such as building and industrial encroachment have taken their toll. Ploughing with or without re-seeding, and the use of artificial fertilisers or herbicides are the main causes of site loss in southern England. During World War II, much downland, which because of its gentle topography could be easily worked agriculturally, was pressed into use, and many suitable habitats for O. ustulata were consequently lost. Overgrazing may also have had a detrimental effect, and still occurs at many sites. This, whilst perhaps not too damaging in the short term will, if prolonged over many years, greatly reduce population numbers, although usually some plants may persist in a vegetative state. Other than physical destruction, an examination of the records for O. ustulata (see also Figure 1) reveals a declining distribution towards the south and east of England, and to those geographical centres of former population density. This reinforces the theory of a deteriorating climatic situation, causing a contraction of range perhaps over the last several decades, and a similar observation has ORCHIS USTULATA IN SOUTHERN ENGLAND 43 been made for Ophrys sphegodes (Hutchings 1987), another orchid of calcareous grassland, which like O. ustulata is also near the limit of its range in southern England and is in decline. If such is the case, recently extinct but still intact sites for O. ustulata would be less likely to be subsequently recolonised in the less favourable conditions pertaining, and the existence of much seemingly suitable but unoccupied habitat in the survey area is typical of a species at the edge of its range — and if now under conditions of a deteriorating climate, is likely to remain so. Other, minor causes of loss of sites include scrub colonisation, quarrying, and building development. Fortunately many of the surviving colonies are protected either as nature reserves or by having S.S.S.I. status, and here O. ustulata should respond to management. Other colonies not so protected usually lie on easily worked ground and at these sites agricultural upgrading will remain a threat. During the course of this survey, some of the more obscure old records have been found difficult to trace and have not yet been fully identified; but work on these continues. Publication of this survey may generate information on these and other sites, which the author will be interested to receive, with a view to presenting a future summary. RECORDS Note: References in the form 00/0.0 refer to 10-km squares, whilst four-figure references placed after localities and in the form (00.00) refer either to tetrads or to 1-km squares; however these latter are not given for large populations or for sensitive localities. S. Devon, v.c. 3: 20/5.4, Wembury (Bovisand), recorded in 1932 (PX); 20/8.3, Start Point, specimen in DMTH, J.R.P. Furze, undated (X); 20/9.5, Berry Head, Brixham, prior to 1860 (X); 20/9.6, Babbacombe Down and Torquay, pre-1860 (X). (An old 19th century record for 20/5.6, Shaugh Vale, near Plymouth is thought to be an error.) Ivimey-Cook (1984) states: ‘“There is no evidence for the continued existence of Orchis ustulata L., the Burnt Orchid, in Devon’’. N. Somerset, v.c. 6: 31/3.6, Worlebury Hill (32.62), last record in 1838 (X); 31/4.5, Mendip Hills, three sites, at two of which it became extinct in last century (near Churchill (44.58) in 1847; and at Wavering Down, W. F. Miller, (40.56) in 1892) (Roe 1981); whilst at the third site, Callow Hill, it was last seen in 1959 by J. Hodgson, shortly after which the site was quarried away (R. G. B. Roe pers. comm.), but there is a specimen in LANC from the Cheddar area collected by E. Hodgson in 1961 in a “limestone copse”’ which may be the same site as Callow Hill (X, X, X); 31/4.7, Weston- in-Gordano (44.74) last recorded in 1850 (X); 31/5.7, Leigh Down (54.72) last recorded in about 1850 (X); 31/7.6, Claverton Down, Bath, (76.62) plentiful up to the end of the 19th century and last recorded in about 1920 by N. Y. Sandwith (Roe 1981), after which it is thought to have succumbed to human pressure and changes in agricultural practice (X); known in the Bath area since c. 1760. N. Wilts., v.c. 7: 31/8.6, South Wraxall (Grose 1957) (X); 41/0.6, Morgan’s Hill, Roundway Hill, Tan Hill, Beckhampton Down, four sites all mentioned by Grose (1957), but agricultural spraying in this area may have eliminated some populations (D. Green pers. comm.) (PX,PX,PX,PX); Cherhill S.S.S.I. (04.68), 8 plants in 1978 (D. Green pers. comm.) (A); 41/1.6 Pewsey Down N.N.R., very strong group of early- and late-flowering populations which include Wansdyke (10.64) (B); Milk Hill (E); Walkers Hill (D); and the late-flowering Knap Hill colony (D); Martinsell Hill, Golden Ball Hill (Grose 1957) (PX,PX); 41/1.7, Rockley Down (Grose 1957) (U); 41/1.8, Wroughton (Grose 1957), last reported in 1888, now very doubtful (D. Green pers. comm.) (PX); 41/2.7, Peaks Downs (24.78), 11 plants in 1985 (D. Green pers. comm.) (B); Aldbourne (Grose 1957) (U); Bailey Hill (26.78), less than 10 plants in 1987 (D. Green pers. comm.) (A); 41/2.8 Russley Down, (26.80), 6 plants in 1987 (D. Green pers. comm.) (A). S. Wilts., v.c. 8: 31/8.3, Mere (Grose 1957) (U); 31/8.5, Westbury Downs (Grose 1957) (U); 31/9.3 (or 9.4), Bishopstrow Down and Stockton Down (Stearn 1975) (U,U); Park Bottom (95.37) (Stearn 1975) (U); near Grant Ridge Wood (Grose 1957) (U); 31/9.4, Scratchbury Camp, Heytesbury, near Warminster, 3 sites given by Grose (1957) (U,U,U); 31/9.5, Great Cheverell Hill, nature reserve, south-facing slope with Platanthera bifolia, up to 500 plants in good years (E), some with pure white flowers (Stearn 1975); Bratton (Grose 1957) (U); 31/9.6, Heddington (Grose 1957) (U); 41/0.2, Bishopstone (Grose 1957) (U); 41/0.3, Stapleford, collected by W. M. 44 M. J. Y. FOLEY Rogers in 1873, LANC (U); Little Langford, Ebsbury Hill, Barford Down, Steeple Langford — Cow Down, 4 sites given by Grose (1957) (U,U,U,U); 41/0.4, Yarnbury Castle, Winterbourne Stoke, around ancient earthwork, heavily grazed, but potentially a strong colony (E); Parsonage Down, two very strong populations, respectively (E,F); West Down, Tilshead (06.48) M.O.D. area with heavy vehicles active, formerly robust plants with Orchis morio (B. G. Tattersall pers. comm.) but now ‘improved’ (X); Orcheston Down, 50 plants in 1988 (B. G. Tattersall pers. comm.) (C); near the Bustard, some with white flowers (Stearn 1975) (U); 41/0.5, near Wilsford (Grose 1957) (U); 41/0.6, Etchilhampton Hill (Grose 1957) (U); 41/1.2, Pennings Lane (11.24) seen in 1987 (A. Hutchison pers. comm) (U); Homington Down (12.24), two small colonies in 1987 (A. Hutchison pers. comm.) (A,A); Odstock Down, very strong population on west-facing steep slope (F); Clearbury Down, extensive population on gentle south-facing slope (F); 41/1.3, High Post (Stearn 1975) (U); Netton, Laverstock, (Grose 1957) (U,U); 41/1.4, Stonehenge, recorded by J. R. Akeroyd in 1966 when (B), W. M. Rogers, specimen in LANC (PX); near Fittleton and Nether Avon, Amesbury, Milston, Larkhill, (Grose 1957) (U,U,U,U); 41/1.5, Milton Lambourne (Grose 1957) (U); 41/2.2, Witherington Down (20.24) currently heavily grazed (A); Dean Hill (Grose 1957) (U); Pepperbox Hill (21.24), occasional in 1985 (A. Hutchinson pers. comm.) (A); 41/2.3, Winterslow (Grose 1957) (U); 41/2.5, Sidbury Hill, Ludgershall, Everleigh, Easton Hill, all given by Grose (1957) (U,U,U,U); 41/3.5, Fosbury (30.56), two plants of late-flowering variant, July 1988 (A. Hutchinson pers. comm.) (A); 41/3.6, between Ham Hill and Rivar Hill (Stearn 1975) (U); Ham Hill Reserve, about ten plants of late- flowering variant, July 1988 (A. Hutchison, S. L. Jury & J. R. Akeroyd pers. comms) (A). (Grose (1957) comments that although there are many sites scattered throughout v.cc.7-8, many have only a very few plants.This is probably even more the case today.) Dorset, v.c. 9: 30/8.8, Lulworth, last recorded in 1895 (X); East Stoke, last record 1895 (X); 30/8.9, S. Milborne St Andrew in 1895 (X); 31/8.0, Stourpaine, last seen in 1904 (X); 31/8.1, Hod Hill, last recorded in 1930, but may still persist (PX); Fontmell Down (H. J. M. Bowen pers. comm.) (A); 31/9.0, Blandford, not seen since 1799 (X); 31/9.1, Gussage St Michael in 1895 (X); Farnham in 1895 (X); Ashmore in 1895 (X); 40/0.8, Old Harry Cliffs (05.82) in 1895 (X); 41/0.1, Bokerley Ditch, Martin Down (05.17), still surviving (H. J. M. Bowen pers. comm.) (A). Wight, v.c. 10: 40/4.8, Garstons Down, discovered in 1973 at a bowl-shaped depression in the chalk, usually 20-30 plants, but over 300 in 1974 (C) (B. Shepard pers. comm.); 40/5.7, Rew Down (54.76) last recorded in 1966, now thought to be extinct due to scrub colonisation (X); 40/5.8, Ashey Down (57.87) last seen in 1966 since which time there has been a change in the grazing pattern (PX). Recorded from some 12 chalk pasture sites on the island since about 1820, but reputedly known at very few to any one generation of botanists (B. Shepard pers. comm.). Hants., v.c. 11: 40/2.9, West Lymington in 1883 (X); 41/0.1, Bokerley Ditch, Martin Down (04.18) few plants on the side of an ancient ditch (B. G. Tattersall pers. comm.) (B); also a second site nearby (A), and variant with white flowers (A. Hutchison pers. comm.); 41/3.2, near Bossington (32.28), chalk alluvial water meadow, late-flowering variant occurring with Gymna- denia conopsea vat. densiflora in atypical habitat for the species, 14 spikes in 1956, last recorded in 1962, but site planted with Populus sp. in c. 1970 (R. P. Bowman pers. comm.) (PX); Romsey area, last known in 1883 (X); 41/4.2, Compton Down (prob. 46.24) recorded last in 1878 (X); 41/ 6.2, Pink’s Hill, Warnford (60.22) in 1848-SO (X). N. Hants., v.c. 12: 41/2.4, Tidworth Golf Links (20.48), small population (Lady A. B. M. Brewis pers. comm.) (A); Kimpton Downs (24.46), ploughed in 1870 (X); 41/3.3 Chattis Hill (32.36) last recorded in 1936 (X); Danebury Hill (32.36) not recorded from 1964 to 1985, and thought to have become extinct due to undergrazing following the demise of the rabbit population until rediscovered in 1986 by P. Wilson (A. J. Byfield pers. comm.) (A); 41/4.3, The Gallups, Worthy Down (44.34) ploughed in 1952 (X); Flower Down (44.30) last recorded in 1799, subsequently built upon (X); 41.45, Woodcott Down (44.56) ploughed after 1892 (X); Litchfield Down (44.54) recorded in 1892 but ploughed at a later date (X); Ladle Hill around ancient earthwork but often heavily grazed, late-flowering variant (C); near Old Burghclere, transplanted here in 1985 from a nearby threatened area, but has still not flowered (P. Brough pers. comm.) (A); Ashley Warren, late-flowering variant, recently threatened by motor-cycle scrambling and undergrazing (C); Watership Down, north of Gallups (49.56), late-flowering variant, two plants in 1988 (A); 41/5.4, ORCHIS USTULATA IN SOUTHERN ENGLAND 45 Downs near Micheldever Station (50.42) last recorded in 1873 and later ploughed (X); near Southwood Farm (58.46) in 1859 and later ploughed (X); 41/5.5, near Wolverton Farm (54.56) in 1870, ploughed (X); White Hill, east-facing slope, discovered here recently, late-flowering variant (P. Brough pers. comm.) (A). W. Sussex, v.c. 13: 41/7.1, Harting 1889 (Arnold 1907) (X); 51/2.0, Portslade, Southwick (Arnold 1907), since built upon (X); 51/2.1, Newtimber Hill, Poynings, refound in 1983 by D. Batchelor when 33 plants, nine plants in 1984, 52 plants in 1985 (M. Briggs, D. C. Lang pers. comm.) (C); this area Newtimber/Saddlescombe/Pyecombe (Arnold 1907; Wolley-Dod 1937) could have been the same formerly more extensive colony (X); Henfield (Arnold 1887) (X). (The locality of a record for ““Summersdean’’, E. Payne (Arnold 1887) has not been traced (X)). E. Sussex, v.c. 14: 50/5.9, known from nine tetrads (Hall 1980), and scattered frequently in the pastures near Beachy Head (Arnold 1907), including Seven Sisters (52.98), 15 plants in 1985 (B); and same area (52.96) (Hall 1980) (U); Cuckmere Haven (50.96) (Hall 1980) (U); Kiln Combe, East Dean (Bullock Down), a north-facing, undergrazed chalk slope (B. G. Tattersall pers. comm) (D); also near East Dean (54.96) (Hall 1980) (U); near Friston (54.98) (Hall 1980) (U); Cow Gap (58.96), (Arnold 1907), ten plants in 1967 (B); downs above Meads (58.98) (Arnold 1907) (U); Whitbread Hollow (58.94) small colony (B); 51/3.0, Castle Hill, Falmer, south-facing downland with several colonies of varying size (B. G. Tattersall pers. comm.) but in total (D); Telscombe Tye (38.00) formerly (B) but destroyed c. 1970 (D. C. Lang pers. comm.) (X); 51/3.1, Standean, Patcham (30.10) formerly (A) but now destroyed (D. C. Lang pers. comm.) (X); 51/ 4.0, Mount Caburn, Glynde, 1000+ plants in a good year (B. G. Tattersall pers. comm.), short grazed turf on and around an ancient earth fort (F); also at (44.08) (U); near Firle Beacon (48.06) (Hall 1980) (U); 51/4.1, Cliffe Hill (43.10) (Arnold 1907) (U); Saxon Down, Glyndebourne (44.10) (A); 51/5.0, Cradle Hill, High and Over (50.00) 20 plants in 1972 (B); The Rails, Alfriston (50.02) four plants in 1965 (A); east of Alfriston (52.02) (Hall 1980) (U); Charleston Bottom (52.00) two plants in 1973 (A); Lullington Heath (54.02) (A), but 50 plants in 1981; Jevington (56.00) recorded in 1979 (A); Meachants, Willingdon, late-flowering variant, often 200+ plants (E) (B. G. Tattersall pers. comm.) and merging with a second colony of c. 15 early-flowering plants on a north-facing chalk slope (Combe Hill) (B. G. Tattersall pers. comm.) (B); 51/9.1, Rye Harbour (Hall 1980) is considered to be an error (M. Briggs pers. comm.). (Some tetrads listed by Hall (1980) in 50/5.9 are now possibly extinct, due to public pressure on the Downs.) E. Kent, v.c. 15: 51/7.6, Bluebell Hill, prior to 1900 (X), some with white flowers (Hanbury & Marshall 1899); 51/8.6, Queen Down Warren, south-facing chalk downland, very small colony with Ophrys sphegodes, Aceras anthropophorum, and Orchis morio (B. G. Tattersall pers. comm.) (A); 61/0.4, Wye Downs (06.44) (A); 61/0.6, Badgen Downs, near Faversham (Hanbury & Marshall 1899) (X); 61/1.3 (or 2.3), west of Folkstone, downland c. 1890 (X); 61/1.5, near Canterbury (Hanbury & Marshall 1899) (X); 61/2.4 Warren Bottom, two small colonies (24.44) with Ophrys sphegodes and Orchis morio (B. G. Tattersall pers. comm.) (A, B); Lydden, three small colonies (26.44) (B. G. Tattersall pers. comm.) (A,A,A); 61/2.5, near Wingham (Hanbury & Marshall 1899) (X); 61/3.4, St Margaret’s at Cliffe in 1901 (X); Langdon Bay (34.42) recorded in 1954 and surviving for perhaps a further ten years (E. G. Philp pers. comm.) (X); downs near Dover, one specimen with white flowers (Hanbury & Marshall 1899) (X). W. Kent, v.c. 16: 51/5.5, between Knockholt and Wrotham, several colonies, last recorded in 1845 (X); 51/5.7, Dartford, recorded in 1821 (X) (E. G. Philp pers. comm.); 51/6.7, Gravesend (Hanbury & Marshall 1899) (X). Surrey, v.c. 17: 41/9.4, Hog’s Back, Guildford, last recorded in 1874 (X); 51/1.4, White Downs, reported in 1976 but not substantiated (Lousley 1976), also very old record for this or the adjacent 10-km square (specimen in BON collected by A. Halley) for Dorking area in 1841 (PX); 51/2.5, Buckland Hills (22.52), small colony in 1946 (X); Walton Downs (22.57) known here since 1927, one plant seen in 1966 and now thought to be lost due to scrub colonization (X); Gatton Park (26.52), last recorded in 1965 (X); Betchworth recorded in 1884 (X); Wingate Hill in 1965 (Lousley 1976) (PX); Juniper Hill, Reigate Hill and Headley Hill, three 19th century sites (Salmon 1863) (X,X,X). N. Essex, v.c. 19: 52/4.4, near Strethall, possibly on the strip lynchets of Coploe Hill in about 1860, lost probably due to arable cultivation (Gibson 1862; Jermyn 1974; K. J. Adams pers. comm.) (X); 52/8.3 (or 8.4), Sudbury. Recorded here by G. S. Gibson in the last century (Gibson 1862), 46 Moy. Yo FOLEY. but some doubt as to whether the site might be actually in Suffolk. Lost probably due to arable development (K. J. Adams pers. comm.) (X). Herts., v.c. 20: 42/9.1, Tring (92.10) recorded in 1815, probably lost due to changing agricultural practices (X); near Tring Station (94.12) in 1883, changes in agriculture and scrub colonisation causing loss) (X); Aldbury (96.12) in 1887 lost due to scrub colonisation (X); 52/0.1, near Isle of Wight Farm, Kensworth (00.18) in 1887, lost to agriculture (X); 52/0.2, Ravensthorpe Castle (08.28) last record 1924, subsequently afforested (X); 52/1.3, High Down (13.30) in 1843 lost to agricultural changes (X); Bury Mead (18.30) recorded in 1843, site now built upon (X); 52/2.3, Arbury Banks (26.38) in small numbers up to the late 19th century, when agricultural methods changed (X); 52/3.3, Therfield Heath, two sites (32.38) — last record here was of a single plant in 1979 (B. Sawford pers. comm.) (A,PX), and also 52/3.4, scattered sparingly in the 1880s but lost with the cessation of grazing (34.40) (X). Middlesex, v.c. 21: 51/0.9, Harefield, pre-1737 record from a chalk pit — J. Blackstone 1737 (X). Berks., v.c. 22: 41/2.8, Kingstone Down and Whitehorse Hill, formerly two sites but no records traced since 1890 and 1964 respectively (X,PX), (but see Steel & Creed (1982) for records for this 10-km square 1970-82) and may survive in small numbers at the latter; 41/3.8, Lambourn Downs in 1838 (X); Wantage, last record in 1859 (X); 41/3.9, Cherbury Camp in 1867 (X); 41/4.8, Gore Hill, last record 1966 (PX); East Hendred in 1961 (PX); Farnborough Downs (Bowen 1968) (PX), (but see Steel & Creed (1982) for records for this 10-km square 1970-82), may still survive in small numbers; 41/5.8, Aston Upthorpe Downs, a nature reserve especially managed for the species (H. J. M. Bowen, B. G. Tattersall, pers. comms.) (B); Blewburton Hill last record 1964 (PX); Streatley in 1950 (PX); Ilsley in 1920 (X); Churn in 1930 (X); Moulsford Downs (Bowen 1968) (PX); 41/6.7, Basildon last recorded in 1879 (X); 42/4.0, Chilswell Hill pre-1820 (X); Botley pre-1820 (X). Oxon., v.c. 23: 42/3.0, Bampton, alluvial meadow in 1930, recorded by P. G. Beak (X); 42/4.1, Yarnton, occasional plant recorded here in the 1970s, possibly still surviving in very small numbers but no recent confirmation (H. J. Killick pers. comm.) (PX). Bucks., v.c. 24: 41/7.8, Hambleden (78.86) prior to 1926 (Druce 1926) (X); 42/6.0, near Albury in 1935, recorded by J. Chapple (X); 42/8.0, Coombe Hill (88.08), last recorded in 1961 (X); 42/9.1, Pitstone Hill (94.14) in 1936 (X); Ivinghoe (96.16) prior to 1926 (Druce 1926) (X). The plant has not been recorded during field work for The flora of Buckinghamshire. E. Suffolk, v.c. 25: 62/3.8, Bungay, specimen in BON pre-1903 (X) — this however could be for E. Norfolk (v.c. 27) in 62/3.9, possibly from Bath Hills. The Shelland and Hadleigh records (see below) given under v.c. 26, may in fact be from E. Suffolk. W. Suffolk, v.c. 26: 52/7.5 (or 7.6), Dalham, last recorded pre-1889, by F. Tearle (Hind 1889) (X); 52/7.6 (or 7.7), Cavenham in 1773, by Sir J. Cullum (Hind 1889) (X); 52/7.6 (or 8.6), Risby Heath, chalk bank in c. 1860 (Hind 1889), also recorded here in 1939 by J. E. Lousley & E. C. Wallace (X); 52/9.5 (or 9.6) or 62/0.5 (or 0.6), Shelland, record in East Anglia Daily Times, F. Woolnough, 1921 (X); 62/0.4, near Hadleigh in 1961, Sir C. Morris (X). (Also 52/6.6, Newmarket Heath in c. 1828 (Hind 1889), but this site may be same as that for Cambs., v.c. 29 (X)). W. Norfolk, v.c. 28: 53/6.0 (or 6.1), Shouldham, old 18th century record by R. Formby (X). No other records (C. P. Petch pers. comm.). Cambs., v.c. 29: 52/4.4, Ickleton before 1860 (X); 52/4.5, Cherry Hinton chalk pit (48.55) last recorded c. 1880 (X); Gog Magog Hills in 1863 (X); between Stapleford and Babraham, last record pre-1860 (X); 52/5.4 near Hildersham before 1860 (X); Linton in 1832 (X); 52/5.5, Balsham Heath pre-1870 (X); Fleam Dyke (53.55) last recorded in 1941 (X); 52/5.6, Devil’s Ditch in 1907 (X); 52/6.6, Devil’s Ditch (62.60) in 1955 (XK); Newmarket Heath pre-1889 (X); Chippenham before 1820 (X). Main causes of extinction in v.c. 29 are thought to be a combination of ploughing after enclosure of grass heaths in the 19th century, and lack of grazing of the remaining chalk grasslands. Beds., v.c. 30: 52/0.2, Ravensburgh Castle (08.28) last recorded in c. 1930 at which time a large heap of manure was deposited on the colony from which it never recovered — site now ploughed (X); Streatley, Sundon, Sharpenhoe, Luton Downs, Warden Hills, Galley Hill (Dony 1953) (X,X,X,X,X,X); 52/0.3, Barton in the Clay, two colonies, one of which at (08.30), was last known c. 1935 subsequently lost to scrub colonisation (X,X); 52/1.3, Knocking Hoe N.N.R., small ORCHIS USTULATA IN SOUTHERN ENGLAND 47 colony of usually 6/20 plants (T.C.E. Wells pers. comm.) (B). Apparently the decline began in c. 1930 when sheep grazing of the downland ceased. Hunts., v.c. 31: 52/0.9, Stibbington (06.98 or 08.98) record based upon a specimen collected in 1890 by Cpt. Vipan and sent to the Marchioness of Huntly who recorded this in her journal (T. C. E. Wells pers. comm.) (X). Northants., v.c. 32: 52/0.8, Biggin (00.88) on chalk drift pre-1930 and thought to have been subsequently ploughed (X); 53/0.0, Barnack (07.04) last recorded c. 1900 by Rev. H. Slater (X); Sutton Heath (08.00) in 1926 by Miss Powell (X); Southorpe (08.02) on rough grass near railway bridge in 1956 by H. F. Tebbs (X) (Biological Records Centre record), also Rigard (1957). E. Gloucs., v.c. 33: 31/9.9, Jackaments Bottom (95.97) small numbers recorded until the 1950s, subsequently lost to agricultural ‘improvement’ (X); 32/9.0, Edgworth, Miserden, Duntisbourne, Cirencester Park area (Riddelsdell et al. 1948) (X,X,X,X); 32/9.1, Brimpsfield to Elkstone (Riddelsdell et al. 1948) (X); Leckhampton Hill in 1862, specimen in CHELB (X); Crickley Hill, scarce in 1914 (Riddelsdell et al. 1948) (X); 32/9.2, Cleeve Common, (X); Prestbury (X) (both Riddelsdell et al. 1948); 42/0.0, Barnsley Downs (Riddelsdell et al. 1948) (X); Ablington Downs (09.08) last record c. 1948 when it was abundant. Site ‘improved’ agriculturally with small remnants which may still hold a few plants (PX); 42/0.2, Sevenhampton (Riddelsdell et al. 1948) (X); 42/0.3 Snowshill (Riddelsdell et al. 1948) (X); 42/1.0, Macaroni Downs (18.07) scattered on both sides of a dry valley, c. 30 plants in 1977, but recently ‘improved’ (X); Fairford (Riddlesdell et al. 1948) (X); 42/1.1, Northleach (Riddelsdell et al. 1948) (X); 42/1.2 Kineton Thorns (11.26) ploughed in 1956 (X); Notgrove and Bourton area (Riddelsdell et al. 1948) (X); 42/1.3, Hornsleasow Rough S.S.S.I. scattered widely over the site, which is on oolitic limestone at the remains of old quarry workings with Orchis mascula and O. morio, 300+ plants in 1965 but now much less (S. C. Holland, B. G. Tattersall pers. comms.), 25+ plants in 1988 (C); Bourton Downs (14.31) ploughed and reseeded c. 1948 (X). W. Gloucs., v.c. 34: 31/5.7, Durdham Down (Riddelsdell et al. 1948) (X); 31/7.7, Wyck Cliffs (Wick Rocks) (Riddelsdell et al. 1948) (X); 31/9.9, Poole Keynes, Rodmarton and Culkerton pastures, very abundant in 1924 (Riddelsdell et al. 1948) (X,X,X): 32/5.1, Symonds Yat, herb E.C. Townsend (X); 32/6.0, near Lydney in 1773 (Riddelsdell et al. 1948) (X). 32/8.0, Minchinhampton (Riddelsdell et al. 1948) (X). Herefs., v.c. 36: 32/5.1, between Rocklands and Coldwell at the foot of Coppett Hill (56.16) in 1849 (X); 32/7.4, Colwall, meadow, prior to 1889 (X). Worcs., v.c. 37: 32/7.4, Mathon (74.44 or 74.46) recorded here in the 19th Century (Amphlett & Rea 1909) (X); West Malvern (76.46) (Amphlett & Rea 1909), site lost early this century to building (X); 32/7.6, Abberley Hill (74.64 or 74.66) in the 19th century (Amphlett & Rea 1909) (X); near Witley (Amphlett & Rea 1909), lost in the 19th century (X); 32/9.3 (or 9.4), Bredon Hill, unconfirmed report in the 1920s (X) (J. J. Day pers. comm). Staffs., v.c. 39: 32/8.8, Kingswinford, about 1820 (Edees 1972) (X); 43/0.4, Weaver Hills, post-1900 (Edees 1972) (also old specimen in BON, pre-1903) (X). Salop, v.c. 40: 32/4.7, Downton Gorge (X); 32/5.7, The Lodge, Ludlow (X); 32/5.8, Upper Millichope (X); 32/5.9, Rowley, site drained in 1904 (X); 32/7.8, The Woodland, Bridgnorth (X); 33/5.0, Harley last recorded in the 19th century (X). No records this century (Sinker et al. , 1985). S. Lincs., v.c. 53: 43/9.2, Easton (X); 43/9.4, Court Leys in 1893 (X); Wilsford (X); Brandon (X); Ancaster Valley, newly discovered site in 1986 (A); 43/9.5, Navenby (X); Wellingore (X); 53/0.1, Bowthorpe in 1836 (X); 53/0.3, Sapperton in 1901 (X); 53/0.5, Temple Bruer (X). Leics., v.c. 55: 43/5.0, near Glenfield, not recorded since c. 1795 and subsequently urbanized (X); 43/5.2, Loughborough/Zouch Mill area recorded in c. 1831 (X) (A. L. Primavesi pers. comm.). Some doubt surrounds the above records since they are not on known calcareous soils. Rutland, v.c. 55b: 43/9.0, Shacklewell Hollow in c. 1875 (X); Ketton Heath, old record, area subsequently quarried for limestone (X); 43/9.1, Exton Park, possibly 1900-15, but later became an Opencast mine site (X) (K. G. Messenger pers. comm.). 48 M. J. ¥. FOLEY ACKNOWLEDGMENTS I would like to thank all those who have helped with records and information, especially many of the B.S.B.I. vice-county recorders of southern England, and also P. Brough, A. J. Byfield, K. Payne and B. G. Tattersall. REFERENCES AMPHLETT, J. & Rea, C. (1909). The botany of Worcestershire. Birmingham. ARNOLD, F. H. (1887). Flora of Sussex. London. ARNOLD, F. H. (1907). Flora of Sussex, 2nd ed. London. Bowen, H. J. M. (1968). The flora of Berkshire. Oxford. Dony, J. G. (1953). Flora of Bedfordshire. Luton. Druce, G. C. (1926). The flora of Buckinghamshire. Arbroath. Epes, E. S. (1972). Flora of Staffordshire, Newton Abbot. . ExstaM, U., JAcoBson, R., Matrson, M. & Porsng, T. (1984). Olands och Gotlands vaxtvarld. Stockholm. Foey, M. J. Y. (1987). The current distribution and abundance of Orchis ustulata L. in northern England. Watsonia 16: 409-415. Grsson, G. S. (1862). Flora of Essex. London. Grose, D. (1957). The flora of Wiltshire. Devizes. Ha t, P. C. (1980). Sussex plant atlas. Brighton. Hansury, J. H. & MARSHALL, E. S. (1899). Flora of Kent. London. Hinp, W. M. (1889). The flora of Suffolk. London. Hutcuincs, M. J. (1987). The population biology of the Early Spider Orchid, Ophrys sphegodes Mill. 1. A demographic study for 1975 to 1984. J. Ecol. 75: 711-727. IvimMEy-Cook, R. B. (1984). Atlas of the Devon flora. Exeter. JERMYN, S. T. (1974). Flora of Essex. Colchester. Lous_Ley, J. E. (1976). Flora of Surrey. Newton Abbot. MEUSEL, F., JAGER, E. & WEINERT, U. (1965). Vergleichende Chorologie der zentraleuropdischen Flora 1. Jena. RIDDELSDELL, H. J., HEDLEY, G. W. & Price, W. R. (1948). Flora of Gloucestershire. Cheltenham. RIGARD, F. (1957). J. Northants. nat. Hist. Soc. 1957: 33. Rog, R. G. B. (1981). The flora of Somerset. Taunton. SALMON, J. D. (1863). Flora of Surrey. London. SINKER, C. A., PACKHAM, J. R., TRUEMAN, I. C., OSWALD, P. H., PERRING, F. H. & PREsTwoop, W. V. (1985). Ecological Flora of the Shropshire region. Shrewsbury. STEARN, L. F. (1975). Supplement to the Flora of Wiltshire. Devizes. STEEL, D. T. & CREED, P. C. (1982). Wild orchids of Berkshire, Buckinghamshire, & Oxfordshire. Oxford. Wo .ey-Dop, A. H. (1937). The flora of Sussex. Bristol. (Accepted March 1989) Watsonia, 18, 49-62 (1990) 49 Introgressive hybridization between Crataegus monogyna Jacq. and C. laevigata (Poiret) DC. in the Upper Thames Valley, England A. G. GOSLER Edward Grey Institute of Field Ornithology, Department of Zoology, South Parks Road, Oxford, OX1 3PS ABSTRACT Where their ranges overlap in Britain, Crataegus laevigata (Poiret) DC. and C. monogyna Jacq. hybridize freely. The extent and causes of introgressive hybridization were investigated in the Upper Thames Valley by population sampling. Hybrid indices were calculated on the basis of leaf characters. A total of 1325 leaves from 265 trees in 17 populations was examined. The composition of Crataegus populations was strongly influenced by habitat, geology and management history. Evidence that coppicing reduced fecundity in the long-term was considered and this tended to encourage the introgression of monogyna genes into laevigata populations. With a reduction in the frequency of coppicing over the last 60 years, a gradual return to original taxon frequencies has occurred in a number of populations. It was suggested that differential insect grazing may have been responsible for the leaf character differences observed in the parental species. INTRODUCTION Two species of Crataegus occur wild in Britain. These are Crataegus laevigata (Poiret) DC. (the Midland Hawthorn) and C. monogyna Jacq. (the Common Hawthorn or May). These are generally accepted as good species and show a range of reliable, correlated characters (Franco 1968; Byatt 1975). Both species are shrubs or small trees growing to about 8 m, and commonly surviving to more than 100 years. Both are found widely in Europe where they are differentiated into a number of distinct geographical races. Those occurring in Britain are C. laevigata subsp. laevigata and C. monogyna subsp. nordica Franco. C. monogyna is a highly variable species in Europe and Browicz (1972) has suggested that interspecific hybridization has been partly responsible for this. Both species are diploid with 2n=2x=34. In Britain, C. laevigata is the rarer plant, occurring in 43/112 (or 38%) of vice-counties (Clapham et al., 1962) whilst C. monogyna is found in all British vice-counties. Under natural conditions, the two species are ecologically isolated. C. laevigata is typically a woodland plant with a strong preference for clay and loam soils. This more or less restricts it to the clay soils of the English midlands and Kentish Weald where it is surrounded by limestones such as chalk, oolite and carboniferous limestone. In contrast, C. monogyna prefers open habitats and is a dominant scrub species. It is rare only on very acid soils such as peat. Where their ranges overlap, these species regularly show introgressive hybridization resulting in a range of variants between the morphological extremes of the parental species. This ‘hybrid swarm’ results from successive crossing and back-crossing over several generations (Anderson 1949). Hybrids, i.e. plants which are intermediate in gross morphology between the parental morphs (Bradshaw 1971; Byatt 1975; Williams 1989), are also intermediate in leaf surface microanatomy, and in the occurrence of flavonoids (Gosler 1981). Byatt (1975) showed that C. laevigata occurred widely in the Kentish Weald but suggested that it had only crossed the chalk by introgression with C. monogyna. She suggested that introgression had been so widespread in that region that pure stands of C. monogyna could be found only on the chalk scarp slopes and that none of C. /aevigata existed. Williams (1989) has found that hybridization between the two species has also been extensive in remnant hedges in the London Borough of Brent. 50 A. G. GOSLER Anderson (1949) described a detailed genetic model of introgressive hybridization between two species. This identified three essential components. Firstly, the characters which allowed the recognition of introgression were controlled by polygenic complexes with a high degree of correspondence between the respective specific genomes. Secondly, there should be some ‘hybridization’ of the habitat which allowed a morphological transition to occur between the two species across an environmental gradient. The third factor was that which caused the initial contact between the two species. This was usually some environmental disturbance which might be natural (as described in Jris by Anderson) or might be due to human activity. Bradshaw (1953) showed that man’s influence was the most important factor causing syntopy in the British Crataegi. Anderson and his students worked mainly with annual and perennial herbs. These are relatively short-lived plants with a rapid life-cycle, but are frequently self-pollinated. Natural selection may act very rapidly in such a population so that significant changes in gene frequencies may be observed after only a few flowering seasons. In a small tree such as Crataegus which does not flower until it is 15 years old (or until 15 years have passed since coppicing) and may live 100 years, selection may act most strongly at a particular time in the life-cycle. The habitat may have little effect on an individual’s survival but may significantly reduce its flower production or fruit-set. Hence the habitat may induce a differential specific fecundity which determines the morphological direction to which the population will move in subsequent generations. Bradshaw (1971) showed that the British Crataegi were outbreeders showing between 30% and 59% fruit-set when cross-pollinated artificially, but only 2% fruit-set when selfed. Similarly, fruit- set was almost as high in interspecific crosses as in intraspecific ones. An examination of stained pollen showed no significant reduction of pollen stainability in hybrid plants, remaining above 95%. This established that full introgressive hybridization was genetically possible, and the need to outbreed greatly increased the probability of hybridization. Since Crataegus flowers are insect- pollinated (largely Hymenoptera and Diptera), vicinism (Grant 1971) plays an important part in producing the spatial distribution of variation observed in a population. Vicinism, longevity and differential fecundity may cause a departure from panmixis which may reduce the need for a smooth environmental gradient. Thus in Crataegus, the transition from open scrub habitats to closed woodland may involve too sudden a change in soil moisture, nutrients or available light to show a clinal change in morphology across the ecotone. This paper considers the nature, extent and causes of introgressive hybridization between C. laevigata and C. monogyna in the Upper Thames Valley by the detailed analysis of selected populations. Agriculture is the principal land use in the region and fields have traditionally been bounded by hedges. In the north, fields tend to be smaller, and here, hedges may be associated with boundary ditches and/or banks indicating the great age of the boundary (Rackham 1987). The area is geologically diverse and of comparable age to the wealden beds of Kent and Sussex — so providing an interesting comparison with Byatt’s (1975) survey of that region. Fig. 1 shows the drift geology and stratigraphy of the present study area (spanning some 135 million years). The soils reflect this geological series but are influenced also by the deposition of alluvium by the Thames and its tributaries. Little has been published on Crataegus hybridization in this area although the range of geology, and especially the close proximity of clays and limestones provides a unique opportunity for such a study. MATERIALS AND METHODS For present purposes, the Upper Thames Valley study area has been defined as that lying between northings 41/70 and 42/30 and between eastings 30 and 70 of the Ordnance Survey National Gnd. This area covers some 2400 km? and includes all of the Thames between Reading and Oxford. The work was carried out during June 1981 by which time the foliage was mature, most flowering had ceased and fruits were forming. Seventeen populations were selected for sampling, and some attempt was made to include C. laevigata populations cited by Bowen (1968). Site details are given in Table 1, and their locations are shown in Fig. 1. In most cases, specimens were collected along transects oriented to run across any apparent woodland/scrub gradient. When sampling hedges, specimens were taken at 5 m intervals. Two-year old twigs of up to 50 cm in length were cut from each of 15 (but see Table 1) trees in each HYBRIDIZATION IN CRATAEGUS 51 SU “ae N 1763360 70 SU70 “o Vo monogyna G))'='e i Clays % hybrid Nilay ee) Clay-with-Flints FE_]Limestones FicureE 1. Map of the Thames Valley study area, showing the drift geology (based on Holmes & Wilson 1960) and the location of the sampled Crataegus populations (numbered 1 to 17 as in Table 1). The pie diagrams indicate the proportion of each taxon in the population. The geological strata are labelled as follows: V alluvium, Ch Chalk, U Upper Greensand, G Gault, Lg Lower Greensand, P Portland beds, K Kimmeridge, Co Corallian, Oc Oxford clays, C Cornbrash, O Great Oolite, L Lias. 52 A. G. GOSLER TABLE 1. POPULATION DETAILS No. of Population name Gridref. n* MHabitats* Shrubs Description and notes 1 Brightwalton Common 41/422.806 15 HEW 7 Hedge along edge of Sycamore wood. 2 Ashampstead Green 41/561.773 15 HW 10 Old hedge along Beech wood. 3 Streatley Gold course 41/583.806 15 H 5 Open boundary hedge. 4 Cock’s Hill 41/662.818 10 HE 10 Old hedge by Beech wood. 5 Exlade Street 41/661.815 15 H 9 Old hedge by Pine plantation. 6 Highland Wood 41/690.788 15 HW 9 Roadside hedge by Beech wood. 7 Steventon Village Wood 41/469.912 15 HWE 8 Deciduous coppiced wood. 8 Stanford-in-the-Vale 41/325.953.. 15. -"HEW E 8 Hedge by wet wood on Corallian. 9 Appleton Lower Common 42/425.005 15 WHE 7 Ancient Oak wood. 10 Eaton Heath Wood 42/442.035 15 WH 5 Disturbed Oak wood. 11 Open Magdalen Wood-west 42/554.057 15 WHES 9 Scrub and Oak wood. 12 Open Magdalen Wood-east 42/557.058 14 WHES 9 Scrub and Oak wood. 13 Holton Wood 42/593.075 15 WHE 8 Coppiced wood. 14 Fernhill Wood 42/658.042 15 HE 4 Hedge running through wet wood. 15 Stratfield Brake 42/499.119 30 WHES 8 Scrub and ancient Oak wood. 16 Rushbeds Wood 42/665.158 15 WHE 11 Ancient Oak wood. 17 Glympton Park Estate 42/433.238 15 HW 10 Sycamore coppice. *n = sample size; Habitats: H hedge, E woodland edge, W wood, S scrub. See also Table 2. population. These were taken from the southern side of the tree and were selected on the presence of short vegetative shoots required for leaf examination. No attempt was made to collect flowers or fruit consistently as these were often too high to sample, although a measure of total flower and fruit production was recorded (see points 4 and 5 below). Hence the present analyses are based largely on vegetative (and especially leaf) characters. Specimens were taken from below 2:5 m high. Although sampling was random with respect to the types of plants collected the sample sizes dictate that some caution should still be exercised in the interpretation of results of ‘between-population’ analyses. In the field, the following data were collected: 1. Transect location. 2. A preliminary determination of identity (L, H or M). 3. Girth of the bole at 30 cm height in uncoppiced trees, or at ground level if coppiced. This gave an indirect indication of tree age. Ages were later grouped into year classes as 1) < 15, 2) 16-30, 3) 31-50, 4) 51-75, 5) 76-100, 6) >100. 4. Flowering was scored on a six point scale so that a non-flowering tree scored ‘0’, whilst a heavily flowering plant scored ‘5’. Although subjective, it was felt that reasonable relative estimates of flower production could be made which were comparable between trees. Fruiting was scored in the same way as flowering. . Tree habit was scored as (1) Coppiced bole, (2) Pollarded bole, (3) Natural (unmanaged bole). Style and/or pyrene number were recorded in all fruiting specimens. The habitat of each specimen was recorded as scrub, edge, hedge, woodland or woodland clearing. 9. A shrub count was made in hedges which formed part of the sampled population. The specimens were packed in presses and dried. In total, 265 specimens were collected. In the laboratory, five undamaged leaves were selected from short vegetative shoots on each specimen. Where possible the oldest leaves were taken. A total of 1320 leaves was measured. Where fruit was present, the presence of hairs on the hypanthium was recorded. Where stipules were present they were recorded as ‘monogyna-like’ or ‘laevigata-like’. The presence of variegated chlorosis indicative of iron deficiency was recorded. The amount of insect damage on the whole specimen was scored on a six-point scale (0-5, where 0 = all leaves complete, 5 = all leaves show some grazing damage). All other characters were scored on the five selected leaves. A hybrid index (I) was calculated for each specimen using the method described by Anderson (1949) where a WO NIAU HYBRIDIZATION IN CRATAEGUS 53 Crataegus laevigata Crataegus monogyna FicurE 2. Diagram showing the leaf characters (A—E, and 10-14) used in the calculation of the hybrid index. The range of variation between the two species is indicated by the two leaves. laevigata character state was scored as 0.0, and monogyna as 1-0. The full character list is as follows (see also Fig. 2): ils . Depth of lowest leaf lobe sinus (mm) (A in Fig. 2) . Lowest lobe length (mm) (B in Fig. 2) . Lowest lobe width (mm) (C in Fig. 2) — SOMNIDMAWN Fruit hypanthium villous or glabrous Length of leaf lamina (mm) (D in Fig. 2) . Length of petiole (mm) (E in Fig. 2) Ratio A/B . Ratio C/B . Ratio E/D . Hairs present in the vein axils of the abaxial lamina (I = 1-0) Hairs absent from the vein axils of the abaxial lamina (I = 0-0) . Number of leaf lobes (subtended by a primary vein) . Leaf veins curve upwards (I = 0.0) Leaf veins curve downwards (I = 1-0) 54 A. G. GOSLER 1-0 A Vv AIB be Q . #, v &X 05 wet 1 2 °O 0-5 ro /B “V “0 % * eae S Beas ts L, 5 Ficure 3. Scatter diagrams of the 17 populations showing ratio A/B plotted against C/B. See population 1 for details of axes. The following information is given for each tree: HABIT A coppiced, @ pollarded, V natural. FLOWER/FRUIT @ none, —@® flower with 1 style, @ 1 and 2 styles, @ 2 and 3 styles, @ flowers/fruit inaccessible. In plots 1-17, solid symbols denote a closed woodland habitat, open symbols an open habitat. The final diagram indicates the population means and their geology as D) Chalk and limestone, A Upper Greensand, and @ Clay communities. Means + 1 S.E. are shown. Note that the axes are not as in the other diagrams. 13. Leaf lobe apex: rounded (I = 0-0) : subobtuse (I = 0-5) : acute (I = 1-0) 14. Leaf margin: serrate (I = 0-0) : entire (I = 1-0) 15. Stipules if present /aevigata-like Stipules if present monogyna-like The hybrid index was calculated as the mean scores of characters 7, 10, 12, 13 and 14. These characters were chosen because, from previous studies, they were considered to be good diagnostic characters for the two species, and/or they could be assessed for every specimen, and because the index was to be used in another study (not reported here) to assess the value of the other characters HYBRIDIZATION IN CRATAEGUS 55 o Population oth means. a) i a at ):3 04 05 06 07 ere FIGURE 3. continued. in specific diagnosis. In the analyses that follow C. /aevigata is defined by a hybrid index (I) of <0-60, hybrids by I = 0-61-0-79 and C. monogyna by I >0-80. Statistical methods follow Sokal & Rohlf (1981) and terminology follows Stearn (1973). RESULTS AND DISCUSSION THE EXTENT OF HYBRIDIZATION IN THAMES VALLEY POPULATIONS Fig. 3 shows scatter diagrams for leaf width and indentation for each population as described by Anderson (1949). The same characters (7 and 8) have been used as were used by Byatt (1975) for comparative purposes. Each point on the diagram shows the position of an individual plant relative to the axes together with its habit, habitat and style number (believed to be one of the most reliable characters). A population with a mean scatter in the upper left-hand corner of the diagram consists largely of C. monogyna whilst the lower right-hand corner represents C. laevigata. The plots have been presented in approximate geographical order from south to north. From these it is seen that chalk and limestone populations are strongly biased in favour of C. monogyna with considerably less 56 A. G. GOSLER variation than populations containing C. /aevigata. This is borne out by the hybrid index values presented in Table 2. Limestone populations showed a mean hybnid index of 0-76+0-06 whilst that for clay populations was 0-57+0-07. This difference was highly signficant (t;;=6-03, p<0-001). Asin the Weald, there were no pure C. /aevigata populations although pure C. monogyna populations were frequent. C. laevigata populations showed a strong preference for clay soils but were considerably hybridized. These diagrams also show a clear habitat shift across the morphological range of the hybrid complex. Hence Crataegus populations in the region showed the same pattern of morphological variation and of geological and habitat preference that was found by Byatt in the Weald. Fig. 1 shows the proportion of each type in each population. The percentage of hybrids (as much as 40% in population 3 and 6) in chalkland populations suggests that both species were present here. Byatt (1975) found a sumilar situation in the Weald. She suggested that either hybridization had occurred between distant populations, or that these populations formerly contained C. Jaevigata and that introgression followed by strong directional selection had produced a monogyna-like population. For such a sudden change in selection pressure to operate there must have been considerable habitat change. It is possible that C. /aevigata was formerly restricted to the clay-with-flints deposits which locally cap the chalk. However, Byatt found no C. /aevigata in populations on the clay-with-flints, and it is unlikely that hybridization would occur between trees more than 30 metres apart (Grant 1971). An alternative hypothesis might be put forward. Most chalkland sampling was done in hedgerows because very little Crataegus could be found in the beechwoods of the region. Since these hedges were almost certainly planted, it is likely that cuttings had to be brought from the clay vales due to the scarcity of suitable shrubs in the local woodlands for taking cuttings. The lowland trees may have come from pure or hybridized C. laevigata populations, in which case gene flow was due to man. However, there is some evidence (though slight) from Ashampstead Green (pop. 2) that C. laevigata may be derived from populations on the clay-with-flints. This woodland population lies on that deposit and shows a significantly lower hybrid index than the surrounding hedge trees (Table 2). FACTORS AFFECTING FITNESS OF C. MONOGYNA AND C. LAEVIGATA I am here concerned with the factors affecting habitat preference and the causes of introgression in the Bntish Crataegi. Since fruit production in Crataegus gives a good measure of fitness I shall consider the effects of predation and habitat on fruit production as equivalent to their role in determining fitness although there are other components of fitness which have not been assessed. TABLE 2. MEAN HYBRID INDEX VALUES BY HABITATS FOR 17 CRATAEGUS POPULATIONS IN THE UPPER THAMES VALLEY Mean hybrid index t value between Population Substrate Woodland Hedge Scrub wood and hedge 1 limestone 0-89 0-84 _ 0-74 ns 2 clay & flint 0-43 0-79 — 4-04 p<0-01 3 limestone a 0-73 os a 4 limestone aS 0-84 —_ — 5 limestone — 0-77 a a 6 limestone 0-71 0-73 -= 0-25 ns 7 sandstone 0-56 0-42 -— 1-08 ns 8 limestone 0-83 0-75 = 1-18 ns 9 clay 0-65 0-88 — 2-78 p<0-05 10 clay 0-60 0-86 — 3-35 p<0-01 11 clay 0-44 0-42 0-87 0-14 ns 12 clay 0-55 0-89 0-88 40-20 p<0-001 3 clay 0-46 0-80 — 3-53 p<0-01 i4 clay 0-49 0-76 — 2-46 p<0-05 15 clay 0-43 0-55 0-81 1-01 ns 16 clay 0-47 0-89 — 3-86 p<0-01 17 limestone 0-74 0-58 — 1-31 ns HYBRIDIZATION IN CRATAEGUS Sy Predation here refers to insect grazing damage to leaves and flowers, rather than fruit loss to birds and mammals which is largely beneficial in dispersing seed. More than 100 invertebrate species are specifically associated with Crataegus in Britain (Pollard et al. 1974). Most of these are herbivores, feeding on fruits, flowers and leaves. Some, such as the hoverfly Episyrphus balteatus (Degeer), require pollen in their diet. The case-bearing caterpillar Coleophora coracipennella (Hubner) (Lepidoptera) also feeds on pollen. This moth larva burrows into young flower buds through one of the closed petals. Once inside, it feeds on the developing style, anthers and pollen. Coleophora may be detected in unopened buds by the small entry hole left near the base of one of the petals. A brief survey showed that as many as 20% of flowers on some C. laevigata in Open Magdalen Wood (pop. ref. 12) had been sterilised in this way. Predation was as high in some scrub C. monogyna trees in the same population. Pollard et al. (1974) suggested that woodland edge and scrub trees maintained a greater diversity of invertebrates than did woodland trees. If true, C. monogyna might be under stronger selection pressure to evolve resistance to predators than is C. laevigata. In the present survey, insect damage was often observed as cavities in the leaf lamina in the vein axil region. This was equally frequent in both species although plants with an open sinus tended to be less damaged. Insect eggs were frequently found in the axils of leaf veins and, again, an open leaf sinus appeared to reduce this. Hence it is possible that the open leaf sinus, axillary hairs and possibly the cuticular wax of C. monogyna are adaptations to reduce insect grazing. To test whether insect grazing could influence future gene frequencies in the population through an effect on fecundity, I plotted the mean fruiting scores of the 171 mature (more than 24 years old) trees against their scores of insect grazing damage (see Methods). The results are presented in Fig. 4. There is a strong negative correlation between insect damage and fruit score (T1469 = —0-342, P<0-001). Although this is highly suggestive, it is not clear whether this is a causal relationship nor indeed which might be the dependent variable. It is possible that the reduction in leaf area caused by selection for an open leaf sinus has reduced the ability of C. monogyna to colonise closed woodland habitats. Gosler (1981) found that for a given leaf length, C. monogyna leaves had a smaller lamina area and that the difference was greatest at the mean leaf length. Note also that the relationship is essentially similar in both the parental and hybrid morphs. However, although there is no significant difference in the mean grazing scores of the three morphs, the correlation was weaker in C. monogyna (tgs; = —0-286, p<0-01) than in either the hybrids (r45 = —0-466, p<0-001) or C. laevigata (135 = —0-407, p<0-05) and the slopes differed significantly (see Fig. 4B). This suggests that fruit production in C. monogyna was affected less by insect grazing than it was in the other taxa. This may help to explain the differences in fitness observed between the two species. Table 3 shows the mean fruiting scores for mature trees of each parental type and hybrids in open and closed habitats on clay soils. In woodland there was no significant difference in fruit production between the parental species. In open habitats fruit production was significantly poorer in C. laevigata then in C. monogyna. However, C. monogyna showed a significant reduction in fruit production in woodland compared with that in open habitats whilst C. /aevigata fruited significantly better in woodland than in open habitats. In both habitats hybrids were intermediate to the parental species in fruit production, although in open habitats C. monogyna trees were not significantly more productive than hybrids (t,g = 1-71, n.s.). We may therefore expect selection to favour introgression TABLE 3. FRUIT SCORES OF MATURE CRATAEGUS IN OPEN AND CLOSED HABITATS ON CLAY SOILS C. laevigata hybrids C. monogyna Mean fruiting score n=22 n=12 n=6 in woodland: 2:09+1-44 1-83+1-40 1-17+2-04 Difference between parental sp. means too = 1-04 ns. in hedges and scrub: n=5 n=7 n=14 0-5 +0-84 2:86+1-78 4-07+0-83 Difference between parental sp. means t17=8-18 p<0-001. t difference between 3-28 p<0-01 i351 n:s. 3-36 p<0-01 habitats. d.f.: 25 iy 18 58 A. G. GOSLER 8J0IS Buljingy ueay aJoIS Buljinuy ueay Grazing Score Figure 4. The relationship between insect damage (grazing) to leaves and fruiting score measured on 171 mature trees. A. shows the mean + 1 S.E. for each grazing score for all trees combined (@). The regression of fruiting score on insect grazing was highly significant (F, j69=21-93, p<0-001). In addition, the group means of populations on clays (i) and limestone ((C) are also shown. B. shows the relationship between fruiting score and grazing damage in the two species and their hybrids separately. Means + 1 S.E. are given for C. laevigata (O), for hybrids (®), and for C. monogyna (@). The three were defined according to their hybrid index values (see methods). The regression equations for each were also significant (F, 34 = 8-1, p<0-01; F,.45 = 12-23, p<0-01; Fi gs = 7:51, p<0-01 respectively). However, note that the slope for C. monogyna was significantly less than that of C. laevigata (t,2> = 5-01, p<0-01); see text. HYBRIDIZATION IN CRATAEGUS 59 in woodland habitats but C. monogyna in open habitats, hence the question remains of why woodland populations on clay are not totally hybridized. Evidence that C. laevigata regeneration is occurring comes from an examination of the age structures of the populations. Fig. 5 shows the age structure of the seventeen populations as kite diagrams. Each diagram shows the proportion of trees in each of the six age classes (see methods) together with the percentage of each type in each age class. Triangular diagrams with the apex at the top indicate a predominance of young plants in the population. These are mostly woodland populations with active regeneration. Triangular diagrams with the apex at the base show no regeneration and are largely hedge and scrub populations in which young plants are removed by grazing cattle or rabbits. The stable age distribution lies between these two extremes. These diagrams illustrate the changes that have taken place over time in the frequencies of the three groups in each population. Essentially four situations are suggested by these figures, although, as stated earlier, sample sizes dictate that some caution be exercised in the interpretation of these diagrams: 1. Populations showing approximately equal proportions of taxa with no changes over time, e.g. 6 and (to some extent) 15. 2. Populations in which hybridization has been extensive following the introduction of a second species, e.g. 3 and 7. 3. Woodland populations in which C. laevigata has been favoured over hybrids following introgression, e.g. 11, 13, and 16. 4. Scrub and hedge populations in which C. monogyna has been favoured over hybrids, e.g. 1 and 4. Some indication of when introgression began is given by the age of the oldest hybrids in a population. Table 4 shows the age of the oldest trees in each population. Although these were rarely hybrids, Table 4 suggests that hybridization was initiated in most populations more than 50 years ago. Since hybrids do not dominate woodland populations this suggests that other factors are operating. The differences in age structure described above have probably resulted largely from differences in habitat and management policy. In many woodlands, coppicing was discontinued more than fifty years ago. This might explain the small number of recently hybridized populations ir the area. Since coppicing affects the woodland trees more than those of the surrounding hedge or woodland edge, it has undoubtedly reduced the fecundity of C. laevigata relative to C. monogyna. Coppicing may have a long-lasting effect on trees. To assess the effect of coppicing on fruit production, fruit scores were standardized for tree age, the openness of the habitat, and insect damage using the regression equation: Fruit score = 0-323 + 0-0276 Age — 0-454 Insect + 0-669 Habitat This equation was derived from a stepwise multiple regression analysis in which these predictors each explained a significant proportion of the variation in fruit score (together with tree habit explaining 30-23% in all). The mean fruiting score for uncoppiced trees, after standardizing for age, habitat and grazing damage was 1-86+0-93 (n=159), significantly higher than that for coppiced and pollarded trees of 1-43+1-05 (n=104) (ty6;=3-33, p<0-001). However, this might be compensated for partly by the tendency for coppicing to increase the tree’s longevity (Rackham 1987). In the present study the mean age of coppiced boles in woodland was 52-:5+31-5 (n=78) which was TABLE 4. THE AGE OF THE OLDEST TREES IN CRATAEGUS POPULATIONS OF THE THAMES VALLEY Number of populations Age of oldest trees (yrs) ce SS (grouped into classes) C. laevigata hybrids C. monogyna 0-15 0 0 0 16-30 Z 0 0 31-50 3 + 6 51-75 3 6 2 76-100 Z 1 2 101+ 3 6 1 60 A. G. GOSLER FiGuRE 5. Kite diagrams showing the age structure of 17 Crataegus populations. Each diagram shows the percentage of all specimens in a given age class, and the percentage of each type in each age class. The final diagram is a key. See text for further details. significantly older than the mean for unmanaged woodland trees of 26-1+20-6 (n=33) (ti99=4-45, p<0.001). However, these results are biassed by the fact that coppicing was discontinued many years ago in most populations, so that the effect of coppicing on longevity cannot be assessed with confidence. Table 2 shows the mean hybrid index values for open and closed habitats in each population. In most woodland populations on non-calcareous soils there is a significant difference between index values in the wood and in the surrounding boundary hedge. However in populations 7, 11 and 15 HYBRIDIZATION IN CRATAEGUS 61 Rae ery 100+ “o C.monogyna 76-100 % hybrid relative SQ 25 0 abundance (%). FiIGuRE 5. continued. which are old woodlands, surrounded by ancient hedges with high shrub diversity suggesting great age (Pollard et al. 1974) and therefore probably planted from cuttings taken from within the wood, there was no significant difference and the low hybrid index values supports the view that C. laevigata is indicative of an old hedge (Bradshaw 1971). Most chalkland woodland populations have undoubtedly developed from hedges as they show no difference in morphology from the planted hedge populations. This is particularly well illustrated by the age structure of the Highland Wood population (6) in Fig. $. Here a young woodland population has grown up from seed from a nearby hedge (this may be assumed as the next nearest Crataegus source was over 1 km away) on the edge of a Beech wood. The frequencies of Crataegus taxa in the hedge and woodland were identical. 62 A. G. GOSLER CONCLUSION Crataegus laevigata and C. monogyna are adapted both anatomically and ecologically to distinct habitats but hybridize readily where their ranges overlap in the Upper Thames Valley. However, in many hybridizing populations, hybrids have not replaced the parental species totally. Two principal factors have contributed to the introgression of C. monogyna genes into C. laevigata populations in the Upper Thames Valley. These are extensive woodland coppicing, which has reduced the fecundity of the woodland C. /aevigata trees both in the short and longer term, and the planting of C. monogyna boundary hedges. The cessation of coppicing in most woodland populations has allowed some slow return to the original frequencies of the species in many woods. This contrasts with Byatt’s (1975) study of introgression in the Weald which predicted a general breakdown in recognisable specific boundaries in these two species. ACKNOWLEDGMENTS The work was carried out in part fulfillment of the M.Sc. degree in Pure and Applied Plant Taxonomy at the University of Reading. I thank the S.E.R.C. for funding this. I thank Prof. David Moore who supervised the project and Dr Stephen Jury for useful discussion and practical help. I am particularly grateful to Dr Alison MacDonald, Harvey Dunkley and an anonymous referee who greatly improved an earlier draft. REFERENCES ANDERSON, E. (1949). Introgressive hybridization. New York. Bowen, H. J. M. (1968). The flora of Berkshire. Reading. BrapDsHAw, A. D. (1953). Human influence on hybridization in Crataegus, in LousLey, J. E., ed. The changing flora of Britain, pp. 181-183. Oxford. BraDsHAW, A. D. (1971). The significance of Hawthorns, in Hedges and local history, pp. 20-29. London. Browicz, K. (1972). Crataegus L., in Davis, P. H., ed. Flora of Turkey 4: 133-147. Edinburgh. Byatt, J. I. (1975). Hybridization between Crataegus monogyna Jacq. and C. laevigata (Poiret) DC. in south- eastern England. Watsonia 10: 253-264. CLAPHAM, A. R., Tutin, T. G., & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed. Cambridge. FRANCO, J. (1968). Crataegus L., in TuTin, T. G. et al., eds. Flora Europaea 2: 73-77. Cambridge. Gos_er, A. G. (1981). Introgressive hybridization between Crataegus laevigata (Poiret) DC. and C. monogyna Jacq. in the Upper Thames Valley. M.Sc. thesis, University of Reading. GRANT, V. (1971). Plant speciation. New York. Homes, S. C. A., & Witson, V. (1960). In SHERLOCK, R. L. British regional geology: London and Thames Valley, 3rd ed. London. PoLitarD, E., Hooper, M. D., & Moore, N. W. (1974). Hedges. London. RACKHAM, O. (1987). The history of the countryside. London. SOKAL, R. R., & RHOLF, F. J: (1981). Biometry, 2nd ed. San Francisco. STEARN, W. T. (1973). Botanical Latin. Newton Abbot. WILLIAMS, L. R. (1989). Crataegus X media Bechst. in Middlesex hedgerows. Watsonia 17: 364-365. (Accepted April 1989) Watsonia, 18, 63-67 (1990) 63 Hypericum X desetangsii Lamotte nm. desetangsii in Yorkshire, with special reference to its spread along railways F. E. CRACKLES 143 Holmgarth Drive, Bellfield Avenue, Hull, HU&8 9DX ABSTRACT Records for Hypericum X desetangsii nm. desetangsii for Yorkshire are given, with an account of the distribution of the parental species, H. perforatum L. and H. maculatum Crantz, in the county. In Yorkshire, the hybrid has almost always been found in the absence of both parental species and only occasionally with H. perforatum L. The hybrid is very variable, particularly in the case of populations on railways sites. A hybrid index has been used to assess the degree of hybridity in individual plants. The distribution of the hybrid and its backcrosses is discussed. The introduction of the hybrid onto railway sites may have been followed by backcrossing with H. perforatum and in the course of time some taxa may have been lost. INTRODUCTION Prior to 1987, Hypericum X desetangsii Lamotte nm. desetangsii (H. maculatum Crantz subsp. obtusiusculum (Tourlet) Hayek X H. perforatum L.) was believed to be rare in Yorkshire, v.cc. 61— 65, having only been recorded ten times (Crackles 1986). Whilst staying in Ilkley, GR 44/11.47, in mid-May, 1987, I tentatively identified the hybrid by vegetative features at a quarry near Burton Leonard, GR 44/32.63, and on three sites adjacent to railway stations on the Leeds to Ilkley railway line. Identification of plants from these sites was confirmed in the flowering season, except for those at Burley-in-Wharfedale which were cut before they had a chance to flower. Interest having been aroused, additional localities for H. x desetangsii have been found in the Ripon area, GR 44/3.6 & 44/2.7, and along railways in v.cc. 61, 62 and 64 during 1987 and 1988. Details of these records are given below and in the forthcoming Flora of v.c. Ol: RECORDS N.E. YORKS., V.C. 62 Railway records: One plant in the disused Pilmoor Junction station and a continuous belt, stretching some 30 m, along the immediately adjacent main York to Edinburgh line GR 44/4.7, 1988, F. E. Crackles (F.E.C.) & J. E. Duncan (J.E.D.). See also the Yorkshire Naturalists’ Union’s annual report (1973). MID-W. YORKS., V.C. 64 Non-railway records: Low Grass Wood, Grassington GR 34/9.6, 1988, H. Lefevre (H.L.); open woodland, north-west of Ripon GR 44/2.7, 1987, D. E. Haythornthwaite (D.E.H.); quarry, Burton Leonard GR 44/3.6, 1987, F.E.C.; Quarry Moor, near Ripon GR 44/3.7, 1972, D.E.H., det. N.K.B. Robson, see also The Naturalist, p. 112 (1976). Railway records: disused line, near Threshfield GR 34/9.6, 1987, H.L.; Embsay station GR 44/0.5, 1987, J.E.D., det. N.K.B.R.; areas adjacent to Ilkley and Ben Rhydding stations GR 44/1.4, 1987, F.E.C. & J.E.D.; Guiseley station GR 44/1.4, 1987, J.E.D.; along the disused Harrogate to Ripon line near Bishop Monkton GR 44/ 3.6, 1987, D.E.H. and at Littlethorpe GR 44/3.6, 1988, F.E.C. & J.E.D.; in quantity by the main line between York station and Dringhouses sidings GR 44/5.4, 1981, T. F. Medd. 64 F. E. CRACKLES N.W. YORKS., V.C. 65 There are no railway records. By R. Swale, above Richmond GR 45/1.0, 1968, J. A. Gilleghan. Additional records are given in Sledge (1961) and The Naturalist 104: 70 (1979). DISTRIBUTION OF PARENTAL SPECIES Hypericum maculatum Crantz subsp. obtusiusculum (Tourlet) Hayek appears to have been always very uncommon in Yorkshire. Perring & Walters (1962) gave the species as having occurred in eight 10-km squares. Watson (1883) gave v.c. 61 in the list of vice-counties in which the species had occurred, but no details have ever come to light and there have been no later records. Baker (1863) stated that the species was ‘‘much less frequent” in N. Yorkshire than H. perforatum and cited seven localities, mainly in hilly country. Lees (1888) described the species as occurring in bushy places, mainly by water courses, “rare, although plentiful enough in certain areas”. He included a record for a stone quarry at Hutton, to the north-east of Ripon, GR 44/3.7, one of the 10-km squares in which the hybrid has been found in recent years. Since 1945, there have been nine Yorkshire records for H. maculatum, in Annual Botanical Reports in The Naturalist, one for v.c. 62, five for v.c. 63, two for v.c. 64 and one for v.c. 65, but none for a 10-km square for which the hybrid has been recorded, with the possible exception of the record for Langton-on-Swale, GR 44/2.9, v.c. 65 (Sledge 1961). Two additional 10-km square records for v.c. 65 are given in Perring & Sell (1968), the specimens in both cases having been examined by N. K. B. Robson. Hypericum perforatum L. is widely, but unevenly distributed in Yorkshire (Perring & Walters 1962), being common in many areas. OCCURRENCE OF THE HYBRID IN RELATION TO THAT OF THE PARENTAL SPECIES In most Yorkshire localities, the hybrid has not been found in the presence of both parents. H. perforatum was also present in five localities: Embsay station GR 44/0.5; woodlands north-west of Ripon GR 44/2.7; disused line, Bishop Monkton GR 44/3.6; Guiseley station GR 44/1.4 and Water Fulford GR 44/6.4 and was often introgressed. Only at Langton-on-Swale, GR 44/2.9, were both parents recorded as present in the same area, but the presence of H. maculatum was later questioned (Sledge 1961). The hybrid and H. maculatum are recorded for the same 10-km square only in the Ripon and Richmond areas (10-km squares 44/3.7 and 45/1.0). VARIABILITY OF THE HYBRID I have examined at least one specimen of the hybrid from each Yorkshire site where it has been found since 1986. The specimens, particularly from railway sites, are so variable that a hybrid index was developed to assess the degree of hybridity in each case. The method of scoring, using only those characters generally accepted as those of the F, hybrid (Robson 1981), is given in Table 1. A summary of the results of hybrid index analysis of specimens examined is given in Figs 1 & 2. Most plants of a population of H. X desentangsii on a roadside verge near Arnold in Holderness, v.c. 61, conform fully to the description of the F; hybrid as do populations on three railway sites. Populations on three sites in the Ripon area including that on the disused line near Bishop Monkton and cultivated plants grown in a garden near Knaresborough and which originated from Quarry Moor, near Ripon, are generally similar to each other and have a high score in the hybrid index analysis (score 6-8). However, the sepals instead of being blunt and apiculate are usually acute and attenuate, but nevertheless broad and erose-denticulate. In some such specimens, the occasional sepal may be blunt and apiculate. It should be noted in this connection that plants of the hybrid found near Langton-on-Swale in 1960 had lanceolate sepals which were pointed (Sledge 1961). First generation hybrids were found immediately adjacent to the platform at Guiseley station and on the same railway line near to Ilkley station. Near Guiseley station there was also a population of H. perforatum and another showing some signs of hybridity (score 4). Specimens from the Ilkley station car park (formerly the station yard) and the population immediately behind the platform at HYPERICUM Xx DESETANGSII IN YORKSHIRE 65 TABLE 1. METHOD OF SCORING THE HYBRID INDEX Hypericum maculatum Intermediate Hypericum perforatum Character Score 2 (or 1) Score 1 Score 0 Leaves No pellucid dots or a Lowest leaves with no All leaves with pellucid few on upper leaves pellucid dots dots only Densely reticulate Distinct, laxly reticulate Inconspicuous minor venation venation veins Stems Stem square for its full Signs of 2nd pair of lines Two raised lines only length or incomplete square- stemming Sepals Width Wide Intermediate Narrow Apex Obtuse and rounded Some sepals obtuse and Acute apiculate Margin near apex Erose-denticulate Slightly erose-denticu- Entire late Petals Black lines (score 1) Number of individuals BNO 1 2 3 a 5 6 af 8 One MON Ae W242 13 Hybrid index score Ficure 1. Histogram of the hybrid index scores for populations sampled on non-railway sites. Bed Rhyddiang station, 1-5 km west of Ilkley, were backcrosses (score 5—6). Most sepals are acute, occasional ones obtuse and erose, with sparse denticulation. N. K. B. Robson examined two specimens from Embsay station, on the former Skipton to Threshfield line, a branch of the Leeds to Ilkley line. Both specimens had numerous pellucid dots on their leaves. One specimen had “rather broad sepals with the apex apiculate and erose-denticulate’”’ whilst the second specimen had “‘acute entire sepals”. Both had “black glandular lines on the petals found in H. maculatum and more reticulate veins than is usual in H. perforatum’”’. These specimens had a hybrid index score of 6 and 3 respectively. Near Threshfield, on a disused section of the same line, Miss H. Lefevre found plants with few pellucid dots on the leaves and sepals which were narrow and acute, but erose and denticulate (score 6). Plants believed to be introgressed H. perforatum, with one or two hybrid features only, have been found with good intermediates at Bishop Monkton and at Embsay and Guiseley stations; they also occur in the absence of the hybrid in a gravel pit and on a waste place near Ilkley and on a disused 66 F, ‘E. CRACKLES Number of individuals No Hybrid index score FicureE 2. Histogram of the hybrid index scores for populations sampled on railway sites. railway line at Deighton, GR 44/6.4, v.c. 61. Introgressants in the Ilkley area are the only ones found on non-railway sites. DISCUSSION The widespread occurrence of Hypericum X desetangsii in Yorkshire almost invariably in the absence of one or both parents is of considerable interest. The occurrence of the hybrid in such circumstances is possible owing partly to a capacity for limited vegetative propagation and partly to its fertility (Robson 1975). The presence of the hybrid may indicate that both parental species once occurred in the area. Near Arnold in Holderness, GR 54/1.3, the hybrid occurs on a remarkable species-rich roadside where both calcicoles, calcifuges and species of both well-drained and wet soils are to be found, including Achillea ptarmica, Anemone nemorosa, Avenula pubescens, Carex spicata, Filipendula ulmaria, F. vulgaris, Lathyrus montanus, Lotus uliginosus, Ononis spinosa, Serratula tinctoria, Silaum silaus, Succisa pratensis and Trifolium medium. With the exception of the plants at Littlethorpe, the populations in the Ripon area (i.e. in 10-km grid squares 44/2.7, 44/3.6 and 44/3.7) are similar, suggesting a common origin. Plants on the stretch of the disused Harrogate to Ripon line near to Bishop Monkton appear to relate to the non-railway hybrid populations in the area. This stretch of disused line is being managed as a nature reserve, its vegetation being regarded as relict grassland of a type present in the district before the railway was constructed. Species present include: Avenula pubescens, Blackstonia perfoliata, Bromus erectus, Dactylorhiza fuchsii, Knautia arvensis, Origanum vulgare, Pimpinella major, P. saxifraga, Rhinanthus minor, Silaum silaus and Trifolium medium. H. maculatum is known to have occurred at some time near to both Ripon and Richmond, in 10-km grid squares 44/3.7 and 45/1.0 respectively. The hybrid tends to occupy habitats intermediate in wetness between those of the parents, which do, however, have overlapping habitat requirements. It is possible that there has been some loss of suitable habitats for H. maculatum in the course of time, whilst environmental changes may have favoured the survival of the hybrid. The presence of the hybrid on so many railway sites in districts where H. maculatum has never been found leads one to the conclusion that the hybrid has been introduced along the railways by air movement generated by trains or by other railway activity. Backcrossing from the hybrid occurs readily in both directions, thus producing a more or less continuous range of variants between the parental species (Robson 1981). Variants of the hybrid, other than F,s, found along railways in Yorkshire are consistent with these being backcrosses to H. perforatum. Introduced first generation hybrids may have come into contact with H. perforatum growing along the line and backcrossed with it, as seems to have happened at Guiseley station. HYPERICUM Xx DESETANGSII IN YORKSHIRE 67 Further away from the point of introduction, one might expect backcrossing to be more pronounced, as it is at both Embsay and Threshfield. Hybrids on disused lines must have been there since the days of the steam-train, indicating that the present situation has arisen over a long period of time during which taxa may have been lost. It is also possible that the initial introduction may have been a backcross or an introgressant. The fact that the Hypericum X desetangsii has been found in quantity along the York—Newcastle— Edinburgh line and by the main York—Doncaster—London line suggests that its spread along railways is unlikely to have been confined to Yorkshire. There are records for the hybrid for railways in other parts of Great Britain. It is significant that in ‘Plant Records’ in Watsonia, there are first vice-county records for disused railway lines in Montgomery, v.c. 47, and Merioneth, v.c. 48, both in 1971 and in Roxburghshire, v.c. 80, in 1975 and a second vice-county record at Crediton station in Kircudbrightshire, v.c. 73, in 1977. The hybrid is known to have occurred in close proximity to both parental species on the disused Waverley line (M.E. Braithwaite pers. comm.) at Acreknowe, GR 36/5.0, and at Longnewton, GR 36/5.2 (Braithwaite 1975). The Waverley line formerly connected Hawick with Edinburgh and was also linked to Carlisle and Newcastle. There is no doubt that H. x desetangsii has been overlooked in Yorkshire even on a nature reserve. Detection of backcrosses requires care. The examination of a single leaf is not enough. In some hybrid plants, median leaves may have many perforations and exceptionally all leaves may have pellucid dots, so that sepals should also be examined. It should also be remembered that all the plants of a population may not have the same combination of characters. _ May I suggest that botanists should examine all railway populations of St. John’s Worts critically and publish their observations and so help to throw further light on this intriguing story. ACKNOWLEDGMENTS I wish to thank Dr N. K. B. Robson for much useful discussion and valuable comments on my observations. I am also indebted to Mrs J. E. Duncan, Mrs D. E. Haythornthwaite and Miss H. Lefevre for providing specimens, and to T. F. Medd for providing information and copies of correspondence. REFERENCES BAKER, J. G. (1863). North Yorkshire — studies of its botany, geology, climate and physical geography. London. BraitHwalteE, M. E. (1975). A railway flora of Teviotdale. Hawick. CRACKLES, F. E. (1986). Botanical comment 4. How came they? Yorks. Nats. Union Bulletin, No. 6: 5—7. York. Legs, F. A. (1888). The flora of West Yorkshire. London. PERRING, F. H. & SELL, P. D., eds (1968). Critical supplement to the Atlas of the British flora. London. PERRING, F. H. & WALTERS, S. M., eds (1962). Atlas of the British flora. London. Rosson, N. K. B. (1975). Hypericum L. in Stace, C. A., ed. Hybridization and the flora of the British Isles, pp. 164-167. London. Rosson, N. K. B. (1981). Hypericum X desetangsii Lamotte in WIGGINTON, M. J. & GRAHAM, G. G. Guide to the identification of some of the more difficult vascular plant species. Banbury. SLEDGE, W. A. (1961). Hypericum X desetangsii Lamotte in Yorkshire. The Naturalist, pp. 43-44. Hull. Watson, H. C. (1883). Topographical botany, 2nd ed. London. (Accepted August 1989) @ reat ary —_ pes q at bse , ac = 7 I Ae § a “ ee eey wr wth @ Bris me he one ot pe eee ameeiy; ~~ af ‘eoiiniriio ty map Esch ti til PPraraci’ = ee a sie a »” oy Leaner oaizs¥ eal sooit si spb eh ed - gralp Reoene ent fed) amass onal f eo: a 2 4 ‘ : pride ol at ye} Sree: m ‘1 tieatiti iso > 1 eta pad a? in ass 10) i ata ie thi | | : d th . Lrerkat iM, ar retje ‘ alll avtt Oona yithameta Ate nye ie va ° ete) uh Rhye at yh rypey sy vals to mong? om emer 6) , Boe. ooh if . ae A Leagan ns Bi. ' neiia ? thet te *e% yy t f = “4 “sth en yy rena wits ~ C+ i, 5 ‘ bi : ayenl tad Th AT ats Wheat i : tee res) Jos Aiea) Sub iS) ears ESN eS ngra * Mesh a ail : r ° hs Cw i we 4 ¢ e a av > A . ae r “ty 4 + 4. >» a wo 4 a, ’ 6 ¢ > - aa 3 J J a we ‘ - Watsonia, 18, 153-172 (1990) 153 An assessment of populations of Dactylorhiza traunsteineri (Sauter) Soo in the British Isles and a comparison with others from Continental Europe hers. y. POLEY 87 Ribchester Road, Clayton-le-Dale, Blackburn, Lancs., BBI 9HT ABSTRACT The acceptance of Dactylorhiza traunsteineri (Sauter) Sod (Orchidaceae) as a British and Irish species has frequently been questioned. Twelve populations of dactylorchids from widely separate parts of the British Isles which have traditionally or recently been referred to this taxon, have been the subject of a critical morphometric evaluation. An inter-population comparison of the mean values of the key diagnostic characters indicated a degree of uniformity through the twelve populations although some geographical trends were identified. The data obtained were compared to, and found to be closely similar to those for populations of D. traunsteineri from the alpine region of Europe, including the type locality, and from Scandinavia, and this is offered as additional evidence for the inclusion of the species in the British and Irish floras. For D. traunsteineri in the British Isles, a comparison has been made with other dactylorchid taxa and their differences highlighted. The principal diagnostic characters of D. traunsteineri are summarised and a provisional distribution map is given. INTRODUCTION In a footnote to his Flora and under the name Orchis Traunsteineri, Reichenbach gave what was the first brief description of a species of marsh orchid from Kitzbihl, Austria, quoting Sauter as the authority (Reichenbach 1831). Shortly afterwards, Sauter himself provided a more detailed diagnosis of O. traunsteineri based upon similar Austrian specimens (Sauter 1837); in current nomenclature this is known as Dactylorhiza traunsteineri (Sauter) So6. Plants with characters corresponding to the description of continental D. traunsteineri were known in the British Isles as long ago as the last century, but were only first considered as such relatively recently (Heslop-Harrison 1953). Before and since then, however, their exact taxonomic status has been the subject of much speculation and various re-classifications have been attempted with treatments at varietal (Godfery 1933), subspecific (Pugsley 1936; Bateman & Denholm 1983) and specific level (in the latter case as Orchis Traunsteinerioides (Pugsley) Pugsley 1940). These treatments have been summarised by Roberts & Gilbert (1963), Tennant (1979), and Roberts (1988), and these workers along with others (Pugsley 1936; Heslop-Harrison 1953; Lacey 1955; Lacey & Roberts 1958) have also listed localities for similar dactylorchids. Morphometric studies of this taxon aimed at clarifying the situation have, in the British Isles, been restricted to the well- known Welsh populations (Cors Geirch, v.c. 49, and Cors Bodeilio, Rhos-y-gad, and Cors Erddreiniog (all v.c. 52)) (Lacey 1955; Lacey & Roberts 1958; Roberts 1966, 1988; Bateman & Denholm 1983), to one in Berkshire, v.c. 22 (Cothill), and also to three in Ireland, v.cc. H19, H20 and H23 (Heslop-Harrison 1953), and two small Yorkshire populations, v.cc. 62 and 64 (Roberts & Gilbert 1963) both of which are now almost extinct. Each of these independent studies has therefore covered only a very limited geographical range. Also, in addition to these differing views on taxonomic status, controversy has also arisen (Bateman & Denholm 1989; Roberts 1989) over the merits of the morphometric techniques employed, the interpretation of the data, and possible associated errors. Within recent years, further populations referred to this taxon have been located in the British Isles, including several in Yorkshire, v.cc. 62 and 65 (Tennant et al. 1983; Foley 1986, 1989; Bolton & Horsman 1987; Tennant 1987), Norfolk, v.cc. 27 and 28 (Petch & Swann 1968; Swann 1975; F. Rose, pers. comm. 1988), Ireland, v.cc. H4, HS, H9 H16, H17, H18, H22, H26 and H29 (Scannell 154 MOI. Y¥. -FOREY 1973; Webb & Scannell 1983; Curtis & McGough 1988; M. Keane, pers. comm. 1989), northern Wales, v.c. 49 (Roberts & Ward 1978), and scattered localities in southern England, v.cc. 6 and 12 (Rose 1975; Hedley 1987). Only very recently however, has it been confirmed from Scotland; Cunningham & Kenneth (1979) first tentatively identified as D. traunsteineri, a population of marsh- orchids from Kintyre, v.c. 101, but this was later refuted (Tennant & Kenneth 1983) when the plants were reassigned to D. majalis (Reichenb.) P. F. Hunt & Summerhayes subsp. occidentalis (Pugsl.) Sell. It was not until the discovery of a population very similar to Orchis francis-drucei Wilmott in W. Ross, v.c. 105, and its subsequent identification as D. traunsteineri (Lowe et al. 1986) that it was finally recognised from Scotland, although its presence there had been suspected for many years. Since then other: populations have come to light (Kenneth et al. 1988), mostly very small and including some in Kintyre, but one larger Scottish population from W. Ross is known and is included in this study. In a recent morphometric evaluation of British and Irish tetraploid marsh-orchids, which included two populations of this taxon from Anglesey, Bateman & Denholm (1983) assigned such plants to the new combination D. majalis subsp. traunsteinerioides (Pugsley) Bateman & Denholm. Roberts (1988), however, has shown that morphological data from one of these Anglesey populations compare well with those obtained by Reinhard (1985) for D. traunsteineri from the alpine region of continental Europe, and that separation of the Anglesey plants from these on the basis suggested by Bateman & Denholm (1983) cannot be justified. The present study, independently instigated, was already in progress prior to the publication of the work of Roberts (1988) and was undertaken in order to assess the degree of similarity or otherwise of various populations representing the full geographical range of the taxon in the Bnitish Isles. A morphometric examination of randomly selected plants has been made for each population, and data obtained compared with those for D. traunsteineri from the alpine region of Europe including the type locality. Comparisons have also been made with plants from Scandinavia, and with other British and Irish dactylorchids. This study therefore addresses two independent questions: (i) are British and Irish populations conspecific with continental D. traunsteineri, and (ii) should this taxon be considered at specific level or as a subspecies of some other dactylorchid, for example D. majalis? POPULATIONS SAMPLED A provisional map indicating the distribution of D. traunsteineri in the British Isles is given in Fig. 1. For the present study, evaluated populations have been selected as representative of the known range. Many of these have been discovered or identified only quite recently and have not been previously assessed morphometrically. Details of the twelve British and Irish populations studied are given in Table 1. Other than at Booton (F) and Killinaboy (K), where the number of plants is small (approx 50), all are relatively strong populations often comprising several hundred flowering plants. This is significant particularly in the case of the Yorkshire sites, since the only previous morphometric study (Roberts & Gilbert 1963) carried out on similar plants from this area, was on colonies of total size nine and 22 plants respectively. Populations examined also include three which have previously been studied morphometrically (Heslop-Harrison 1953; Roberts 1966, 1988; Bateman & Denholm 1983). These are in Berks (v.c. 22) at Cothill (G), and in Anglesey (v.c. 52) at Cors Erddreiniog (H) and at Rhos- y-gad (1), and again all comprise a relatively large number of plants. These twelve populations which are the subject of this study each occur in eutrophic, base-rich fens or flushes, some of which are quite small in area and are isolated. Schoenus nigricans L. is invariably present and grows in close association with these dactylorchids, whilst Valeriana dioica L., and where their distributional ranges allow, Pinguicula vulgaris L. and Primula farinosa L., are usually present also. Other associates can include Carex lepidocarpa Tausch, Menyanthes trifoliata L., Epipactis palustris (L.) Crantz, Listera ovata (L.) R. Br., and in Anglesey and Ireland, Ophrys insectifera L. Other dactylorchids are quite often absent, but Dactylorhiza incarnata (L.) S06 subsp. incarnata and subsp. pulchella (Druce) So6 sometimes occur nearby but not usually in direct association or intermixed with D. traunsteineri. D. majalis subsp. purpurella (T. & T. A. Steph.) Moore & So6 is a rarer associate, but subsp. praetermissa (Druce) Moore & So6 occurs at sites in southern and eastern England, whilst D. fuchsii (Druce) S06 and D. maculata (L.) S06 also occur MORPHOLOGY OF DACTYLORHIZA TRAUNSTEINERI 155 CriaraaEt ISL ANOS PLOTTED ON UIM GRID FicureE 1. The distribution of Dactylorhiza traunsteineri in the British Isles plotted on a 10-km square basis (provisional). @ recorded sites; © doubtful or unconfirmed records. within the general vicinity of most populations. Hybrids with other dactylorchids have been recorded by the author and others at several of the sites studied; the hybrid with D. maculata is known at Thornton Dale, Dalby Forest (Foley 1986), Cors Erddreiniog and Rhos-y-gad; that with D. fuchsii at the two Anglesey colonies; with D. incarnata at Wharfedale and again at the two Anglesey colonies; and with D. majalis subsp. praetermissa at the three sites in southern and eastern England. In all cases they are relatively infrequent and often very rare. No hybrids were found at Fylingdales although that with D. fuchsii may have been recorded here recently (Horsman 1989). In addition hybrids with the spotted orchids are suspected to occur at W. Ross, but no hybrids were 156 M. J. Y. FOLEY TABLE 1. BRITISH AND IRISH POPULATIONS OF DACTYLORHIZA TRAUNSTEINERI STUDIED Grid Number Population Habitat Reference* _ of plants First record (if known) Dalby Forest, Fen 44/8— 8— 100-200 Foley (1986) v.c. 62 (A) Thornton Dale, Base-rich 44/8— .8— 1000+ Originally thought to be D. majalis v.c. 62 (B) flushes subsp. purpurella until redetermined by Tennant, Wright & Wright (1983) Fylingdales, Base-rich 44/9—.9— 500+ Old record recently rediscovered by v.c. 62 (C) flush Bolton & Horsman (1987) Wharfedale, Base-rich 34/9—.6— 300-500 Recorded here by P. M. Hall & W. A. v.c. 64 (D) flush Sledge in 1934. Also Pugsley (1935) Beeston, Fen 63/165.424 500+ Discovered by F. Rose, see Petch & v.c. 27 (E) Swann (1968) Booton, Fen 63/1—.2— ec. 0 Discovered by F. Rose (F. Rose, pers. v.c. 27 (F) comm. 1988) Cothill, Fen 41/461.998 100+ Known here for many years. See also v.c. 22 (G) Heslop—Harmison (1953) Cors Erddreiniog, Fen 23/475.821 1000+ Lacey & Roberts (1958) v.c. 52 (H) Rhos-y-gad, Fen/ 23/510.788 100-200 Roberts (1960) v.c. 52 (I) meadow West Ross, Base-rich 18/7—.4— 500-1000 Discovered by M. R. Lowe in 1985 v.c. 105 (J) flushes Killinaboy, Fen 11/2—.9— c. 50 Not known (M. Keane, pers. comm. v.c. H9 (K) 1989) Kilmacduagh, Fen 11/3—.9— 100+ Known for some years; see Webb & v.c. H9 (L) Scannell (1983) * Owing to the fact some populations are in areas of very restricted access or are otherwise sensitive localities, full grid references are not always given. Full details are held by the author and by the Biological Records Centre. observed at Killinaboy or Kilmacduagh although D. majalis subsp. occidentalis occurs in the vicinity. In all cases where mixed populations of dactylorchids occurred there was no difficulty in assigning plants to the correct taxon, or in recognising hybrids as such. More than 85 localities have been recorded for D. traunsteineri in the British Isles, mostly restricted to calcareous fens and flushes. The plant exhibits a rather fragmented distributional pattern as can be seen from the provisional map (Fig. 1). However it has not been possible to examine specimens from all of the localities shown. Some populations occurring in Ireland have had their taxonomic status questioned (Webb & Scannell 1983) — as have many elsewhere — and it is not too clear how many Irish localities are still extant (Curtis & McGough 1988). In the British Isles plants are normally in full flower in late May in western localities including Anglesey, and by early/mid-June inland, at higher altitudes, and to the north. METHODS Morphological characters were recorded from ten randomly selected plants in each population. The characters selected (and method of measurement), were in accord with those of other workers (Bateman & Denholm 1983; Roberts & Gilbert 1963; Roberts 1966, 1988), namely, plant height, total number of all developed and developing leaves (excluding the basal scale-like leaf which often decays), length of fully open inflorescence (from apex to the junction of the lowest flower with the MORPHOLOGY OF DACTYLORHIZA TRAUNSTEINERI 157 stem), the number of flowers in the inflorescence, and the maximum dimensions of the second leaf above the stem base (which is also usually the largest leaf). These characters were all measured on field material. At the same time the degree of leaf spotting was assessed (0 = absent, 1 = present) and the extent/distribution and size of the spots noted; an estimate was also made of the degree of anthocyanin staining of the upper stem and floral bracts (0 = absent, 1 = light, 2 = heavy). A single flower was removed from below the mid-point of each inflorescence, and from it, the maximum labellum width, the labellum length from the apex of the central lobe to the spur opening, the depth of the deeper inter-lobe sinus (if different) measured parallel to the vertical axis through the labellum from the base of the sinus to a point level with the apex of the adjacent lateral lobe, and the spur length from its tip to the junction with the labellum as well as the spur width at this point (with spur detached), were all obtained from flattened specimens which had been pressed within 48 hours of sampling. The shape of the labellum and the type and distribution of markings upon it were noted on similarly flattened specimens and also in the field. The peripheral cells of a floral bract selected from below the mid-point of the inflorescence were also assessed for shape in profile and measured under microscopic examination (100). In order to overcome any variation in shape and size of individual cells within and between bracts from the same and different plants, the total length of ten contiguous peripheral cells was measured at a point approximating to the mid-length on five separate bracts each from different plants within the population. The mean length and range of the cell lengths was calculated and any characteristic cell shapes noted. Any other distinct differences between populations which were readily apparent in the field, were also recorded, and at some localities, pH measurements were made on samples of surface water taken from within the immediate vicinity of the plants. Results for most of the characters studied are recorded as population mean values, and where appropriate the standard error has also been calculated. From the above measurements two further characters, floral density (number of flowers/cm of inflorescence length), and mean leaf index (second leaf from the base of the stem, mean length/ mean (maximum) width) were calculated. All the data were obtained during the period 1985-1989. RESULTS AND DISCUSSION The main diagnostic characters of D. traunsteineri were given in some detail by Sauter (1837), viz: long narrow leaves, a lax, few-flowered inflorescence with relatively large flowers, each having a tri- lobed labellum. In addition other continental plants of D. traunsteineri have been described or illustrated as being typically slender, possessing relatively few, spotted or unspotted leaves, which are well-spaced along the stem, and with relatively broad labella (e.g. Landwehr 1977; Reinhard 1985; Kalteisen & Reinhard 1986; H. R. Reinhard, pers. comm. 1988). VARIATION WITHIN THE BRITISH AND IRISH POPULATIONS STUDIED For the populations studied, Table 2 lists data for the above-mentioned distinctive features. The mean length and shape of the peripheral cells of the floral bracts (character 19) are included in Table a. The average of the character means and the range which encompass the twelve British and Irish populations examined have been extracted from these data and are also given in Table 2 where it can be seen that there is close inter-population agreement in many key characters namely:— (a) Mean number of flowers per plant (character 3). There is close agreement between most populations, falling within a narrow range 7-0—9-0; the slightly higher values for some populations (E-L), especially those from Ireland (K and L), reflects the presence of more robust plants at these sites. (b) Mean floral density (character 4). Again there is good agreement throughout with little regional trend indicated. Inflorescences are variable in shape or very roughly cylindrical/tapering. (c) Mean total leaves per plant (character 5). No significant regional trends were apparent, with good inter-population agreement. The mean number of non-sheathing leaves (character 6) is often considered to be a partially diagnostic character of D. traunsteineri (range 0-1) since the value for other British dactylorchids is much higher. Here, where known, the range is 0-6-1-0. In all cases it 237 ¥. POLLEY 158 (§-0-0:0) (=) ) =) (=) (—) f—) (=) (—) (—) (—) (—) (—) sunods jeoy 7:0 8-0 1-0 0:0 0-0 0-0 1-0 0-0 0-0 S-0 0-0 1-0 €-0 Jo 90189q ‘O] (08-ZI-T€-L(6r-S)) (—) (—) (=) (=) (—) (=) (—) (—) (—) (—) (=) (—) gXOpul 9L°8 09:71 O8-ZI 6S pL:L 9€-L €1-6 06:6 85-0 €S-S 01-6 6S-L le-L yea] URI “6 (p1-1-€8-0) (80:0) (80-0) (0-0) (S00) (S00) (S0:0) (90-0) (€0-0) (90-0) (€0-0) (40-0) (€0-0) (Wid) yea] puodes Z0°1 ail 60:1 16-0 €0'T O11 00-1 10-1 €6:0 €0°1 €8-0 O11 pO'l jo yIpIm “xew URI °g (L€-r1-00°S) (LL:0) (€9:0) (Or-0) (6b-0) (8br-0) (89-0) (TZ:0) (95-0) (81-0) (O€-0) (O€-0) (9-0) (Wid) yea] puodes 96:8 Lev S6-€I 00°S L6:L O1l'8 €1-6 00-01 8-6 OL:S SSL S€-8 09-4 Jo yysus] URI “7 (0-1-9-0) (c1-0) (60-0) (21-0) (ST-0) (60-0) (00-0) (SI-0) (60-0) (€I-0) (SI-0) (SI-0) (60-0) SoAra] Suryyeoys 8-0 8-0 6-0 8-0 9-0 6:0 0-1 L-0 6:0 8-0 L:0 9-0 6-0 -uOU JO ‘OU URI “9 (0-b-€:€) (ST-0) (ST-0) (81-0) (ST-0) (STO) (ST-0) (91-0) (1-0) (2-0) (€I-0) (91-0) (ST-0) SdAB9] JO ‘OU 9-€ L€ E-€ C-€ L€ r-€ L€ CE 0-r 6-€ 8-¢ S-€ E-€ [210} URI “¢ (26:7-90:7) (—) =) (—) (=) =) om) (—) (—3) (—) (—) (=) (—) €r-7Z S77 69-7 0€-Z €£-Z 767 69-7 L0-Z Or-Z 69-7 €S-Z 97-7 90-7 pAUSUSP [RIO] “p (p-Z1-0:L) (9L'1) (Sp-T) (bL:0) (rL:0) (66:0) (O€-1) (40-1) (68-0) (ZE-0) (68-0) (Sr-0) (€S-0) SIOMOY JO L‘6 v-Z1 7-71 LL €-6 rl EIT Z-01 9:6 0:6 9-8 LL O-L Joquinu urs] “¢ (S-S-+-€) (8p-0) (Or-0) (Z1-0) (27-0) (07:0) (rE-0) (67-0) (17-0) (81-0) (1-0) (02-0) (61-0) (Wid) soUDDsa10yUI O-r ¢-S 9-p pe 0-r 6-€ a C6" 0-r re re re re Jo yBua] URI *Z (S-O€-7:Z1) (Spl) (Ip) (96:0) (60-1) (Ir-0) (0-2) (drt) (90-1) (€9-0) (79-0) (82-1) (0-1) (wd) 143104 6-81 S-0€ 0-0€ (ara! LST 9-SI S-07 9-€Z 6:81 LZ S-r1 S-81 al jue]d ues *] oduel ysenpoeur = foq ssoy pes soru uo} gepoy sojepsur ajeq uo} = 3sd104 Jajoereyy pue ueoW [[P19AC “AT Bu “Isom “¢ -A-SouY ‘] -oIppiq [[IMIOD ‘HD UOJOOg “4 -soog *q -IeEYUM'G -IAY4 ‘> -woyL ‘gq Aqreq ‘v 10) ‘'H IMANIDALSNOVUL VZIHYOTALIVG AO SNOILV1NdOd AA TAML JO HOVE NI SLNV'Td GaLOATAS ATWOGNVY NAL YOU (SASHHLNAUVd NI) SHOUNT GUVGNVLS GNV SLNAWAUNSVAW YALOVUVHO NVAW ‘7 ATAVL 159 MORPHOLOGY OF DACTYLORHIZA TRAUNSTEINERI ‘AABOY = Z ‘IWS = | ‘Ussqe = Q ‘dulurTe}s UTUeADOYIUP JO 99199q , ‘juasoid = | ‘yUasqe = Q ‘(jUaSoId UDYM JoJOWeIP WU | jnNoge pure iYsT] AIDA SAME) BUINIOdS Jeo] JO ddI30q_, "g JajovleyO// JoyoeleyD = xopul jes] ues], ‘7 Jajoeleyy/¢ Jo}oeIeyD = Aysusp [e104 , (2q0] (2q0] ye1Uus9 ye1Uu95 yuoUT qUOUI -wioid -wioid AIOA) AOA) [ep [ep — P2egOTL Peqo[L poqojly poeqgo[ly peqoyi4-loquioyi1-loqwuioys (QSIMIO410 Aounsip Ajounsip AoUuNsIp -qns -qns 0} poqoyIn pogo] JO) Wn][Oqe] Jo /P2QOTLN Psqo[LN peqgoylN Apounsip peqgoyLy ApouNsip paqoypiy ainjeUu poqgojuL, “ST (LI-I-L7-0) (01:0) (40-0) (ZT-0) (ST-0) (07:0) (€T-0) (60-0) (81-0) (11-0) (07:0) (91-0) (91-0) (wu) yidap ZL-0 LL‘0 L7:0 LUI 76:0 (0) Z€-0 Lv-0 OL-0 81-0 8-0 68-0 9¢-0 snuls ue */] (06:0-ZL:0) (€0:0) (20-0) (0:0) (70-0) (+0-0) (€0-0) (€0-0) (€0-0) (€0-0) (10-0) (20-0) (20-0) (wo) yisus] 18-0 06:0 8-0 SL-0 8-0 78-0 LL‘0 08-0 98-0 ZL-0 SL‘0 L8-0 6L:0 inds ues] 9] (9¢-0-87:0) (10:0) (10-0) (10-0) (70-0) (20-0) (10-0) (10-0) (10-0) (10-0) (10-0) (20-0) (10-0) (two) YIpIm ZE-0 T€-0 87-0 67:0 Z€-0 €£-0 T€-0 0€-0 I€-0 ZE-0 re-0 ZE-0 9€-0 inds ues] “CT (L8-0-ZL:0) (p00) (10-0) (720-0) (€0-0) (40:0) (€0-0) (€0-0) (€0-0) (20-0) (70-0) (20-0) (€0-0) (Wid) ysuUdZ] 08-0 78-0 LL‘0 ZL:0 08-0 LL-0 18-0 6L:0 L8-0 6L:0 €L:0 08-0 L8-0 wny[aqe] URS] “pT (Z1-1-06:0) (90:0) (70-0) (€0-0) (€0-0) (S00) (€0:0) (40-0) (€0-0) (40-0) (70-0) (20-0) (0-0) (Wid) YIpPIM 00-1 Z1-1 06-0 L6:0 76-0 60°T 76:0 96-0 LO-1 76-0 56:0 90:1 LOT uNyjoqe| URI] “ET (0-7-9-0) (=) G) (>) Ga) G—) () () €) (=) =) (=) G-) psurureys el 8-0 9-0 C1 aa 0-1 aa 0-1 ET 0:2 S-1 0-2 0-1 uruefooyjue — sjorig °Z] (S-I-Z-0) (=) (=) (=) (==) (a) C) ~) () (—) (=) (3) G-) pourureys utueAs 8-0 9-0 Z-0 0-1 8-0 0-1 8-0 Z-0 L-0 C1 0-1 C-1 $-0 -oyjue — woys Joddy ‘TT a3uerl ysenpoew oq ssoy pes solu uo} gjepoy soyepsur seq uo} 4so104 Jajoeleyy pue ueow Yel =I TT PUNT “ISOM f -A-SOUY "T -FOIPpIg [IMIO0D “HD uojoog “y -seeg’g ~yeyYM'd -Ad'D -WoYyL a Aqred “Vv JOD “H pduod 7 ATEVL 160 M. J. Y: FOLEY TABLE 3. VARIATION IN BRACT PERIPHERAL CELLS OF DACTYLORHIZA TRAUNSTEINERI AND RELATED TAXA Mean cell length and typical cell shape based upon ten contiguous peripheral cells from at least five bracts from separate plants in each population. Population mean cell length (um), and range of means for individual Typical cell shape in Species and population bracts in parentheses profile D. traunsteineri (Britain and Ireland) A. Dalby Forest 98 (80-115) elongated subacute to serrate B. Thornton Dale 83 (80— 90) ie C. Fylingdales 85 (73-100) se D. Wharfedale 74 (63— 80) a E. Beeston 77 (65— 88) F. Booton 83 (70-108) a G. Cothill 78 (65— 93) x H. Cors Erddreiniog 93 (85-105) i I. Rhos-y-gad 85 (68-100) ef J. W. Ross 88 (78-108) ¢ K. Killinaboy 85 (70- 98) - L. Kilmacduagh 89 (78-100) ee D. traunsteineri (Continental Europe) Schwarzsee, Kitzbihl (Austria) 89 (80-112) elongated subacute to serrate Schmerikon, St. Gallen (Switzerland) 89 (78-103) ‘ys Njurunda (Sweden) 76 (68-88) a Visby (Sweden) 84 (70-103) “ D. incarnata subsp. cruenta (Ireland) Knockaunroe, v.c. H9 46 (40- 53) rounded-crenate D. incarnata subsp. pulchella (Cumbria) Milnthorpe, v.c. 69 56 (S0- 58) rounded-crenate Sunbiggin Tarn, v.c. 69 46 (43— 50) . D. majalis subsp. purpurella (Cumbria) Beckfoot, v.c. 70 69 (60- 80) rounded-obtuse Sandscale Haws, v.c. 69 68 (S8- 78) “ Sunbiggin Tarn, v.c. 69 69 (SS— 83) Bh D. majalis subsp. praetermissa (England and Wales) Sandwich, v.c. 15 66 (58- 78) rounded-obtuse Elstead, v.c. 17 67 (S8- 75) te Kenfig, v.c. 41 57 (50- 65) i D. majalis subsp. occidentalis (Ireland) Ballynaleckan, v.c. H9 78 (70- 85) rounded-obtuse was noted that leaves were well spaced along the stem and not crowded towards the base as is the case in some species of dactylorchid. (d) Mean leaf index of second leaf (character 9, incorporating characters 7 and 8). The mean leaf index (length/naximum width of second leaf from stem base) showed some inter-population variation with the Wharfedale (D) population particularly low. Conversely the East Anglian populations (E and F) possessed relatively high values and the Irish (K and L) the highest of all, a character which is quite noticeable in the field. The Fylingdales plants (C) had the narrowest leaves (character 8) and showed little variation in width — this again was noticeable in the field. The mean maximum leaf width varied between 0-83-1-14 cm in the populations studied. (e) Degree of leaf spotting (character 10). Leaf spots, when they occurred, were always small in size, about 1 mm diameter, solid, never densely distributed, and often spread over most of the upper surface of the leaf. In many plants in all populations spotting was absent, and in the samples examined, was not recorded from the Yorkshire population (C), nor from either of the East Anglian populations (E and F), nor those from Anglesey (H and I) (although recorded in a small proportion MORPHOLOGY OF DACTYLORHIZA TRAUNSTEINERI 16] pee ae aed Siete eee wie le aN a Rosters Nee (Gor Saab Ss a pcp pg ll ee 3 ar eee i eee es este birreedll ted trea /TO TI, i LRG ORE MAT ERT Bd ae waisslaanenenis cneitd headtinhiwtuminaadny woags SLE, Salat) MY OCTET seats TSA AON Bie Total el a= Ficure 2. Profiles of peripheral cells of floral bracts (scale bar = 100 um). 1. D. traunsteineri (Schmerikon, St. Gallen, Switzerland), 2. D. traunsteineri (Schwarzsee, Kitzbiihl, Austria), 3. D. traunsteineri (Visby, Sweden), 4. D. traunsteineri (Thornton Dale, v.c. 62), 5. D. traunsteineri (Dalby Forest, v.c. 62), 6. D. majalis subsp. purpurella (Beckfoot, v.c. 70), 7. D. majalis subsp. praetermissa (Elstead, v.c. 17), 8. D. majalis subsp. occidentalis (Ballynaleckan, v.c. H9), 9. D. incarnata subsp. pulchella (Milnthorpe, v.c. 69), 10. D. incarnata subsp. cruenta (Knockaunroe, v.c. H9). of the plants at colony I (Roberts 1960)), nor that from Scotland (J). In contrast, two Yorkshire colonies (A and D) were extensively leaf-spotted as was one of the Irish populations (L). (f) Mean labellum width (character 13) and mean labellum length (character 14). Taken together these two characters showed no significant inter-population trend although five, Dalby Forest (A), Thornton Dale (B), Beeston (E), Cors Erddreiniog (H), and Kilmacduagh (L), had the greatest mean labellum widths. In all cases labella were slightly to semi-reflexed along the vertical axis. (g) Mean depths of sinuses and trilobed nature of labellum (characters 17 and 18). This composite character was partly assessed subjectively on flattened specimens of labella from the plants sampled, together with measurements of inter-lobe sinus depths. The Anglesey populations (H and I) and that from W. Ross (J) could be separated from the others in being the most distinctly trilobed and possessing deeper inter-lobe sinuses (mean depth 0-92-1-17 mm), whereas the southern England populations (E, F and G) were trilobed/sub-rhomboidal with generally much less prominent sinuses (0:32-0:70 mm) and with the Cothill (G) and W. Ross (J) plants, especially, displaying very prominent central lobes. The Yorkshire populations had on average rather more pronounced sinuses (0-36—-0-89 mm) than the southern plants. The Irish plants were somewhat variable and intermediate in this respect. (h) Bract peripheral cells — mean length and shape (character 19). The shape in profile and length of 162 M:4, Y.\ FOLEY. the peripheral cells of the floral bracts is known to vary between some species of dactylorchid (Hylander 1966). When such cells of various species and subspecies are examined under a magnification of 100, it is readily apparent that, whilst these cells vary somewhat in shape and length even within the same bract, certain characteristics emerge when analysed as described above. The results for cell length measurements for various populations are given in Table 3 and typical cell shapes taken from photomicrographs are shown in Fig. 2. For the twelve British and Irish and four continental populations of D. traunsteineri, there was very close agreement in peripheral cell length and shape — these being typically elongated, subacute to serrate, angled in the direction of the bract apex, of mean length 63-115 um (average 85 um) and up to 65 um in width. Populations of other species/subspecies of dactylorchids examined in the same manner were found to possess peripheral bract cells of a different shape and length; those of D. majalis subsp. purpurella and subsp. praetermissa were rounded-obtuse, shorter, mean length 50-83 um and up to 50 um in width, whilst for a population of subsp. occidentalis (Ballynaleckan, Co. Clare, v.c. H9), although rather longer (mean length 70-85 um and up to 50 um wide), were of similar shape to subsp. purpurella and subsp. praetermissa. Two subspecies of the diploid D. incarnata— subsp. pulchella and subsp. cruenta — had even shorter cells, mean length 40-58 um and up to 35 wm wide, and were characteristically rounded-crenate giving a beaded appearance to the bract edge. Whilst it is stressed that there is some variation in shape and size within and between bracts from both the same and different plants in all taxa, nevertheless a clear overall picture emerges, and they can be used as characters to identify D. traunsteineri and separate it from other dactylorchids. (1) Other characters (1) Upper stem and bracts — anthocyanin staining (characters 11 and 12). These two characters appear to be related, with the bracts being invariably more strongly pigmented than the upper stem. Whilst this character is semi-subjective, it was found nevertheless that the Yorkshire plants (A, B, C and D) were on the whole the most deeply anthocyanin stained, with the Irish plants (K and L) the least so. (2) Spur dimensions (characters 15 and 16). This would appear to be a less important character than those listed above. In all populations the spurs were relatively straight in lateral profile, especially on the under side, slightly curved on the upper and tapering to a blunt tip. (3) Labellum colour and markings. Base colour varied between deep pink and light red-purple with markings of a similar much deeper colour. All the Yorkshire colonies (and especially C and D) were generally noticeably deeper in base colour than the others and marked overall by many small spots and flecks. On the other hand the Anglesey plants (H and I) were much lighter in base colour yet marked with much more contrasting heavier semi-continuous lines and blotches, and the Irish plants (K and L) possessed a very characteristic lilac-pink base colour. The remaining populations (E, F and G) were similar in base colour to the Anglesey plants but much more lightly and less contrastingly marked. The degree of variation in all major characters between the populations studied is relatively small and throughout there is good agreement between the results presented here and those for the same populations studied by previous workers, e.g. Roberts (1988). Regional differences may be summarised as follows:— The Yorkshire plants (A, B, C and D) have fewer flowers of a deeper base colour with labella relatively finely marked, and with a proportion of plants with lightly spotted leaves. The upper stem and bracts are invariably strongly stained with anthocyanin. Other populations of similar plants have been given varietal status by Godfery (1933) as Orchis latifolia var. eborensis, and more recently as D. majalis subsp. traunsteinerioides var. eborensis by Bateman & Denholm (1983). The latter imply that such plants are always leaf marked, such marks more often than not being annular in shape, that inflorescences have often more than eight flowers, and that labella rarely exceed more than 7-5 x 9-5 mm, often more or less as broad as long and with a base colour only occasionally dark. Such diagnostic characters for the Yorkshire plants are not upheld in this study, and although there is some regional variation as outlined above, this appears to be within acceptable limits, and suggests that a separate varietal status may not be justified. Furthermore as Roberts & Gilbert (1963) acknowledge, in the two very small and isolated Yorkshire populations which they studied, there may have been appreciable genetic deviation from the norm, and the results now obtained seem to confirm this. The southern England populations (E, F and G) have longer narrow leaves (high leaf index), with MORPHOLOGY OF DACTYLORHIZA TRAUNSTEINERI 163 labella trilobed to sub-rhomboidal, and relatively lightly coloured but not heavily spotted or loop marked. Plants from the Anglesey populations (H and I) have much more lightly coloured flowers which are more heavily and contrastingly marked on the labella, these being distinctly trilobed, with relatively deep inter-lobe sinuses. The Scottish population (J) comprises plants of small stature with relatively short leaves, and flowers which are markedly trilobed, with deep sinuses and a prominent central lobe. Plants in both Irish populations (K and L) are relatively robust with a somewhat larger number of flowers in the inflorescence, these being a distinctive lilac-pink colour. They possess the highest leaf index of the populations examined and also have the longest leaves, and one population (L) has a high incidence of leaf-spotted plants; spurs are somewhat down-curved. Some of these characters agree closely with those possessed by plants in a population of D. traunsteineri from Njurunda, Sweden which is discussed in more detail below, and is thought to show some affinity to D. russowii (Klinge) J. Holub. From examination of the above data, it is apparent that plants from the twelve populations included in this study, because of their relative geographical isolation from other populations of this taxon, understandably exhibit some inter-populational differences. Nevertheless, the populations show good agreement in the key diagnostic characters and it is therefore concluded that they should be treated as a single species, possibly with varietal status accorded to some Irish populations. COMPARISON OF THE BRITISH AND IRISH POPULATIONS WITH DATA FOR D. TRAUNSTEINERI AT CONTINENTAL SITES The average and range of the mean character measurements obtained for the twelve British and Irish populations taken together are included in Table 4. These are compared with data supplied by H. R. Reinhard of Ziirich (pers. comm. 1989) for D. traunsteineri at the type locality, Schwarzsee, Kitzbuihl, and with other composite data obtained by Reinhard for populations in Austria and Switzerland (Reinhard 1985) (note — data given by Kalteisen & Reinhard (1986) and described as “Tocus classicus’’ differ slightly from the Kitzbihl data since they also include plants from Zell am See, Salzburg). My observations and measurements on bract peripheral cells for D. traunsteineri populations from the British Isles and continental Europe are given in Table 3. When the diagnostic characters 3, 4,5, 8,9, 13, 14, 18 and 19, are examined for British, Irish and continental European populations (Table 4), very close agreement is obtained in each case. In addition Reinhard (pers. comm. 1988) states that in plants from Austria and Switzerland the leaves can be either spotted or unspotted. The only significant difference lies in spur length which is found to be appreciably lower in this study (0-81 cm) than that found by Reinhard (1-11 cm and 1-09 cm). This is not a diagnostic character, but it is still surprising in view of the fact that Reinhard’s data (Reinhard 1985) for spur length of D. lapponica (Laest. ex Hartman) So6 (0-78—0-96 cm), a taxon closely related to D. traunsteineri, are consistently lower and in close agreement with the present results. This difference from Reinhard’s spur length data 1s also displayed in the results of Roberts (1988) and earlier workers. Nevertheless, the overall good agreement in the principal diagnostic characters supports the view of Roberts (1988) when concluding that the Rhos-y-gad plants (I) are indeed D. traunsteineri; similarly, the conclusion is now drawn that the British and Irish populations covered in this study are also conspecific with D. traunsteineri. However, if further work shows that British and Irish populations differ in some small respect from continental ones (e.g. as in spur length above), then the former should be placed as a subspecies of D. traunsteineri. Tables 3 and 4 also include the author’s data for a dactylorchid population from Njurunda, Sweden. This, whilst acceptably complying with the diagnostic characters for D. traunsteineri, nevertheless possesses some features applicable to the relatively elusive and rather vaguely described D. russowii, recorded for the Baltic area of Scandinavia and north-eastern Europe. This latter is a robust plant with a relatively long, rather dense-flowered inflorescence with a large number of flowers, each with comparatively small labellum dimensions, somewhat down-curved spurs, and with leaves quite heavily spotted and blotched. The Njurunda population occurs in neutral to slightly calcareous conditions (pH 6-9) growing with Epipactis palustris, the only locality for the latter in northern Sweden (R. Lidberg, pers. comm. 1989). Again most characters of the British populations agree well, and those from Ireland, (mean number of flowers, floral density, spur shape) especially so. Further work might show that D. russowii itself warrants consideration 164 M. J. Y. FOLEY only as a variety or subspecies of D. traunsteineri, possibly intergrading with the type, with the Njurunda population representing an intermediate variant. Morphometric data have also been obtained for a further population of D. traunsteineri from Visby, Gotland, Sweden which shares some similar characters with the Njurunda plants (inflorescence length, number of flowers). In the Visby plants, labella are slightly trilobed to sub-rhomboidal often with a prominent central lobe; the mean dimensions (n = 10) for labella are 0-96 cm (wide) x 0-79 cm, and for flattened spurs, 0-30 cm (wide) x 0-88 cm, with the floral bracts appreciably anthocyanin-stained (mean value 1-5). These data again agree closely with those for British and Irish populations, and fall well within the range of their appropriate character means. As already commented, D. traunsteineri in the British Isles is usually restricted to calcareous fens and flushes with an approximate pH range 7-0~7-5, although it can occur in slightly acidic conditions as at Fylingdales (pH 6-5). Its existence in calcareous habitats is also the case at many sites in the alpine region of continental Europe and in Scandinavia where its associates can include calcicoles such as Eriophorum latifolium Hoppe and Schoenus ferrungineus L. (Simonsson & Lindstr6m 1977; Jonsell 1982; Devillers-Terschuren & Devillers 1986). D. traunsteineri is also recorded from calcareous fens in the Haute-Marne area of north-eastern France growing with Schoenus nigricans, S. ferrugineus, and Epipactis palustris (Tyteca 1981), and from calcareous coastal fens in Pas-de- Calais (F. Rose, pers. comm. 1989). Records for D. traunsteineri from somewhat acidic habitats in continental Europe suggest a tolerance to such, possibly as a specialised ecotype of the species. Other records from distinctly acidic localities may be a result of confusion with a similarly described, taller, relatively narrow-leaved but more dense-flowered taxon accorded specific status as D. sphagnicola (Hoppner) Soo. This latter has been recognised within the geographical range of D. traunsteineri, and has been recorded for West Germany (Rube 1972), for Belgium and north- eastern France (Tyteca 1986), for Sweden (Birkedal & Danielson 1981; Ericsson 1982), and is also claimed to occur along with the taxon described as D. traunsteineri subsp. curvifolia (Nyl.) Soo in eastern Finland (Rasanen & Saari 1987). It is known to inhabit acidic habitats as shown by such associates as Juncus acutiflorus Ehrh. ex Hoffm., Drosera rotundifolia L., and Erica tetralix L. (Landwehr 1977; Tyteca 1981). Illustrations of continental D. traunsteineri closely similar to plants encountered in this study are given by Landwehr (1977) (see p. 159, nos. 1—5), as well as others in the same work from Ireland (p. 166, nos. 1-2). However those shown by him as D. traunsteinerioides (Pugsl.) Landw. (p. 166, nos. 3-5) from the same Inish locality are not typical of British and Irish plants, being much more dense- flowered and robust and are possibly of hybrid origin. A series of colour photographs of examples of D. traunsteineri from various sites in Switzerland and in Austria have been provided by H. R. Reinhard, and all show a very close similarity to plants from Bnitish and Irish populations. In addition, Reinhard has examined detailed colour photographs of typical plants from the British and Irish populations studied and has confirmed that all agree very well with D. traunsteineri from Kitzbuhl — “‘Alle Ihne Bilder stimmen recht gut mit D. traunsteineri von Kitzbuhel tberein’”’. COMPARISON OF THE BRITISH AND IRISH POPULATIONS OF D. TRAUNSTEINERI WITH OTHER DACTYLORCHIDS In Britain and Ireland, dactylorchids which may occasionally bear a superficial similarity to D. traunsteineri are the various subspecies of D. majalis:— subsp. occidentalis, subsp. purpurella, and sometimes subsp. praetermissa, and also and in particular, D. incarnata subsp. pulchella. Occasionally D. incarnata subsp. cruenta can exist in a form somewhat similar to D. traunsteinert. Comparative mean characters for D. majalis subsp. purpurella from populations in Cumbria (v.cc. 69 and 70) and for subsp. praetermissa from S. Lancs (v.c. 59), are included in Table 5, and can be seen to differ from the principal diagnostic characters of D. traunsteineri which are indicated in column 2. (It should be emphasised that not all the characters shown in Table 5 are useful in separating D. traunsteineri from the various subspecies of D. majalis.) D. majalis subsp. occidentalis, as observed in western Ireland, also differs noticeably from D. traunsteineri particu- larly with respect to flower density, leaf number and shape, and in having significantly greater maximum leaf width; it also differs in spur shape, and all three subspecies of D. majalis differ from D. traunsteineri in that they possess leaves which are relatively crowded towards the base of the stem. Bract peripheral cell data (Table 3) also help separate D. traunsteineri from these taxa. D. incarnata subsp. pulchella is readily separated from D. traunsteineri by its denser inflorescence MORPHOLOGY OF DACTYLORHIZA TRAUNSTEINERI 165 TABLE 4. COMPARISON OF DATA FOR THE TWELVE POPULATIONS (BRITISH ISLES) OF DACTYLORHIZA TRAUNSTEINERI STUDIED HERE WITH THOSE FROM EUROPEAN LOCALITIES For each character, means are given together with either the standard error or range in parentheses. Character 1. Mean plant height (cm) 2. Mean length of inflorescence (cm) 3. Mean number of flowers 4. Floral density* 5. Mean total no. of leaves 6. Mean no. of non- sheathing leaves 7. Mean length of second leaf (cm) 8. Mean max. width of second leaf (cm) 9. Mean leaf index* 10. Degree of leaf spotting* 11. Upper stem — antho- cyanin staining* 12. Bracts — anthocyanin staining* 13. Mean labellum width (cm) 14. Mean labellum length (cm) 15. Mean spur width (cm) 16. Mean spur length (cm) 17. Mean sinus depth (mm) from tip of lateral lobe (where available) 18. Trilobed nature of labellum (or otherwise) Twelve populations (A-L) from the British Isles 18-9 (12-2-30-5) 4-0 (3-4-5-5) 9-7 (7-0-12-4) 2-43 8-96 (5-0-14-37) 1-02 (0-83-1-14) 8-76 ((5-49)7-31-12-80) 0- 1-00 (0-90-1-12) 0-80 (0-72-0-87) 0-32 0-72 (0-27-1-17) usually distinctly Schwarzsee, Kitzbuihl, Austria (locus classicus) n = 13 (data ex Reinhard, pers. comm. 1989) 22-4 (0-96) 4-7 (0-23) 8. distinctly trilobed trilobed, occasionally with lines, flecks approaching sub- rhomboidal, dotted lined or flecked and loops “See Table 2 for an explanation of these characters. bn = 19. wa — 17. “True diameter (not flattened). nd = No data available. Composite data from eight separate European populations n = 75 (data ex Reinhard 1985) distinctly trilobed with sinuses, occasionally sub-rhomboidal, with lines, flecks and loops Mackelmyra, Njurunda, Sweden n= 10 (author’s data) 25-7 (1-07) 5: more or less trilobed M3). FOEEY 166 (=) ce) j—} (—) (08-ZI-1€-L(6b-S)) gXopul 19°€ ese COP SS°€ 9L°8 + yea] ues “6 (L1-0) (01-0) (90-0) (01-0) (pI-1-€8-0) (Wid) yea] puosas LO 69-1 it VEC cO-T + JO UIpIM “xeu URI *g (L9-0) (0€-0) (S7Z-0) (LZ-0) (L€-p1-00°S) (wid) Jeo] puosas S€-rl L6°S 06-L 1€-8 96:8 _ Jo yysua] ues */ (67-0) (60-0) (61-0) (00-0) (0-1-9-0) Sako] SuryRoYs VC jae 81 0-1 8-0 + -uou JO ‘OU URI “9 (87-0) (07-0) (61-0) (61-0) (0-r-€:€) SOARQ] JO “OU OL £9. 8° 8S 9-€ + [e}0} URI “CS (=) (>) C) (=) (%6:7-90°2) eL:b< 7E-S< eb-S< LL:S< Eb-~ + gAllsuap [PIO] “p =) = (=) (—) (p-ZI-0:L) SIMO JO 0b0< 07< ST< F< L‘6 + Joquinu uesypy “¢ (€S-0) (L1-0) (67-0) (p7-0) (S-S-r-€) (Wid) sdUadSaI0 YU! o°8 6°t 9 os 0O-P ~ JO ysus] URI “7 (99-2) (6b-0) (SL-0) (SS-0) (S-0€-Z:Z1) (wd) ysI9y4 L-OV v6 {Litt Ler 6:81 - juejd ues “| (Z'p/pe) (L'I/v€) 69 *9°A (0°9/S€) 69 *9°A (S'0/S¢) (T-v) eoour}oduil JayoeIeYyD 6S ‘OA ‘UOJSUOT ‘SMEP{ JROSpuRS ‘ule uiddiqung OL (9'A “JOOJYWVIg suonefndod dsajom Jayoevieyy pssiudajavid pjjaandaind pyjaandand ojjaandind 149U19]SUND4] °C ‘dsqns ‘dsqns ‘dsqns ‘dsqns syolou syolou *‘q syolou “q syolou ‘q ‘sosoyjuored Ul o8URI JO JOIID PIepULIS DY} JOYITA YIM J9YI9IO}] UDAIS oe SURDUT ‘19]9RIBYO YRS 104 VSSINYALAVUd “dS€NS ANV V7T7TAeNdYNd ‘dSANS SITVIVW ‘d JO SNOILV1NdOd HSILIMA HLIM IYANIDALSNOVYL VZIHYOTALOVG AO (SHTSI HSILLIMH) SNOLLVINdOd AATAML AO NOSINVdWOO ‘S ATAVL 167 MORPHOLOGY OF DACTYLORHIZA TRAUNSTEINERI ‘QdUSNSIIOYU! OY} SUIAONSOp Noy Ajasiooid suTWID}ap 0} INOYTIG , ‘s1o}ov1BYO 9soy} JO UONeULIdxo UP JOJ Z BIQUI 99S, ‘Tnyasn jou = — ‘ynjasn AyTensed = —/+ ‘[njosn = + ‘g a[qeRJ, Ul paysi] exe) oY} WI wauiajsunval ‘gq Sunesedas ut s19}9eIeYd JO ssouynjosn oy] , peqoy peqoy peqo| Ppayooy JO pour] ‘psyj0p “Ly APYstys “Uy Apystys “Uy Apysiys ‘Teploquioys-qns SOWT}IWIOS SOUITJIWIOS SOWIOWIOS Suryoeoldde Aqye (AsIM194}0 poqoyl} ‘jeploquioyl ‘jeproquioys ‘jeploquioyl -UOISeD90 ‘paqoyLy 10) win}[oaqey] jo /jeploqwuoyl SS9] IO JIOU SS9] IO JIOU! SSO] JO JJIOW Ajounsip Ayyensn —/+ oInjeu Peqo]l1L QT (Z0-0) (Z0-0) (€0-0) (10-0) (06:0-ZL:0) (wid) yy8u9] 76-0 69-0 08-0 18-0 18-0 ~ Inds ues ‘97 (Z0-0) (10-0) (10-0) (10:0) (9€-0-87-0) (Wd) YIpPIM ZE-0 87-0 ZE-0 re-0 Z€-0 = inds ues “ST (Z0-0) (Z0-0) (Z0-0) (Z0-0) (L8-0-ZL:0) (wo) Yyysu9] 98-0 79-0 99-0 19-0 08-0 - wINn]jaqe] URI] ‘pT (€0-0) (Z0-0) (€0-0) (€0-0) (ZI-1-06-0) (wo) YIpIM L71 18-0 78-0 98-0 00-1 —/+ wINn[aqe] UB “ET (=) Cs) (<=) (>) (0-7-9-0) qSUlure}s $-0> 0-1 0-1 S-0 €-1 —/+ ulueAosoyjue — sjoelg *Z] ) (==) i) Ce) (S-I-Z-0) qsutureys ulueAd ¢-0> 9-0 9-0 ¢-0 8-0 —/+ -oyjue — ways Joddy “TT =) (Ge (=) eS) (8-0-0:0) qsuniods yea] 1-0 €-0 0-0 0-1 Z:0 - Jo 90189 “0 (Z'p/re) (L'T/p€) 69 “O°A (0°9/SE) 69 “9°A (S'0/SE€) (T-v) ,oour}oduit JayovleyD 6S [oA ‘u0j3u07T “SMEH o[eospues ‘ule ulssiquns OL TO )e/s\ ‘JOOJYIOG suoneindod DATOM L Jojyoerey) pssiusajavid pjjaindind pjjaindind vjjaandind 149U12JSUNDA] *G ‘dsqns ‘dsqns ‘dsqns ‘dsqns syolou ‘q syolou *q syolou *q syolou *q Psuor § ATEVL M. J. 'Y. FOLEY 168 EE E———EE——E—E——E————E eee irre (x1 —x, Ajjensn) opi uey} 193u0] XS-¢ ISRO] 1B yea] JS9dI1e] ‘9}e[OSNUR] -reouly ‘payjods Apysty Sunjods = ceifate = = ae ae er os + ze io poyodsun soaray ~=—s /adeys jea] Wd 07-1 Je yIpIn Es = “ a = + te + + + JSoSie] YIPIM WNWIxey yeay p JO WNWIxew [B}0} ‘ways SuoTe SOAB9] os Sch = Ps a te + ek ag ae pooeds-|Jom Soave] JO ON yisu2] a0usdsa10yul Jo wid Jed (ued) sIOMOY ¢€ URYI Ayisuop = = ae = = te a + uk fe SSO] — DOUDOSOIOYUI Xe] [e10L4 QoUddSdIOYUI Jod p] JO WNWIxeul 0} SIOMOY —/+ | = oa =| tr + + tT 7 + L WOJJ — palaMmoy-Mo4 jO “ON 6H ‘2A 69 ‘O°A 6S ‘OA 69 ‘O°A 6H ‘9°A UspemMg —§ gUapamg ysuoneindod ,(snazssvj2 (1-v) uondusseq JaqoeieyD soluneysouy = eds oy UTI u0j3Uu0'T] wie ueyooeudyeg ‘pueyog ‘epunin{y 4310 §njo]) pourmexo uIddIqung ‘Kasia ‘erAwoyoRW —(uesdoing) jynqziry suoneindod SOT}TBIO] ysu] ouldiy ysHug IV EE ———E———E_ aan ae pjuanso pyayajnd —s vsstusajavid —_—ojjaandand sijpjuap1990 tuaulajsunval 021440] 4}90q ‘dsqns ‘dsqns ‘dsqns ‘dsqns ‘dsqns pjpusvoul ‘q = vjpuavoul‘q_ = syjvivu *q syolou *q syvlou “q ON OOS ie Sie hala Se AAS AAA TE nee ene enc ee a eee SEK { a ae f ‘UMOYS BIDYM Jdaoxe SUORAIISQO PUP S]USWINsPIW Pay S.Joy Ne 34} uodn paseq Biep [|TV ‘uonduosop YIM sooisesip = — ‘UodOsop Joy sUTJJap10g = —/+ ‘uoNddsSap YIM SddITe = + SBLIA}LO BS9Y} JO JSOW O} WAIOJUOS pinoys Auojoo Aur ut syuejd poyoajas AjwWopues +0] JO sIskyeUe UP JOJ SON[LA ULL SY} ‘USAID SIU] OY} SpIs}NO a] SOUTOWOS Aew syueyd yenpiaipul ysnoyyy VXVL VZIHYOTALOVG HSIUI GNV HSLLIMd WAHLO HLIM GaaVdWOO IMANIA.LSNOVUL VZIHYOTALIVE AO SUALOVUVHD 9 TTEVL 169 MORPHOLOGY OF DACTYLORHIZA TRAUNSTEINERI ‘pouteys AyfeotdA} you yng poyods sjovig , "sooeJins YI0Q UO payiods A[IABOY S2AP2q ITY _ ‘ponods Ajiavoy uayo soava’T , ‘NMossn4 ‘q 0} AWUYJY , ‘UURWUYOSIAA “J pure preyUIoYy UH Ag payddns jetajyew oat] awWOg “686 ‘886T “SUIUIOD ‘siad ‘preyUIoYy “Yy “H pur ‘(Sg6T) pieyuloy xo oe BLsNY pue pUk]I9Z}IMG Ul SUONe|Ndod Joj spiejaq , Sr re a (win Cg adeioAe ‘un CL T-09 YIU] uvoul [eo1dd}) a3e1108 0} 9]Nde-qns poyesuO|S See) = = ae =e = ae a. ae 26 a s][90 Jessyduad jovig pelg sutode} Anysys Ajuo pue ‘[eJUOZIIOY ssa] JO JIOW odeys = — —/+ _ _ + = + + + pure ysnqod ‘yori Inds Inds sasnuls o[qesonou pue ‘oqo [e1U90 pojesuoje ‘jusuTWoId YUM UDO ‘pogoy odeys sa! = = = “a aye a at ae ae Ayyensn winyjaqe’] winyjoqe’] apiM wo yipiM = = 4 as Be “1 ok fe fe a 06-0 1se9] 18 WUNTTOqQe’T] winyjoqe’] pourejs-urueAsoyyue UONeINO]OS ae + = Hs = a + + + + Ajqesonou usaqo sjovig porig Se a) ee Se eS Seer Se 6H ‘o'A 69 ‘O'A 6S “A 69 OA 6H ‘o'A uspaMs quopemg ,suoneindod _,(snorssvj9 (J-v) uondussaq qoyoeleyo soluneysouy «= ado yu uojsUu0T] ule], ueyogfeudyeg ‘puejog ‘epuniniyy 1319 §n20]) pourmexa uld3iquns ‘Aqsta = ‘eiAwjayoey —(ueodoingq) ynqziry suonejndod SOnTedO] ysuy ouldiy ysHig IV eee pjuanso pyjayajnd ~— vssiuusajavad ~— ojjaandind s1]0JUap1990 Maulajsund4é] DZ1YAO]Ajo0Gq ‘dsqns ‘dsqns ‘dsqns ‘dsqns ‘dsqns DIDULDIUI ‘GQ = dioUADIUI-q = syvipU ‘Gq syolou -q syolou gq er I ee ee = SL SE ee ee eee P dod 9 AIGVL 170 M.I>¥. FOLEY (often 20 or more flowers per plant and floral density >5 flowers/cm of inflorescence), its relatively small, strongly reflexed, and not noticeably trilobed labella (mean 0-67 cm wide x 0-56 cm long (details from Milnthorpe, v.c. 69)), relatively broad leaves, and again and most characteristically, by the rounded-crenate and consistently shorter peripheral cells of the floral bracts (Table 3). This latter character is an effective method of separating all subspecies of D. incarnata from D. traunsteineri. D. incarnata subsp. cruenta is also readily separated on characters similar to those which separate subsp. pulchella from D. traunsteineri. In Table 6, based mainly upon data obtained by the author, the key characters useful in identifying British and Irish D. traunsteineri are applied to various other dactylorchids as well as to D. traunsteineri. From this, the latter is clearly shown to be separate from the other dactylorchids. CONCLUSIONS The mean values for the principal characters of the twelve British and Irish populations examined were found to be similar, although some regional trends could be detected. Whilst individuals within any population may deviate appreciably from the typical variant, generally populations exhibit a uniformity in diagnostic characters, particularly the characters of floral density, leaf index, maximum leaf width and labellum width. The data obtained for all populations were very close to those recorded for continental D. traunsteineri, including plants at the type locality. Also, when these data were compared to those for populations of other dactylorchid taxa, clear differences were found. It is therefore concluded that plants from the British Isles are conspecific with D. traunsteineri from the classic localities of continental Europe, and should be retained at specific rank and not placed under D. majalis as suggested by Bateman & Denholm (1983). SYNONYMY Dactylorhiza traunsteineri (Sauter) Sod, Nom. nov. gen. Dactylorhiza 6 (1962). Orchis Traunstei- neri Sauter ap Reichenbach, Flora Germ. Exc. (1831). Orchis latifolia L. var. eborensis Godfery, Mon. Icon. Br. nat. Orchidaceae 219 (1933). O. majalis Reichenbach subsp. Traunsteinerioides Pugsley in Proc. Linn. Soc. Lond. 148: 124 (1936). O. Francis-Drucei Wilmott in Proc. Linn. Soc. Lond. 148: 128 (1936). O. majalis Reichenbach subsp. traunsteinerioides Pugsley var. eborensis (Godfery) Pugsley inJ. Bot., Lond. 77: 54 (1939). O. traunsteinerioides (Pugsley) Pugsley in J. Bot., Lond. 78: 179 (1940). Dactylorchis traunsteineri (Sauter) Vermeulen, Stud. Dactyl. 66 (1947). Dactylorchis traunsteinerioides (Pugsley) Vermeulen, Stud. Dactyl. 66 (1947). Dactylorhiza traunsteineri (Sauter) S06 subsp. traunsteinerioides (Pugsley) So6, Nom. nov. gen. Dactylorhiza 6 (1962). D. traunsteineri (Sauter) Sod subsp. francis-drucei (Wilmott) So6, Nom. nov. gen. Dactylorhiza 6 (1962). D. traunsteinerioides (Pugsley) Landwehr in Orchideeén 37: 80 (1975). D. traunsteineri (Sauter) So6 subsp. hibernica Landwehr in Orchideeén 37: 79 (1975). D. majalis (Reichenbach) P. F. Hunt & Summerhayes subsp. traunsteinerioides (Pugsley) Bateman & Denholm in Watsonia 14: 372 (1983). D. majalis subsp. traunsteinerioides var. traunsteinerioides Bateman & Denholm in Watsonia 14: 373 (1983). D. majalis subsp. traunsteinerioides var. eborensis Bateman & Denholm in Watsonia 14: 373 (1983). D. majalis subsp. traunsteinerioides var. francis- drucei Bateman & Denholm in Watsonia 14: 373 (1983). DESCRIPTION The principal diagnostic features of populations of D. traunsteineri in the British Isles are summarised as:— Variable in height (12-30 cm tall), distinctly slender in appearance, often with flexuous stems. Leaves few (usually 3-4 including 0-1 non-sheathing leaf), narrow, linear-lanceolate, strict to occasionally subarcuate, usually more than 7X as long as broad, (second lowest (usually also largest) leaf 8-12 mm at maximum width), well-spaced out along the stem, unspotted or lightly spotted (spots solid, c. 1 mm diameter). The inflorescence is often rather variable in shape or roughly cylindrical/tapering, often subsecund, few-flowered (usually 7-12) and distinctly lax, the flowers coloured deep pink/red-purple, relatively large with + semi-reflexed, broad (c. 10 mm) labella which are flecked and loop-marked in a similar but much deeper colour, usually distinctly trilobed, sometimes with a pronounced central lobe. The floral bracts and upper stem are often MORPHOLOGY OF DACTYLORHIZA TRAUNSTEINERI 171 stained red-brown with anthocyanin, and the peripheral cells of these bracts are characteristically elongated, sub-acute to serrate, 60-115 um in mean length. Plants are in full flower from late May to mid-June, and are usually found in calcareous fens and flushes. A provisional distribution map of D. traunsteineri in the British Isles (Fig. 1) is included, but it is very likely that further field work will lead to a consolidation of its distributional pattern. Colour photographs of plants, samples of pressed and mounted labella, spurs and bracts of the plants examined, photomicrographs of bract peripheral cells, and site details of the populations studied, are all retained by the author. ACKNOWLEDGMENTS I am very grateful to R. H. Roberts for his helpful comments and advice whilst drafting this paper and to L. Blainey, M. Keane, M. R. Lowe, A. J. Richards, R. H. Roberts, F. Rose, W. A. Sledge, B. G. Tattersall and D. A. Webb for contributing information relating to the location of populations of this taxon. I am also very grateful to H. R. Reinhard for commenting on photographs of the populations studied and for providing information on European D. traunsteineri and for data relating to the type locality, to B. G. Johannsson and R. Lidberg for help with Swedish populations, to M. S. Porter and B. G. Tattersall for assistance in the field, and to G. Halliday (University of Lancaster) for providing herbarium facilities. REFERENCES BATEMAN, R. M. & DENHOLM, I. (1983). A reappraisal of the British and Irish Dactylorchids, 1. The tetraploid marsh-orchids. Watsonia 14: 347-376. BATEMAN, R. M. & DENHOLM, I. (1989). Morphometric procedure, taxonomic objectivity and marsh-orchid systematics. Watsonia 17: 449-455. BIRKEDAL, S. & DANIELSON, J. (1981). Forsta fyndet av Mossnycklar (Dactylorhiza sphagnicola) i Skane. Svensk bot. Tidskr. 75: 313-314. Bo ton, J. H. & Horsman, F. (1987). Botanical records for 1985. Naturalist, Hull 112: (No. 980) 25. CUNNINGHAM, M. H. & KENNETH, A. G., eds. (1979). The flora of Kintyre. Wakefield. Curtis, T. G. F. & McGouau, H. N. (1988). The Irish red data book. 1. Vascular plants. Dublin. DEVILLERS-TERSCHUREN, J. & DEvILLERS, P. (1986). Distribution et systématique du genre Dactylorhiza en Belgique et dans les régions limitrophes. Naturalistes belg. 67: 143-155. Ericsson, B. C. (1982). Mossnycklar och skogsfru vaxter pa Oland. Svensk bot. Tidskr. 76: 1-4. Fo.ey, M. J. Y. (1986). Dactylorhiza maculata (L.) So6 Xx D. traunsteineri (Sauter) Soé in N. E. Yorks. Watsonia 16: 175-176. Fotey, M. J. Y. (1989). Dactylorhiza traunsteineri (Sauter) So6: variants in north-east Yorkshire. Watsonia 17: 355-356. Goprery, M. J. (1933). Monograph and iconograph of native British Orchidaceae. Cambridge. HEDLEY, S. (1987). Plant records. Watsonia 16: 450. HeEs.op-Harrison, J. (1953). Studies in Orchis L., II. Orchis traunsteineri Saut. in the British Isles. Watsonia 2: 371-391. Horsman, F. (1989). Botanical Report for 1987. Naturalist, Hull 114: 51. Hy anper, N. (1966). Nordisk Karlvaxtflora 2: vii. Stockholm. JONSELL, B. (1982). Angsnycklar och sumpnycklar i nordligaste Uppland. Svensk bot. Tidskr. 76: 103-111. KALTEISEN, M. & REINHARD, H. R. (1986). Orchideen in zentralen italienischen Siidalpenraum. Mitt. bl. Arbeitskr.-Heim. Orch., Baden-Wiirtt. 18: 1-136. KENNETH, A. G., Lowe, M. R. & TENNANT, D. J. (1988). Dactylorhiza lapponica (Laest. ex Hartman) So6 in Scotland. Watsonia 17: 37-41. Lacey, W. S. (1955). Orchis traunsteineri Saut. in Wales. Proc. bot. Soc. Br. Isl. 1: 297-300. Lacey, W. S. & Roserts, R. H. (1958). Further notes on Dactylorchis traunsteineri (Saut.) Vermeul. in Wales. Proc. bot. Soc. Br. Isl. 3: 22-27. LANDWEHR, J. (1975). Het geslacht Dactylorhiza. Orchideeén 37: 76-80. LANDWEHR, J. (1977). Wilde orchideeén van Europa. Vol. 1. Amsterdam. Lowe, M. R., TENNANT, D. J. & KENNETH, A. G. (1986). The status of Orchis francis-drucei Wilmott. Watsonia 16: 178-180. Petcu, C. P. & Swann, E. L. (1968). Flora of Norfolk. Norwich. |i M. J. Y. FOLEY Pucs.ey, H. W. (1935). On some marsh orchids. Bot. J. Linn. Soc. 49: 553-592. Pucs.ey, H. W. (1936). New British marsh orchids. Proc. Linn. Soc. Lond. 148: 121-125. Pus ey, H. W. (1939). Recent work on dactylorchids. J. Bot., Lond. 77: 50-56. Pucs ey, H. W. (1940). Further notes on British dactylorchids. J. Bot., Lond. 78: 177-181. RASANEN, J. & Saari, H. (1987). KaitakammekAn (Dactylorhiza traunsteineri) muuntelusta Pohjois-Karjalassa. Lutukka 3: 35-39. REICHENBACH, H. G. L. (1831). Flora Germanica Excursoria, p. 140. Lipsiae. REINHARD, H. R. (1985). Skandinavische und Alpine Dactylorhiza-Arten (Orchidaceae). Mitt. Bl. Arbeitskr.- Heim. Orch., Baden-Wiirtt. 17: 321-416. Roserts, R. H. (1960). The Wicklow marsh orchid in Anglesey. Nature Wales 6: 14-17. Roserts, R. H. (1966). Studies on Welsh orchids. III. The coexistence of some tetraploid species of marsh orchids. Watsania 6: 260-267. Roserts, R. H. (1988). The occurrence of Dactylorhiza traunsteineri (Sauter) So6 in Britain and Ireland. Watsonia 17: 43-47. Roserts, R. H. (1989). Errors and misconceptions in the study of marsh-orchids. Watsonia 17: 455—462. Roserts, R. H. & Givsert, O. L. (1963). The status of Orchis latifolia var. eborensis Godfery in Yorkshire. Watsonia 5: 287-293. Roserts, R. H. & Warp, S. D. (1978). Plant records. Watsonia 12: 177. Rose, F. (1975). Plant records. Watsonia 10: 433. Ruse, G. (1972). Dactylorhiza sphagnicola (H6ppner) So6 in der Liineburger Heide. J. ber. naturw. Ver. Wuppertal. 25: 138-139. SAUTER, J. (1837). Flora 20: Beilbl. 3. SCANNELL, M. J. P. (1973). Dactylorhiza traunsteineri (Sauter) So6 in east Cork, mid Cork, Offaly, Meath, Leitrim and east Mayo. Ir. Nat. J. 17: 426. Simonsson, P. & Linpstr6M, B. M. (1977). Myrar i vastra Ramsele, Angermanland. Svensk bot. Tidskr. 71: 419-428. SwANNn, E. L. (1975). Supplement to the Flora of Norfolk. Norwich. TENNANT, D. J. (1979). Dactylorhiza traunsteineri in Yorkshire. Naturalist, Hull 104: 9-13. TENNANT, D. J. (1987). Plant records. Watsonia 16: 450. TENNANT, D. J. & KENNETH, A. G. (1983). The Scottish records of Dactylorhiza traunsteineri (Sauter) Soo. Watsonia 14: 415-417. TENNANT, D. J., WRIGHT, B. & WriGHT, A. (1983). Plant records. Watsonia 14: 431. TytEca, D. (1981). Observations sur quelques Dactylorhiza de Belgique et du nord de la France. Bull. Soc. r. Bot. Belg. 114: 15-30. TyTeEcA, D. (1986). Observations orchidologiques en Belgique et dans les territoires voisins: bilan 1981-1985. Dumortiera 34-35: 107-111. VERMEULEN, P. (1947). Studies on dactylorchids. Utrecht. Wess, D. A. & SCANNELL, M. J. P. (1983). Flora of Connemara and the Burren. Cambridge. Wivmotr, A. J. (1936). New British marsh orchids. Proc. Linn. Soc. Lond. 148, 126-130. (Accepted November 1989) Watsonia, 18, 173-177 (1990) 173 Displacement of Elodea canadensis Michx by Elodea nuttallii (Planch.) H. St John in the British Isles D. A. SIMPSON The Herbarium, Royal Botanic Gardens, Kew, Surrey, TW9 3AE ABSTRACT Elodea nuttallii (Planch.) H. St John has spread rapidly since its introduction into the British Isles in 1966. A feature of its spread has been its displacement of E. canadensis Michx in places where the latter was well established. One possible reason is that E. nuttallii is able to achieve more rapid stem elongation, as well as produce a greater number of axillary stems over a given period of time, resulting in E. canadensis eventually being shaded out of a locality. Cultivation experiments suggested that stem elongation and axillary stem production was significantly (P<0-001) greater in E. nuttallii under nutrient-poor and -rich conditions and a range of light intensities. Stem elongation in E. nuttallii appeared to be most rapid immediately after establishment, while in E. canadensis there was a gradual elongation as time progressed. These characteristics may lead to more rapid formation of a canopy of stems and leaves by E. nuttallii at or near the water surface, a factor which may help to bring about the eventual displacement of E. canadensis in many places. INTRODUCTION Elodea canadensis Michx and E. nuttallii (Planch.) H. St John are native to North America, but as adventive species they have become widely distributed around the British Isles. E. canadensis has long been established, but E. nuttallii was first recorded in 1966, since when it has spread rapidly. An interesting feature of its spread was its apparent displacement of E. canadensis in many localities where the latter species had been well established. This was observed by a number of workers (cf. Briggs 1977; Lund 1979; Simpson 1984) who noted that E. nuttallii more-or-less replaced E. canadensis over a period of one or two years, with only a few plants of the latter remaining after this time. The speed of displacement initially led to suggestions that E. nuttallii was a phenotypic variant of E. canadensis, although this has been disproved (Simpson 1988). Nevertheless the displacement of E. canadensis appears to be continuing and, indeed, personal observations made in southern England over the past two years suggest that here, at least, E. canadensis is now much less common than E. nuttallii. However it should be noted that there are some sites where there seems to be little competition between the two species, and others where E. canadensis is still the only species present. A number of incidental observations were also made on material of both species which had been planted at the same time in a nutrient-rich substrate in a large tank. Within three to four weeks of planting E. nuttallii had become well-established, producing many stems, whereas E. canadensis remained short and more or less decumbent with comparatively few stems being produced. E. canadensis remained in a similar condition throughout the growing season, while E. nuttallii continued to produce stems in quantity. One possible reason for the displacement of E. canadensis in natural habitats is that E. nuttallii is able to achieve more rapid stem elongation, as well as produce a greater number of axillary stems over a given period of time, with the result that E. canadensis may eventually be shaded out. To examine this in more detail, three cultivation experiments were carried out in which stem elongation and production of axillary stems were measured in both species under differing environmental conditions. 174 D. A. SIMPSON MATERIALS AND METHODS Material was collected from the Lancaster-Kendal Canal, Burton-in-Kendal, Westmorland, v.c. 69 (E. canadensis) and R. Lune, Skerton, Lancaster, v.c. 60 (E. nuttallii) in August 1983. The three experiments, also carried out in August 1983, were as follows: (i) 5 cm long apical stem sections of each species were planted in a 5 cm deep, nutrient-poor coarse sand/gravel mixture, which had been put into six 30 cm diameter, 40 cm deep polypropylene bins. 15 stem sections of each species were planted in each bin. The bins were placed in pairs ina glasshouse where supplementary illumination was Provided by 20 Atlas ‘Daylight’ fluorescent tubes, giving a maximum light intensity of 250 umol m~s~!. The lights were set to give a 16 hour photoperiod and temperature was maintained at 22+5°C. Ten plants of each species were harvested on each date and immediately pressed. Stem length, together with the number of axillary stems produced by each plant, were recorded after 10, 20 and 30 days. Analyses of variance were carried out on species data for each harvesting date. (ii) Plants were grown at four surface light intensity levels, 250 (high), 35 (medium), 2 (low) and 0 umol m~*s~'. Otherwise, culture methods were similar to those in experiment (i). Ten plants of each species were grown at each light intensity. After 35 days stem length and the number of axillary stems per plant were recorded. Analyses of variance were carried out on stem length data between species at each light intensity. (iii) Similar to experiment (ii) except the plants were grown in nutrient-rich river sediment. RESULTS AND DISCUSSION The results of experiment (i) are shown in Fig. 1. E. canadensis showed almost no stem elongation after ten days, and had grown to only about 7 cm after 30 days. Conversely, E. nuttallii showed a marked amount of stem elongation after ten days, when the mean stem length was 9-5 cm. By 20 days mean stem length was 12-5 cm, and this was followed by a further, although smaller increase after 30 days. The difference in stem length between the two species was highly significant (P<0-001) on each sampling date. A similar rate of stem elongation in E. nuttallii was noted by Kunii (1982). The results of experiments (ii) and (iii) are shown in Fig. 2 and Table 1. As expected, no plants 20 Mean length of plant (cm) Days Figure 1. Growth of Elodea canadensis (@) and E. nuttallii (A) under conditions of constant temperature, light intensity and photoperiod. DISPLACEMENT OF ELODEA CANADENSIS BY E. NUTTALLII 175 survived in total darkness. Otherwise both species demonstrated greater stem elongation in nutrient-rich substrate, and again, this was expected considering the impoverished nutrient regime provided by the coarse sand/gravel mixture. However in both experiments E. nuttallii showed significantly (P<0-001) greater stem elongation than E. canadensis after 30 days at the two lower intensities. In material growing on nutrient-poor sand/gravel this difference was greatest at medium light intensity, but in plants growing on nutrient-rich river sediment it was greatest at high light intensity. The number of axillary stems per plant decreased in both species with decreasing light intensity in both substrate regimes. However at medium and high light intensities the number of branches was considerably higher in E. nuttallii, again in both substrate regimes. 50 40 WwW Oo 20 Length of plant (cm) 0 50 100 150 200 250 Light intensity (\umol m7? s~!) FicureE 2. Growth of Elodea canadensis (@) and E. nuttallii (A) after 30 days at differing light intensities in nutrient-poor (- -) and nutrient-rich (—) subtrates. The data from experiment (i) also suggest that stem elongation in E. nuttallii is most rapid in the period immediately after establishment, while in E. canadensis there is only a gradual elongation as time progresses. As only nutrient-poor substrate was used in this experiment, it could be argued E. TABLE 1. MEAN NUMBER OF AXILLARY STEMS PER PLANT ON ELODEA CANADENSIS MICHX AND ELODEA NUTTALLII (PLANCH.) H. ST JOHN AFTER 30 DAYS GROWTH ON NUTRIENT- POOR AND -RICH SUBSTRATES UNDER THREE DIFFERING LIGHT INTENSITIES E. canadensis E. nuttallii Substrate type Light intensity (mean+SE) (mean+SE) Nutrient-poor low 1-00+0-001 1-00+0-001 medium 1-00+0-01 3-00+0-01 high 1-00+0-001 5-:00+0-01 Nutrient-rich low 1-00+0-001 1-00+0-001 medium 1-75+0-001 3-25+0-01 high 8-75+0-02 11-00+0-01 176 D. A. SIMPSON canadensis might show more rapid stem elongation on nutrient-rich substrate. However experiment (ili) suggests that, even when nutrient-rich substrate is used, E. nuttallii is still capable of faster growth. Although these experiments are by no means exhaustive, they do suggest that E. nuttallii has a greater rate of stem elongation and axillary stem production over a given period of time. One way in which these could be related to the displacement of E. canadensis is in the formation of a canopy. Canopy formation is of particular importance to submerged macrophytes which, because of the light attenuating properties of water, often grow in low light intensities. One of the main problems for such plants is to obtain sufficient light for photosynthesis, and to overcome this they concentrate photosynthetic tissue towards the water surface, as near as possible to the light (Barko & Smart 1981). In shallow water this results in the formation of a canopy at the water surface which is composed of a large number of densely crowded stems and leaves. Such a feature is characteristic of Elodea and related genera. An important feature of the canopy is that light levels below it are much reduced. This was amply demonstrated in Hydrilla verticillata (L.f.) Royle by Haller & Sutton (1975) who recorded a 95% reduction in light intensity under a canopy of this species. The effect of this would be to shade out other species present in the same habitat. Canopy production confers an obvious advantage as it severely restricts the competitive ability of other species (Haller & Sutton 1975; Titus & Adams 1979; Barko & Smart 1981). The implications of this are that if stem elongation and axillary stem production is more rapid in E. nuttallii, this species may produce a canopy more quickly than E. canadensis and thereby shade out the latter. In deep water, aquatic plants are often unable to elongate to the water surface (Barko et al. 1982) and they do not produce a canopy of the type described above. Nevertheless a canopy may still be formed, although in this case it consists of longer, less densely crowded stems and leaves. Again, the formation of a canopy may be more rapid in E. nuttallii. This is supported by observations made in Mitchell Wyke Bay, Windermere where E. nuttallii rapidly displaced E. canadensis (Lund 1979), although both species grow at depths of up to 3 m or more and never reach more than 1-35 m below the water surface (Simpson 1983). The results obtained from experiment (i) may also be analogous to stem elongation occurring early in the growing season, which might be another significant factor in determining the success of E. nuttallii over E. canadensis. Assuming that both species commence active growth at the same time, stem elongation may proceed more rapidly in E. nuttallii; thus canopy formation in this species is already well advanced before E. canadensis is able to reach an equivalent stage. As a result, E. canadensis may only be able to survive in gaps within the stand of E. nuttallii, or on its fringes, through the remainder of the season. Observations in the field would seem to fit in with this hypothesis, since E. canadensis is usually seen in this type of situation when the two species are growing in the same habitat. Incidental observations were also made of the time taken for new roots to appear on the stem pieces. This was faster in E. nuttallii, and the first roots were usually seen about four days after planting. Those of E. canadensis took about 10-14 days to appear. In the British Isles both Elodea species spread by vegetative means. The stems are brittle and, when small pieces of stem break away from the parent plant, roots often form at the nodes, allowing the piece to establish itself as a new plant. Thus E. nuttallii may have a further advantage over E. canadensis in being able to establish itself faster by rooting more quickly. It should be emphasised that the experiments described were of a fairly simple nature, and that measurements were made after only a short period of cultivation. Therefore it is conceivable that some of the differences may have occurred due to prevailing conditions in the original habitats. Nevertheless, although more detailed and lengthy work is needed to confirm the results, they do shed some light on the problem under discussion. The reasons why E. nuttallii seems to be more vigorous than FE. canadensis in some habitats and not others also need to be determined and would be worthy of further investigation. ACKNOWLEDGMENT I thank Dr G. Halliday for his continued advice and encouragement throughout this work. DISPLACEMENT OF ELODEA CANADENSIS BY E. NUTTALLII 177 REFERENCES Barko, J. W. & SMart, R. M. (1981). Comparative influences of light and temperature on the growth of selected submersed freshwater macrophytes. Ecol. Monogr. 51: 219-235. Barko, J. W., Harbin, D. G. & MatrHews, M. S. (1982). Growth and morphology of submerged freshwater macrophytes in relation to light and temperature. Can. J. Bot. 60: 877-887. Briacos, M. (1977). Elodea nuttallii. B.S.B.I. News 15: 9-10. HAL Ler, W. T. & Sutton, D. L. (1975). Hyacinth Control J. 13: 48-50. Kuni, H. (1982). The critical water temperature for the active growth of Elodea nuttallii (Planch.) St John. Jap. J. Bot. 32: 111-112. Lunp, J. W. G. (1979). The mystery of Elodea Michx in Great Britain. Watsonia 12: 338. Simpson, D. A. (1983). Experimental taxonomic studies of Elodea Michx in the British Isles. Ph.D. thesis, University of Lancaster. Simpson, D. A. (1984). A short history of the introduction and spread of Elodea Michx in the British Isles. Watsonia 15: 1-9. Simpson, D. A. (1988). Phenotypic plasticity of Elodea nuttallii (Planch.) H. St John and Elodea canadensis Michx in the British Isles. Watsonia 17: 121-132. Titus, J. E. & Apams, M. S. (1979). Coexistence and comparative light relations of the submersed macrophytes Myriophyllum spicatum L. and Vallisneria americana Michx. Oecologia 40: 273-286. (Accepted December 1989) ul] s - = ; } ve a ~~ ; ia’ L, ry =| a oc ome Tih Pe a. ae aa? —~ “a ‘fie age rt ot sient: at hose ‘> - e 7 _ . 7 — ra ‘ Act i iw The I ? rh a SR ‘ ‘ = = te q = "4 F x "5 oN a r tar ne 9% neve ; ome vy a u2 A a - Ash d q rom aint Ayden Ppl! fa it ii! i nl nl : 7 j ad et ah : +, A ios 1% i s r ® Ya * ? o i. wonrrnre sisi 4 13 iw ni ee Sai b ‘ { ‘ antl 4 fi i ve | ~~ " on fea \ 1 “aat nighat veh ie eh pon tte: 5 , 2 ) et > ah - f i, ‘ = ~ ft mh } a ; if f ~ ] T yd og — ‘tay Watsonia, 18, 179-187 (1990) 179 Additions to the flora of Mull and adjacent small islands J. W. CLARK Tigh Sodlais, Onich, by Fort William, Inverness-shire, PH33 6SA and A. C. JERMY Natural History Museum, Cromwell Road, London, SW7 5BD ABSTRACT Vascular plant records, new to the modified 10-km squares used by the British Museum (Natural History) Survey of Mull and adjacent islands, made over the past ten years are listed. Thirty species and two hybrids of flowering plants, and one horsetail, are recorded as new to the archipelago, and two further species are confirmed. Eighteen species are recorded from the islands only for the second time, and eight species have been refound after a period of at least 40 years. INTRODUCTION The British Museum (Natural History) Survey of the Isle of Mull and adjacent small islands, carried out between 1966 and 1970, was published in 1978 (Jermy & Crabbe 1978). Between the time that the resulting Flora went to press and the end of 1988, when Joan Clark handed over the B.S.B.I. Recordership of v.c. 103, a number of vascular plant records have been made and are published here to chronicle the changes taking place in the flora of the island. The authors are not in a position to assess the changes in land-use that will have altered the ecology of the island. However, a few comments can be made. Cottages on the island are frequently bought and lived in by people from the mainland, who often develop or expand the small gardens, and a number of alien plants are thus introduced and may escape to become established locally: e.g. Ficus carica, Lysichiton americanus, Montia perfoliata, Poa chaixii, Sedum spurium and Soleirolia soleirolii. Another recorded alien is the North American Epilobium ciliatum, the seed possibly having been brought in with other garden plants. Forestry has taken its toll, like elsewhere in Western Scotland, and the maturation of trees in plantations has meant closer canopies and the elimination of a ground flora. The most serious change has been widespread planting by the Forestry Commission in the Coladoir River valley S.S.S.I., where the interesting bog complex has been divided and surrounded by trees. Similar large-scale planting on very wet mires in the Belat valley, Dervaig, will change the character of this extensive alluvial area which, although not outstandingly rich, is an area of floristic potential. With forestry planting comes the introduction of alien tree species, which may establish themselves locally and beyond. One such species, A/nus incana (L.) Moench, obviously planted in the Salen Forest around Loch Frisa (square 7), was recorded in 1979. Other species seen outside obviously planted areas, and possibly established from seed (or more likely planted by a landscape-conscious tenant/owner), and not hitherto recorded, are: Larix kaempferi (Lamb.) Carriére (11) Doire Darach, 1988, Picea abies (L.) Karsten (13) Rubha na Gall, 1987, Pinus contorta Douglas ex Loudon (11) Doire Darach, 1988, Quercus cerris L. (4) Ulva, 1987, Salix calodendron Wimmer (4) Ulva, 1983 and Tsuga heterophylla (Rafin.) Sarg. (11) Doire Darach, 1988. The density of sheep is still very high, and numbers of deer similarly so (P. Wormell, pers. comm.). There appears to be little management of the number of deer within or across estates. Fewer sheep and deer, and more cattle, would improve the botanical richness. Fish-farming is 180 J. W. CLARK AND A. C. JERMY established in and around Mull, as elsewhere in western Scotland, the largest salmon-rearing unit being in Loch Spelve. The effect of this on the marine flora, and eventually on the littoral vegetation, is an unknown quantity. The island has attracted a number of botanists, some on a regular basis, and a few resident, over the last ten years. This has resulted in 30 species and two hybrids of flowering plants (not including the trees listed above), and one horsetail (Equisetum hymale), being recorded as new to the area. A further 18 species, known previously from single records, have been recorded for the second time. Also of interest are the following species, refound after at least 40 years, or in need of confirmation for the island: Barbarea vulgaris (1879:1981), Hypericum maculatum subsp. maculatum (1889:1984), Malva sylvestris (1939:1977 & 1981), Polygonum bistorta (1879:1988), Rubus radula (1894:1984), Silene alba (1850:? 1945, confirmed 1987), Spergularia rubra (1877:? 1939, confirmed 1988) and Pseudorchis albida (1848:1988, refound in the area of Boswell-Syme’s original locality). In the following text, records new to the island are marked with an asterisk (*), and those recorded for a second time only with two asterisks (**). The order of the species (with the exception of Rubus, which follows Edees & Newton (1988)) follows Dandy (1958), with the later additions inserted appropriately. The system of modified 10-km square as used in the British Museum Survey is indicated throughout (in bold type). Voucher specimens, when available have been deposited in BM or E and indicated as such. Records made by Joan Clark are indicated by the initials JWC, and those made by Alan Stirling by AMcGS; other recorders’ names are given in full. SYSTEMATIC LIST FERNS AND FERN ALLIES Lycopodum clavatum L.:7. Speinne Mor, 1983, R. W. M. Corner. 10. Beinn na Croise, 1983, R. W. M. Corner. 14. Allt M6r Coire nan Eunachar, 1984, A. Kelham. Diphasiasirum alpinum (L.) Holub: 7. Speinne Mor, 1983, R. W. M. Corner. 10. Beinn na Croise, 1980, A. Keiham. Equisetum fluviatile L.: 4. Ditch, Ulva, 1988, A. Walker. E. hyemale L.: 6. Ardow burn, 1986, M. North, E. First localized record. Ophioglossum vulgatum L.: 4. Three clumps in short turf at separate sites, Gometra, 1988, M. Jones. Botrychium lunaria (L.) Swartz: 2. Loch Staoineig, 1980, H. Rhodes. 3. Ardalanish, 1984, A. Kelham. 11. Abundant in grassland on sand and shingle shore, Loch Ba, 1988, G. Rothero; on southwestern slopes of Ben More, 1988, L. Farrell & R. Scott. Cryptogramma crispa (L.) R.Br. ex Hook.: 8. Creach Bheinn, 1979, A. Kelham, BM. Asplenium adiantum-nigrum L.: Entry in Bangerter & Cannon (1978) should read 1-17. A. trichomanes subsp. quadrivalens D. E. Meyer emend. Lovis: 4. Farm dyke, Ulva, 1982, JWC. 14. Rock crevice, Garmony, 1981, JWC, BM. 17. Rock crevice, Leckruah, 1983, JWC. A. ruta-muraria L.: 14. Old chapel, Pennygown, 1981, JWC. Cystopteris fragilis (L.) Bernh.: 17. Port na Tairbeirt and south of Grass Point, 1984, AMcGS. Polystichum aculeatum (L.) Roth: 17. Woodland, Auchnacraig, 1988, A. Walker & E. Stewart. P. setiferum (Forsk.) Woynar: 7. Above Loch Frisa, 1983, R. W. M. Corner. Dryopteris filix-mas (L.) Schott: 11. Allt an Toll-dhoire, 1988, G. Rothero, det. A. C. Jermy. CONIFERS AND FLOWERING PLANTS Pinus sylvestris L.: 14. Hillside between Pennygown and Garmony. Juniperus communis L. subsp. alpina (Neilr.) Celak. (subsp. nana Syme): 14. On wall of ruined chapel, Pennygown, 1981, JWC. 17. Shore south of Grass Point, 1984, AMcGS. Trollius europaeus L.: 17. Beside burn, Leckruah, 1983, JWC. Ranunculus sceleratus L.: 1. Bac Mor, 1987, L. Farrell. 6. Near shore, Dervaig, 1987, B. Rae. 7. Muddy shore below irises, Lagganulva, 1988, JWC. Thalictrum alpinum L.: 10. Beinn na Croise, 1983, R. W. M. Corner. Papaver somniferum L.: 5. Schoolhouse, Kilninian, 1983, JWC. 13. Rubha nan Gall, 1982, A. Wright. In both instances plants appeared after cultivation of former garden ground. Meconopsis cambrica (L.) Vig.: Spreading from mainland, now widely naturalized. 3. Bunessan, ADDITIONS TO THE FLORA OF MULL 181 1981, JWC. 4. Near farm buildings, Ulva, 1982, JWC. 10. Shore, Pennyghael, 1981, JWC. 17. Garden, Torosay Castle, 1981, JWC. Dicentra formosa (Haw.) Walp. **: 17. Naturalized in a wood at Craignure, 1981, JWC, BM. Corydalis claviculata (L.) DC.: 4. Growing over broom and bracken by shore, Ulva, 1982, JWC, E. Fumaria bastardii Boreau: 3. Arable field, Ardalanish, 1981, JWC, BM and E. F. muralis Sonder ex Koch subsp. boraei (Jordan) Pugsley: 3. Garden weed, Erraid, 1988, JWC, det. M. Lidén, E. Fumaria officinalis L.: 4: Arable field, Ulva, 1982, JWC, BM. Brassica napus L.: 7: Waste ground, Lagganulva, 1982, JWC. B. rapa L.: 3. Waste ground, Ardalanish, 1981, JWC, BM. 4. Old garden, Gometra House, 1988, JWC. 9. Knockan, 1981, JWC, BM. Sinapsis arvensis L.: 3. Knockvologan, 1977, A. Kelham. 7. Lagganulva, 1982, JWC. 14. Corrynachenchy, 1981, JWC. Lepidium heterophyllum Benth.: 17. Shore, Java, 1984, AMcGS. Coronopus squamatus (Forskal) Aschers.: 2. In gravel, Baile Mor, Iona, 1987, JWC, E. Confirms previous records. Thlaspi arvense L. **: 6. Garden path, Croig, 1986, A. A. Slack. 13. Garden, Rubha nan Gall, 1984, A. Wright, E. Confirmation of 1967 record for this square. Capsella bursa-pastoris (L.) Medicus: 15. Garden, Scallastle, 1983, JWC. Cochlearia officinalis L. subsp. officinalis: 11. Shingle shore, Knock, 1987, JWC. C. danica L.: 3. Shore, Erraid, 1988, JWC. Cardamine hirsuta L.: 10: Waste ground, Pennyghael, 1981, JWC. Barbarea vulgaris R. Br. **: 12. Verge of A849, Glenforsa, 1981, JWC, E. 14. Verge of A849, near Pennygown chapel, 1981, JWC, E. Hesperis matronalis L.: 2. Field margin near church, Iona, 1979, JWC. Sisymbrium officinale (L.) Scop.: 3. Waste ground near garden, Erraid, 1988, JWC & A. Walker. 7. Abundant in arable fields, Lagganulva, 1982, JWC, E. Arabidopsis thaliana (DC.) Heynh.: 3. Sand-topped rocks, Fidden, and at Knockvologan, 1981, JWC, E. 6. Nursery garden, Glengorm Castle, 1987, JWC. 8. Allt na Teangaidh, 1981, A. Kelham. 9. Eas Dubh, 1982, A. Kelham. 14. Car park, Pennygown, 1981, JWC. 17. Waste ground near car- park, Duart, 1986, JWC. Polygala vulgaris L.: 11. North-western shore of Loch Ba, 1988, G. Rothero. Field record. Hypericum maculatum Cranz subsp. maculatum. **: 12. Verge of A84, Glenforsa, 1984, AMcGS & JWC, det. N. K. B. Robson, BM and E. Previous record (1889) was from Aros Castle. H. humifusum L.: 11. Doire Darach, 1988, L. Farrell & R. Scott. Silene latifolia Poiret subsp. alba (Miller) Greuter & Burdet (S. alba (Miller) E. H. L. Krause): 2. Bishop’s House garden and nearby waste ground, Iona, 1987, JWC. Considered in Jermy & Crabbe (1978, pp. 11, 13) as possible error for S. dioica (L.) Clairv.; now confirmed. Stellaria graminea L.: 10. On grassy waste ground, Pennyghael, 1984, JWC, E. Arenaria leptocladus (Reichb.) Guss. *: 3. On loose sand, Knockvologan, 1981, JWC, det. M. McCallum Webster, E. Spergularia rubra (L.) J. & C. Presl: 9. Forestry road near Shiaba, 1988, I. C. Christie. This species although listed by G. Ross, 1877, was considered an error by J. Cannon et al. This record now confirms this species for the island and it is possible that Ross was correct. Spergularia marina (L.) Griseb.: 1. On small rock outcrop, Bac Mor, 1987, L. Farrell. Montia fontana L. subsp. fontana: 4. Damp hollow in field, Ulva, 1982, JWC. M. perfoliata (Willd.) Howell *: 13. Garden, Rubha nan Gall, 1984, A. Wright, E. Probably brought in with mainland garden plants and now well-established. M. sibirica (L.) Howell: 17. Naturalized near houses, Java and Torosay, 1981, JWC. Chenopodium album L.: 17. Shore, Port nan Crullach, 1983, JWC, E. Beta vulgaris L. subsp. maritima (L.) Arcangeli: 1. Staffa, 1986, P. Wormell, E. Atriplex glabriuscula Edmondston: 11. Shingle shore, Knock, 1987, E. Stephenson, det. J. M. Mullin, BM. Malva sylvestris L. **: 2. By jetty, Iona, 1981, JWC, E. 3. Ardtun, 1977, A. Kelham, BM. Confirms unlocalized 1939 record. Aesculus hippocastanum L.: 9. West of Carsaig, 1977, A. Kelham. 182 J. W. CLARK AND A. C. JERMY Buxus sempervirens L. *: 14. Naturalized under brambles by field dyke, Garmony, 1981, JWC. Probably originated as seedling from old hedge in farm garden. Cytisus scoparius (L.) Link subsp. scoparius (Sarothamnus scoparius (L.) Wimm. ex Koch). 4. Near ferry, Ulva, 1982, JWC. 9. On verge of A849 east of Bunessan, 1977, A. Kelham. 17. On verge of A849 at Duart signpost, and also at Duart, 1981, JWC. Trifolium campestre Schreber: 12. Quarry track southwest of Salen, 1978, R. Coomber. T. dubium Sibth.: 15. On verge of A849, Alterich, 1983, JWC. Vicia orobus DC.: 4. Two sites on south coast of Ulva between Ulva House and Ormaig, 1980, A. Kelham. 8. On cliffs, Dun Bhuirg, 1988, P. Wormell. According to Bangerter & Cannon (1978) the record needed confirmation. V. sylvatica L.: 16. Loch Buie, 1956, H. Milne-Redhead; field card record omitted from Bangerter & Cannon (1978). V. sepium L.: 14. Corrynachenchy, 1981, JWC. V. angustifolia L.*: 3. Sandy shore, Ardalanish, 1981, JWC, BM and E. Rubus spectabilis Pursh: 7. Near farm, Lagganulva, 1982, K. Leitch. 12. In hedge near Kellan farmhouse, 1981, A. Kelham, det. A. Newton. R. plicatus Weihe & Nees: 15. Roadside, Torness, 1984, AMcGS. R. lindleianus Lees **: 2. Iona, 1987, JWC. 4. Ulva, 1982, JWC, E. 7. Southeast of Kilbrennan, 1982, JWC, E. 17. Craignure, 1984, AMcGS. R. dumnoniensis Bab.: 4. Ulva, 1982, JWC, E. 5. Calgary, 1984, AMcGS. 7. Achnacraig, 1984, AMcGS. 8. Balevulin, 1984, AMcGS. 9. Knockan, 1984, AMcGS. 10. Dererach, 1981, JWC; 11. Knock, 1984, AMcGS. 12. Salen, 1981, JWC, E. 13. Gualann Dhubh, 1984, AMcGS. 15. Torness, 1984, AMcGS. R. nemoralis P. J. Mueller (R. selmeri Lindeb.): 3. Riverbank, Bunessan, 1981, JWC, E. 5. Calgary, 1984, AMcGS. 6. Dervaig, 1984, AMcGS. 8. Dhiseig and Balevulin, 1984, AMcGS. 9. Knockan, 1981, JWC. 11. Knock, 1984, AMcGS. 13. Aros Park, 1984, AMcGS. 14. Fishnish, 1981, JWC. R. polyanthemus Lindeb.: 4. Ulva, 1982, JWC, E. 5. Calgary, 1982, JWC, E. 7. Rubha Mor, Loch na Keal, 1981, JWC, E. 8. Balevulin, 1984, AMcGS. 9. Knockan, 1984, AMcGS. 10. Near Rossal, 1984, JWC. 11. Loch na Keal, 1981, JWC, E. 13. Gualann Dhubh, 1984, AMcGS. 14. Fishnish, 1982, JWC, E. 15. Torness, 1984, AMcGS. 16. Croggan, 1983, JWC. 17. Craignure, 1984, AMcGS. R. septentrionalis W. R. C. Wats.: 2. Drainage ditch, Iona, 1987, JWC, E. 5. Calgary, 1984, AMcGS. 6. Dervaig, 1984, AMcGS. 10. Killiemore, 1984, AMcGS. 11. Knock, 1984, AMcGS. 12. Gruline, 1984, AMcGS. 13. Gualann Dhubh, 1984, AMcGS. 17. South of Grass Point, 1984, AMcGS. R. subinermoides Druce*: 4. Garden, Ulva House, 1982, JWC, E. 7. Near shore, Lagganulva, 1988, JWC, E. R. mucronulatus Borr.**: 4. Ulva, 1982, JWC, E. Gometra, 1988, JWC. 7. Achnacraig, 1984, AMcGS, E. 12. Gruline, 1984, AMcGS. 17. Port na Tairbeirt, 1984, AMcGS. R. radula Weihe ex Boenne. **: 9. Calgary, 1982, JWC, E. 8. Port Uamha Beathaig, 1984, AMcGS, E. 13. Aros Park, 1984, AMcGS, E. R. rufescens Muell & Lefév. *: 13. Aros Park, 1984, AMcGS. E. R. latifolius Bab. *: 4. Old garden, Gometra House, 1988, JWC. 6. Roadside, Croig, 1986, JWC. 8. Port Uamha Beathaig, 1984, AMcGS. 12. Roadside near Aros Castle, 1982, JWC, E. Potentilla sterilis (L.) Garcke: 13. Aros Park, 1986, JWC. Geum urbanum L.: 17. Woodland, Auchnacraig, 1988, A. Walker & E. Stewart. Alchemilla alpina L.: 3. Southwest of Ardalanish, 1977, A. Kelham. Unconfirmed field record. Rosa rubiginosa L.: 11. Sandy delta on northwestern shore of Loch Ba, 1988, G. Rothero, field record. Prunus spinosa L.: 14 Garmony, 1981, JWC. P. avium (L.) L.: 4. Ulva, 1982, JWC. 5. Treshnish House woods, 1983, JWC. 6. Dervaig, 1983, JWC. 7. Lagganulva, 1986, H. Leitch. 14. Garmony, 1981, JWC. 16. Croggan, 1983, JWC. P. padus L.: 5. Calgary and Tostarie, 1984, M. Barron. 8 Glen Seilisdeir, 1984, M. Barron. 11. Scarisdale, 1984, M. Barron. 14. Pennygown, 1984, M. Barron. Cotoneaster microphyllus Wall. ex Lindl.: 2: Cliff behind manse, Iona, 1979, JWC. 4. Widespread over an acre of hill behind Gometra House, 1982, J. M. Howard. 17. On rocks, Java Point, 1981, JWC., ADDITIONS TO THE FLORA OF MULL 183 Sorbus intermedia (Ehrh.) Pers.: 4. On cliffs, Gometra, 1988, A. Walker, det. AMcGS. S. croceocarpa P. D. Sell: 10. Pennyghael, 1971, A. G. Kenneth (recorded as S. Jatifolia in Bangerter & Cannon 1978). S. rupicola (Syme) Hedl. 17. Port na Tairbeirt, 1984, AMcGS. Additional site record. Sedum spurium Bieb. *: 7. In roadside rock crevice, Lagganulva, 1986, H. Leitch, E. Presumably escape from distant garden. S. acre L.: 11. Shore, Knock, 1987, JWC. Saxifraga stellaris L.: 10. Beinn na Croise, 1980, A. Kelham. S. x urbium D. A. Webb: 12. Glenforsa Lodge woods, 1981, JWC, BM. Saxifraga hirsuta L. **: 3. Roadside near car-park, Aros Park, 1986, JWC, E, (det. C. A. Stace). S. tridactylites L. **: 2. Rocks above Sandeels Bay, Iona, 1981, JWC, E. 3. Fidden rocks, 1981, JWC, E. S. aizoides L.: 11. Scarisdale river, 1983, R. W. M. Corner; Allt na Coille Moire, 1988, L. Farrell & R. Scott. S. oppositifolia L.: 11. Maol na Damh, 1988, A. & J. Gardner. Chrysosplenium oppositifolium L.: 1: Lunga, 1983, R. Coomber. Ribes rubrum L.: 13. Growing in garden wall, Rubha nan Gall, 1986, A. Wright. R. nigrum L.: 5. Reudle, 1983, JWC. 6. Druimghigha, 1986, M. North. 8. Balevulin, 1984, JWC. Lythrum salicaria L.: 17. Port na Tairbeirt, 1984, AMcGS. Epilobium hirsutum L. *: 13. Waste ground, Tobermory, 1978, R. Coomber, E. E. parviflorum Schreber: 11. Forestry plantation, Maol na Coille Méire, 1988, L. Farrell & R. Scott. E. ciliatum Rafin. *: 4. Ulva House garden, 1982, JWC, E. E. obscurum Schreber: 5. Haunn, 1983, B. Rae. 11. Forestry plantation, Maol na Coille Moire, 1988, L. Farrell & R. Scott. Chamerion angustifolium (L.) J. Holub (Epilobium angustifolium L.): 11. Loch Ba, 1988, A. Walker & E. Stewart; Forestry plantation, Maol na Coille Médie, 1988, L. Farrell & Scott. Fuchsia magellanica Lam.: 4. Naturalized, Ulva, 1982, JWC. 8. Tiroran, 1984, AMcGS. Circaea lutetiana L.: 11. Wooded ravine, Allt an Toll-dhoire, 1988, G. Rothero. Callitriche stagnalis Scop.: 1. Staffa, 1986, P. Wormell. 14. Garmony, 1981, JWC. C. hamulata Koch (C. intermedia subsp. hamulata Koch): 2. Pool on top of Dun I, Iona, 1979, JWC, BM, E. Eryngium maritimum L.: 1. Lunga, 1983, B. Rae. 2. At time of British Museum Survey not seen on Iona, but refound at the north end of the island in 1979 by H. Rhodes. Anthriscus sylvestris (L.) Hoffm.: 3 Cul a’Bhaile, 1984, A. Kelham. Conium maculatum L.: 12. Arla farm, 1978, R. Coomber. Heracleum sphondylium L.: 14. Widespread, 1981, JWC. H. sphondylium L. var. angustifolium Huds. *: 2. Jetty, lona, 1987, JWC, E. Euphorbia dulcis L. *: 12. Glenforsa Lodge woods, widely naturalized, 1981, JWC, E. Polygonum aviculare L. sensu lato: 11. Shore, Knock, 1987, JWC. P. bistorta L. **: 4. Ulva, 1988, JWC. Well established on waste ground beside farm road. Probable escape from Ulva House garden. Polygonum amphibium L.: 4. Shore, Ulva, 1982, JWC. 17. Rubha na Sroine, 1981, JWC. P. hydropiper L.: 8. Ardmeanach, 1984, AMcGS. 16. Leckruah, 1983, JWC. Polygonum campanulatum Hooker fil.: 8. Tiroran, 1984, AMcGS. Reynoutria sachalinensis (F. Schmidt Petrop.) Nakai (Polygonum sachalinense F. Schmidt Petrop. ): 12. Verge of B8073 at head of Loch na Keal, and at Aros, 1987, JWC. 17. Torosay Castle woods by railway, 1987, JWC. R. japonica Houtt. (Polygonum cuspidatum Sieb. & Zucc.): 4. Near church, Ulva, and roadside, Gometra, 1988, JWC. Rumex longifolius DC. *: 10. Roadside, Glen Leidle, 1984, AMcGS, E. A new vice-county record. R. obtusifolius L.: 14. Widespread, 1981, JWC. R. sanguineus L. var. viridus Sibth.: 7. Lagganulva, 1982, JWC, E. Parietaria judaica L. (P. diffusa Mert. & Koch) *: 17. Wall, Duart Castle, 1983, R. W. M. Corner, E. Soleirolia soleirolii (Req.) Dandy *: 4. In cracks in steps and round base of Gometra House, 1988, M. Jones, E. Introduced. 184 J. W. CLARK AND A. C. JERMY Humulus lupulus L. *: 6. Ruins of clachan in Glengorm afforested area, 1962, A. Wright (photo, BM). Ulmus glabra Huds.: 9. Near Scoor, 1978, A. Kelham. Ficus carica L. *: 13. Rock cleft near gun emplacement, Tobermory, 1983, A. Wright. Alnus incana (L.) Moench: 7. Moribund plantation, Loch Frisa, Salen Forest, 1979, A. Kelham. Populus tremula L.: 17. Cliffs south of Grass Point, 1984, AMcGS. P. nigra agg. **: 8. Clachandhu, 1982, A. Kelham. 14. Garmony, 1981, JWC, BM. Salix alba L.: 8. Kilfinichen, 1984, AMcGS, E. S. herbacea L.: 5. Carn Mér, 1983, R. W. M. Corner. 7. Speinne Mor, 1983, R. W. M. Corner. 10. Beinn na Croise and An Sgulan, 1983, R. W. M. Corner. Rhododendron penticum L.: 4. Ulva, 1982, JWC. Arctostaphyllos uva-ursi (L.) Spreng.: 10. Island in Loch Fuaron, 1977, A. Kelham. Orthilia secunda (L.) House **: 13. Near Sput Dubh, 1981, D. Kelly. Empetrum hermaphroditum Hagerup: 11: Beinn a’Ghraig, 1983, R. W. M. Corner. Trientalis europaea L. *: 16. Above birch wood in flattish area of bracken and heather, A’Bhuigneach, 1987, A. Gardner. 17. Flat ground with Sphagnum and young birch trees, Java Point, 1981, JWC, BM and E. Anagallis tenella (L.) L.: 17. Port Donain, 1978, R. Coomber. A. arvensis L.: 4. Gometra, 1982, J. E. Howard; arable field, Ulva, 1982, M. Cowe, E. Gentianella campestris (L.) Borner: 1. Staffa, 1986, P. Wormell. Symphytum officinale L.: 3. Waste ground near coastguard cottages, Erraid, 1988, JWC & A. Walker. 4. Naturalized, old garden, Ulva, 1982, JWC, E. S. X uplandicum Nyman: 3. Waste ground behind croft house, Erraid, 1988, JWC & A. Walker. S. tuberosum L. **: 14. Near jetty, Fishnish, 1981, JWC, E. Borago officinalis L. *: 2. Waste ground and gardens in Baile M6r, Iona, 1987, JWC. Introduced for bees, and now spreading widely. Pentaglottis sempervirens (L.) Tausch: 6. Garden escape, Glengorm Castle, 1987, JWC. 17. Naturalized outside garden of Duart Castle, 1981, JWC. Mertensia maritima (L.) Gray: 4. Little Colonsay, 1982, J. M. Howard. 5. Only one small plant seen at Calgary, but colony at Ensay Burn luxuriant and spreading. 12. North shore of head of Loch na Keal, 1986, R. Coomber. 11. Extensive stretch on spit at head of Loch na Keal, north of fish hatchery, 1988, JWC. Calystegia sepium (L.) R.Br.: 2. Baile M6r, Iona, 1987, JWC. 4. Farm road, Ulva, 1982, JWC, E. On shore, Gometra, 1988, M. North. 17. Waste ground, Java, 1987, JWC. C. pulchra Brummitt & Heywood: 7. Lagganulva, 1982, H. Leitch. Solanum dulcamara L. **: 17. Shore, Port Donain, 1978, R. Coomber. Scrophularia nodosa L.: 17. Port na Tairbeirt, 1984, AMcGS. Mimulus moschatus Douglas ex Lindley: 17. Waste ground, Java, 1984, AMcGS. Sibthorpia europaea L. *: 11. Beside track between Mausoleum and Gruline House, 1988, A. Walker & E. Stewart, GL. Probably introduced. Veronica scutellata L.: 3. In flush, Erraid, 1988, A. Walker. V. montana L.: 17. Wood edge, Port Donain, 1978, R. Coomber. V. agrestis L.: 3. Island House, Dunan Mor, 1977, A. Kelham. V. filiformis Sm.: 3. Lawn, Bunessan, 1981, JWC, E. 15. Lawn, Scallastle, 1983, JWC. Rhinanthus minor L.: 1. Bac Mor, 1987, L. Farrell. R. borealis (Sterneck) P. D. Sell: 11. Scarisdale river, 1983, R. W. M. Corner. Odontites verna (Bellardi) Dumort.: 1. Lunga, 1983, B. Rae. 11. Shore, Knock, 1987, JWC. Orobanche alba Willd.: 8. South Ardmeanach, near shore, 1976, P. Wormell. Mentha arvensis L.: 7. Field edge, Torloisk, 1982, JWC, E. Mentha X piperita L. *: 12. Marsh by shore, Salen, 1983, R. W. M. Corner, det. R. M. Harley. M. suaveolens Ehrh. *: 13. Naturalised, Rubha nan Gall, 1984, A. Wright, det. R. M. Harley. M. x villosa Huds. nm. alopecuroides (Hull) Briq. *: 2. Martyrs’ Bay, Iona, 1981, JWC, det. R. M. Harley, E. 12. Roadside verge, Salen, 1983, R. W. M. Corner, det. R. M. Harley. 17. Grass Point and Java, 1984, AMcGS. Lycopus europaeus L.: 14. By jetty, Fishnish, 1981, JWC. Stachys palustris L.: 11. Field verge, Knock, 1987, JWC. ADDITIONS TO THE FLORA OF MULL 185 S. x ambigua Sm.: 17. By shore, Java, 1984, AMcGS, E. Lamium amplexicaule L.: 6. Garden, Druimghigha, 1986, M. North. L. hybridum Vill. **: 3. Garden, coastguard cottages, Erraid, 1988, JWC & A. Walker, E. L. album L.: 6. Glen Gorm Nursery Garden, 1984, B. Rae. Scutellaria galericulata L.: 15. Shore, Alterich, 1983, JWC. Campanula latifolia L.: 4. Garden escape, roadside, Ulva, 1982, JWC. Sherardia arvensis L.: 9. Near Scoor, 1981, M. Collier, E. Galium album Miller: 12. On verge of A849, Pennygown, 1984, AMcGS, E. G. uliginosum L.: 11. By burn, Airigh Lag nan Cluas, 1988, L. Farrell & R. Scott. Sambucus nigra L.: 14: Corrynachenchy, 1981, JWC. Symphoricarpos albus (L.) S. F. Blake var. laevigatus (Fernald) S. F. Blake (S. rivularis Suksd.): 8. Beside B8035, Balevulin, 1984, JWC. 14. Corrynachenchy, 1981, JWC. Valerianella locusta (L.) Laterrade: 16. Loch Buie, 1979. R. Coomber. Dipsacus fullonum L. *: 6. Casual on waste ground, Croig, 1983, B. Rae. Senecio sylvaticus L.: 12. Quarry track south-west of Salen, 1979, R. Coomber. 17. Pier, Craignure, 1985, C. Murray. S. viscosus L.: 6. Waste ground, Dervaig, 1983, B. Rae. S. vulgaris L.: 14. Pennygown, 1981, JWC. 15. Garden, Alterich, 1983, JWC. Tussilago farfara L.: 11. Forestry plantation, Maol na Coille Moire, 1988, L. Farrell & R. Scott. 14. Pennygown, 1981, JWC. Inula helenium L.: 5. Roadside near school, Kilninian, 1983, JWC. Eupatorium cannabinum L.: 17. South of Grass Point, 1984, AMcGS. Tripleurospermum maritimum (L.) Koch: 11. Shore, Knock, 1988, JWC. 15. Scallastle, 1983, JWC. Matricaria matricarioides (Less.) Porter: 15. Alterich, 1983, JWC. Tanacetum parthenium (L.) Schultz Bip.: 15. Alterich, 1983, JWC. 17. Waste ground, Java, 1983, JWC. T. vulgare L.: 17. Beside old house, Grass Point, 1984, JWC. Arctium minus Bernh. subsp. nemorosum (Lej.) Syme.: 14. Forestry roadside, Fishnish, 1981, JWC. Cirsium helenioides (L.) Hill (C. heterophyllum (L.) Hill) 17. Verge of A849, Torosay, 1984, AMcGS. Leontodon taraxacoides (Vill.) Mérat: 13. Car-park, Aros Park, 1984, AMcGS. Sonchus arvensis L.: 8. Shore, Balmeanach, 1984, JWC. 11. Shore near Knock, 1987, JWC. S. asper (L.) Hill: 11. Forestry plantation, Maol na Coille Moéire, 1988, L. Farrell & R. Scott. Hieracium rubiginosum F. J. Hanb.: 11. Beinn Fhada, 1967, A. Kenneth, CGE. Omitted from Flora of Mull (Bangerter & Cannon 1978). H. pilosella L. (Pilosella officinarum C. H. & F. W. Schultz subsp. officinarum): 17. Shore, Grass Point, 1984, AMcGS. Crepis capillaris (L.) Wallr.: 11. Shore near Knock, 1987, JWC. 14. A849 verge, Garmony, 1981, JWC. Taraxacum duplidentifrons Dt. (= T. raunkiaerii Wiinst): 2. Dunes and machair, Iona, 1989, JWC. 3. Sea wall, Bunessan, 1981, JWC, E. 17. Java Point and Craignure, 1981, JWC. Taraxacum euryphyllum (Dahlst.) M. P. Christensen: 3. Uisken, 1981, JWC, E. T. landmarkii Dahlst.: 17. Java Point, 1981, JWC, E. T. maculosum A. J. Richards (T. maculigerum H. Lindb. f.): 3. Machair, Fidden, 1981, JWC, E. 14. Fishnish, 1981, JWC, E. T. spectabile Dahlst.: 10. Pennyghael, 1981, JWC, E. T. unguilobum Dahlst.: 14. Fishnish, 1981, JWC, E. T. stictophyllum Dahlst. *: 17. Torosay Castle grounds, 1981, JWC, E. Triglochin maritima L.: 15. Shore, Alterich, 1983, JWC. Polygonatum multiflorum (L.) All.: 4. Garden escape, Ulva, 1982, M. Cowe. Scilla verna Huds.: 3. Fidden, 1972, C. Murray. 9. Garbh Eilean, 1981, J. Edwards. Juncus squarrosus L.: 4. Moor, Ulva, 1982, JWC. J. triglumis L.: 11. Maol Meadhonach, 1988, A. Gardner. Allium ursinum L.: 14. Doire Dorch, 1981, JWC. Leucojum vernum L. *: 13. Naturalized in glen, Tobermory, 1979, R. Coomber, E. 186 J. W. CLARK AND A. C. JERMY Cephalanthera longifolia (L.) Fritsch: 13. Loch a’Churrabain, 1979, R. Coomber. Listera ovata (L.) R.Br.: 3. Dunes, Traigh Gheal, Erraid, 1988, A. Walker. 17. Meadow, Auchnacraig, 1988, A. Walker & E. Stewart. Neottia nidus-avis (L.) Rich.: 10. Under lime trees above Carsaig Bay, 1985, A. Gardner. 17. Hazel/ ash wood, Druim Mor Aird na Drochaide, 1987, E. Campbell & A. Walker. Pseudorchis albida (L.) A. & D. Léve: 3. Ben Talaidh, 1977, A. Kelham. 9. Knockan, 1981, JWC. 16. Ardura, Loch Spelve, 1988, A. & J. Gardner. Previous record 1848. 17. Abundant on hill slope, Gorten, 1981, JWC, E. Platanthera chlorantha (Custer) Reichenb.: 6. Grassy slope, Croig, 1983, B. Rae. Orchis mascula L.: 13. Riverbank, Tobermory, 1976, R. Coomber. Dactylorhiza fuchsti (Druce) S60: 17. Meadow, Auchnacraig, 1988, A. Walker & E. Stewart. D. incarnata (L.) So6 subsp. incarnata: 7. Meadow, Oskamull, 1983, H. Leitch. D. incarnata (L.) So6 subsp. pulchella (Druce) H. Harrison fil. *: 17. Port na Tairbeirt, 1984, AMcGS, E. D. incarnata (L.) So6 subsp. coccinea (Pugsl.) H. Harrison fil. *: 5. Machair, Port Langamull, 1984, R. Coomber. Lysichiton americanus Hulten & St. John *: 14. Shore among irises at burn outflow, Fishnish, 1988, B. Rae & R. Douglas. Arum maculatum L. **: 13. One plant on shore near pier, and colony of many hundreds on wooded slope below Courthouse beside Prison Brae, 1986, A. Wright, E. Scirpus maritimus L.: 9: Brackish pool, Port nan Droigheann, 1988, J. Muscott & J. Winham. S. tabernaemontani C. C. Gmel: 9. Freshwater marsh, Port nan Droigheann, 1988, J. Muscott & J. Winham. Eleocharis uniglumis (Link) Schultes: 11. Airigh Lag nan Cluas, 1988, L. Farrell & R. Scott. Blymus rufus (Huds.) Link: 9. Shore, Knockan, 1981, JWC. Rhynchospora alba (L.) Vahl: 6. Druimghigha, 1986, M. North. Carex lepidocarpa Tausch *: 4. Near shore, Ulva, 1982, BM and E. 11. Grassy sand/shingle delta, northwest shore Loch Ba, 1988, G. Rothero. Field record. C. viridula Michx subsp. viridula (C. serotina Mérat): 17. Shore, Craignure, 1984 AMcGS, E. C. extensa Gooden.: 3. Shore, Erraid, 1988, JWC & A. Walker. 8. Balmeanach, 1984, AMcGS. C. sylvatica Huds.: 9. Cliff scrub, Port nan Droigheann, 1988, J. Muscott & J. Winham. 11. Wood, Glen Clachaig, 1988, A. Walker & E. Stewart. 17. Woodland, Auchnacraig, 1988, A. Walker & E. Stewart. C. vesicaria L.: 6. Silted-up dam, Quinish gardens, 1986, M. North. C. bigelowii Torr. ex Schwein: 10. Beinn na Croise, 1983, R. W. M. Corner. Carex paniculata L.: 7. East side of Carn Mor, 1986, M. North. 9. Freshwater marsh, Port nan Droigheann, 1988, J. Muscott & J. Winham. C. remota L.: 17. Druim Mé6r Aird na Drochaide, 1987, A. Walker. C. curta Gooden: 10. Beinn na Croise, 1985, A. Kelham. Glyceria declinata Bréb.: 8. Balmeanach, 1984, AMcGS. Festuca arundinacea Schreber: 5. Calgary, 1984, AMcGS. F. tenuifolia Sibth.: 7. Gravel bank, Fanmore, 1982, JWC; E. Puccinellia distans (L.) Parl.: 17. Shore, Java, 1984, AMcGS. Poa angustifolia L. **: 1. Found in dense grassland on Staffa, 1969, British Museum Survey, BM. Omitted in error from Cannon & Bangerter (1978). P. subcaerulea Sm.: 4. Shore track, Ulva, 1988, A. Walker. P. chaixii Vill. *: 17. Naturalized on pond edge, Torosay gardens, 1987, JWC, E. Catabrosa aquatica (L.) Beauv. subsp. minor (Bab.) Perring & Sell: 7. Shore, Lagganulva, 1988, JWC. Bromus ramosus L.: 17. Woodland, Auchnacraig, 1988, A. Walker & E. Stewart. B. hordeaceus L. subsp. hordeaceus: 15. Waste ground, Scallastle, 1983, JWC. Elymus caninus (L.) L. (Agropyron caninum (L.) Beauv.): 7. Ash wood, Acharonach, 1988, A. Walker & E. Stewart. Deschampsia cespitosa (L.) Beauv., viviparous variant *: 11. Scarisdale river bank, 1983, R. W. M. Corner. ADDITIONS TO THE FLORA OF MULL 187 Aira caryophyllea L. subsp. multiculmis (Dumort.) Aschers. & Graebn. ex Hegi *: 5. Roadbank, Kilninian, 1982, JWC, E. Calamagrostis epigejos (L.) Roth: 11. Wet woodland northwest shore of Loch Ba, 1988, G. Rothero. One plant only. Alopecurus pratensis L.: 4. Roadside, Ulva, 1982, JWC. Phalaris arundinacea L.: 14. Corrynachenchy, 1981, JWC. ACKNOWLEDGMENTS The authors would like to thank the following for identification of critical genera: Rubus, E. S. Edees (up to 1984) and A. McG. Stirling; Taraxacum, A. J. Richards and the late C. C. Haworth; Mentha, R. M. Harley; and J. M. Mullin for checking nomenclature. We would also like to thank Peter Wormall who gave us information on land-use changes. Joan Clark thanks all those visiting or living on Mull, Iona, Ulva and Gometra, who helped by gathering and sending in new records, and keeping in touch; and especially her co-author Clive Jermy, without whose help and encouragement this list would not have been attempted nor polished for publication. REFERENCES BANGERTER, E. B. & CANNON, J. M. F. (1978). Flowering plants and conifers, in JeErMy, A. C. & CRABBE, J. A., eds. The Island of Mull: a survey of its flora and environment. London. Danpy, J. E. (1958). List of British vascular plants. London. JerMYy, A. C. & CRABBE, J. A., eds. (1978). The Island of Mull: a survey of its flora and environment. London. EpEEs, E. S. & Newton, A. (1988). Brambles of the British Isles. London. (Accepted February 1990) : : ’ “ f ' q a Gre exe wes = ahiie ben. ) : ‘ wed +. ; : alee ‘ . , a 7 4. i ict, -& : . as ’ Laat ar iS a3 B.S y : 7 ‘if? SA ’ : + . . — ss 0 | Ems E at tik ora teed ebrem ‘ ” F - “ve. é 4 7 * fit cat rn Se SITIO 3 way = ad F i oad *c, prt in taut al pol eee + a eae p! ed Tiere alt bas..aiocs on im Sass VER esanbeg 7G Kp ioe is errs 4 wolhe tendiive re evil 1 Dhe-0o ye xin . : F ~ % . . pa = at ) , ay Soli. stu wih Deleted 100 Hur Salad Q r y ’ : 4 a "1 ca leg " . 4 . Ls : é & 7 i i] a>. - a ifn. hs saa ‘X we ¢ € h ts ria MvsS LAS wood ea rs ah re ey tHicick Pe Paes s " Lint hd ‘ieee 7 y*) ey > uti wi. io eel ‘ as ve | ol ot £2 158 wre r 3 aA) 5 med : = —- % 75) or freer PAWS) 3 : ‘ ‘ \ 4 3 - 4 i ¢ ‘ os | = » * i ' — « b . 3 mé n a* / , ..% Vv _ ‘ " : @ <4 > th etiah fh vane -- : a = 4 * & @ _ ‘ a , _ iP av Watsonia, 18, 189-198 (1990) 189 Five brambles from Wales A. NEWTON 10, The Fairways, Leamington Spa, Warwickshire, CV32 6PR and M. PORTER Aberhoywy, Cyffredin Lane, Llangynidr, Crickhowell, Powys, NP8 1LR ABSTRACT The history of the study of brambles in Wales is summarised and five new species are described: Rubus tavensis Newton & Porter, sp. nov., R. aquarum Newton & Porter, sp. nov., R. gallofuscus Newton & Porter, sp. nov., R. biloensis Newton & Porter, sp. nov., and R. merlini Newton & Porter, sp. nov. INTRODUCTION Valuable pioneering studies of the Welsh brambles were made in the last decade of the nineteenth century by J. W. Griffith in the Bangor district, A. Ley, who collected in many parts of Wales and the Marches and W. M. Rogers who visited the upper Wye Valley and adjacent areas in 1898. Riddelsdell (1907), assisted by Rogers and Ley, compiled a lengthy Glamorgan list and with W. C. Barton visited the Dolgellau and Portmadoc areas in 1922-23, making extensive collections of the local plants (now in BM). The results of these endeavours can be studied in most of the major British herbaria and provide a solid foundation for systematic enquiry. Knowledge of the Welsh Rubus flora has steadily increased since the brief visit of Watson (1950). A Welsh bramble foray (Newton 1972) described the fruits of a tour through northern and central Wales. Specimens collected by T. A. W. Davis in Pembrokeshire and Carmarthenshire were annually examined by E. S. Edees and A. N. and further surveys of South Wales produced new taxa described by Newton (1974) and an account of the Rubi of Glamorgan and adjacent areas (Newton 1976). In 1978 a B.S.B.I. field meeting based at Lampeter added considerably to the records for Cardiganshire and northern Carmarthenshire, updating the account of Rubus in Salter (1935). Visits to North Wales were made sporadically and the accumulated wisdom of these excursions, together with Welsh records from NMW and other herbaria, were collated by A. N. in Ellis (1983), which included some distribution maps. Also incorporated were records from M.P. who had been investigating the Rubi of Breconshire and gathering information for a county Flora since 1970. A joint visit in 1983 allowed many of the problematic local species of Breconshire to be studied in situ. The B.S.B.I meeting based at Carmarthen reported by Pryce (1988) provided information from previously unrecorded areas, which was included in the monograph by Edees & Newton (1988). Gradually it has become possible to discern the extent of some recognised but hitherto unnamed taxa and to establish new distribution data for others. Even now not all Welsh brambles can be satisfactorily named, but few of these are other than single bushes or of limited local occurrence. The following descriptions are therefore the outcome of lengthy and painstaking study in the field and herbarium. Rubus tavensis Newton & Porter, sp. nov. (Series Rhamnifolii) Turio altiarcuatus in apricis purpureus pruinosus angulatus faciebus planis leviter pilosus aculeis ad angulos limitatis aequalibus validis patentibus rectis vel raro leviter curvatis e basi compressa 190 A. NEWTON AND M. PORTER paulatim contractis armatus. Folia quinata pedata vel subdigitata pusilla inferne griseoviridia pilosa; foliolum terminale late obovatum vel suborbiculare apice cuspidatum basi integra margine aequaliter serrata longe petioluatum. Inflorescentia longa angustata inferne 2—4 foliis ternatis superne 1-2 foliis simplicibus ornata; ramuli medii + patentes 2-5 flori alte divisi. Rachis vix flexuosa dense pilosa aculeis curvatis validis nonnullis armata. Flores 2-5 cm diam.; sepala reflexa tomentosa grisea; petala rosea obovata pubescentia; stamina stylos virides basi roseos vix superantia; anthera glabra vel sparsim pilosa; carpella pilosa; fructus parvus rotundus. Stem arching, dull purple on exposed side, pruinose, angled with flat sides, very thinly hairy with few short simple or tufted hairs and more frequent stellate hairs, scattered sessile or subsessile glands, prickles (¢c. 10 per 5 cm) on the angles, equal, strong, about as long as the stem diameter, straight or rarely slightly curved, + patent, dull purple with straw coloured tips. Leaves 5-nate, pedate or subdigitate, leaflets rather small, usually not contiguous, dull green and glabrous or sparsely strigose above, pale greyish-green and hairy beneath with numerous short or very short simple hairs, sparse to frequent stellate hairs and occasional inconspicuous sessile glands; terminal leaflet broadly obovate or suborbicular with entire or, rarely, emarginate base and shortly cuspidate tip; margin flat or slightly undulate, finely and evenly serrate, petiolule long (c. 45% length of lamina); intermediate leaflets only slightly smaller than the terminal; basal leaflets with petiolules 3— 4 mm; petioles about the same length or longer than basal leaflets, coloured and clothed like the stem and with 12-15 small falcate prickles (2-3 mm). Flowering branch long and cylindrical, with 2— 4 ternate leaves at the base subtending ascending peduncles usually shorter than the leaves, and 1-2 simple leaves, not leafy to the apex; peduncles of the ultra-axillary part almost patent, divided well above the middle and bearing 2-5 flowers. Rachis slightly flexuose, densely hairy with short, patent, simple or tufted whitish hairs, stellate hairs, some inconspicuous sessile or subsessile glands, strongly armed with stout curved prickles (4~7 mm); peduncles and pedicels clothed as rachis but with smaller curved prickles. Flowers about 2-5 cm in diameter; sepals reflexed, greyish felted, with simple hairs and dense stellate hairs; petals pink or pale pink, obovate, c. 13 x 8 mm, downy; stamens about the same length or rather longer than the styles, anthers glabrous or sparsely hairy, filaments pink or white; styles pale green or pink-based; young carpels hairy; receptacle glabrous; fruit round, small with few drupelets. Flowering in July and August. 1 2 3 4 Ficure 1. Distribution of Rubus tavensis Newton & Porter FIVE BRAMBLES FROM WALES 191 Ho otype: near Wern Plemys wood, Ystradgynlais, Brecs., v.c. 42, GR 22/790.095, 21 July 1988, M. Porter (NMW). This bramble was named Rubus godroni Lec. & Lam. var. foliolatus by W. M. Rogers and A. Ley. Their very brief description in J. Bot. 44: 58 (1906) forms the basis for the entry in the addendum of Edees & Newton (1988), but also covered originally the Cheshire bramble subsequently described as Rubus robii (W. C. R. Watson) Newton. Rogers and Ley noted that the plant was first observed at Aberpergwm in the Nedd valley about 1890. The earliest herbarium specimen we have traced is at BIRM, collected by A. Ley on 21 July 1899 from Ystalyfera in the Tawe valley. The bramble is frequent in the south-west of Brecs. (v.c. 42) and adjacent parts of Glam. (v.c. 41) and is a particular feature of the valleys of the Tawe and Nedd. The long narrow leafy panicle and the small leaflets are distinctive features. Representative exsiccata: Glam., v.c. 41, Neath, 25 July 1905, H. J. Riddelsdell (NMW). Brecs., v.c. 42, Ystalyfera, 21 July 1899, A. Ley (BIRM); Ystradgynlais, 25 July 1899, A. Ley (LIV); Penwylit, 4 August 1899, A. Ley (BIRM); Cellwen, 2 August 1906, A. Ley (BIRM); Pen-y-Cae, 27 July 1977, M.P. (herb. M.P.); Cwm Twrch, 1 August 1977, M.P. (herb. M.P.); Abercraf, 17 August 1978, _M.P. (herb. M.P.); Cwm Giedd, 27 July 1982, M.P. (herb. M.P.). The known distribution is shown in Fig. 1. Rubus aquarum Newton & Porter, sp. nov. (Series Mucronati) Turio primo arcuatus tandem procumbens faciebus planis vel concavis in apricis erubescens vel fusco-purpureus angulatus fere glaber glandulis aciculisque sparsis brevibus, aculeis plerumque ad angulos limitatis rectis tenuibus e basi vix compressa et aculeolis brevioribus raris obsitus. Folia (3—) S5nata pedata vel subdigitata inferne sparsim pilosa; foliolum terminale obovatum basi subcuneata apice cuspidatum margine denticulata. Inflorescentia diffusa inferne foliis ternatis (foliolo medio basi cuneata), superne uno folio ovato ornata apice aphylla + pyramidata pedunculis pedicellisque longis; ramuli medii 1-4 flori. Rachis flexuosa aculeis tenuissimis falcatis declinatisve armata superne dense pilosa et tomentosa glandulis brevibus aciculisque nonnullis obsita. Flores (2—)2-5—3 cm, in umbrosis minores stellati, sepala reflexa griseoviridia tomentosa glandulifera aciculata; petala obovata roseo-lilacina ciliata; stamina stylos superantia, anthera pilosa; filamenta lilacina basi vinoso-purpurea. Carpella pilosa; fructus subglobosus. Stem low-medium arching at first, then trailing or scrambling, angled with flat or slightly furrowed sides, reddish-brown or purplish-brown in exposure, obscurely striate, almost glabrous but with rare short simple or tufted hairs, locally frequent sessile glands and occasional short stalked glands and acicles; prickles (c. 10 per 5 cm) mostly on the angles, slender, declining from a slightly compressed base, coloured as the stem or with yellow tips, + equal (c. 5-6 mm) but with an occasional shorter prickle. Leaves pedate or subdigitate; leaflets (3-)5, not contiguous, upper surface dark green, thinly strigose or almost glabrous, sometimes weakly plicate, lower surface sparsely clothed with short and very short simple hairs and sessile glands. Terminal leaflet c. 7 x 4cm, obovate with sides tapering to a narrow rounded or cuneate base, apex cuspidate, margin flat or undulate with fine slightly irregular teeth; petiolule c. 30-40% length of lamina. Petiole as long or longer than the basal leaflets, coloured as the stem, with about 12-15 small falcate prickles and sparsely clothed with simple hairs and sessile glands, few short or very short stalked glands, rare acicles and some stellate hairs. Inflorescence diffuse and showy, with 2-4 ternate leaves having markedly cuneate central leaflets, subtending ascending peduncles as long as or shorter than the leaves, usually one broadly ovate simple leaf, not leafy to the apex; the upper part of the inflorescence more or less pyramidal with long peduncles and pedicels so that the flowers are well spaced, middle peduncles with 14 flowers. Rachis flexuose, with a few very slender falcate or declining prickles (to 4 mm), rather thickly clothed especially on the upper part with frequent short simple or tufted hairs and stalked glands mostly shorter than the hairs, a few acicles and dense grey felt; pedicels mostly 1-2 cm, clothed as upper rachis but with more stalked glands and acicles exceeding the short hairs and with a few straight, patent, very fine pricklets (up to three times the diameter of the pedicel). Flowers (2-) 192 A. NEWTON AND M. PORTER 2-5-3 cm diameter (smaller in shade), starry; sepals reflexed, greyish-green, closely felted and with short mainly adpressed simple hairs, few to many acicles and short stalked glands; petals c. 14 x 7 mm, obovate, narrowed to a short claw, lilac-pink, hairy and with ciliate margins; stamens longer than styles, anthers cream with lilac sutures, pilose; filaments lilac shading to bright reddish-purple at the base; styles biscuit or pale green; young carpels hairy; receptacle hairy; fruit subglobose of about 20-30 drupelets. Flowering in July and August. Ho otyPe: hedge west of Llanwrtyd Wells, Brecs., v.c. 42, GR 22/872.459, 18 July 1988, M. Porter (NMW). A. Ley first collected this bramble from a wood at Llanwrtyd Wells in 1897. It was determined by W. O. Focke as Rubus lejeunei W. & N. and appears thus in Rogers (1900), although the author was unusually sceptical about Focke’s naming in this case. Later, the same plant, which Ley had gathered from several places in northern Breconshire, was equated with Rubus breconensis W. C. R. Watson (Watson 1958). Subsequently both Edees and Newton noted that Ley’s bramble did not match the type specimen of R. breconensis in BM. Although the taxa have some affinity, especially in floral characters, the northern Breconshire plant has almost glabrous stems with rare acicles and fewer prickles which are more or less equal and restricted to the angles. The inflorescence is more spectacular having larger well-spaced lilac flowers with conspicuous magenta filaments. Because it is such a distinctive plant and features prominently in the literature it was thought desirable to describe it, in spite of a limited distribution. As it has been found across northern Breconshire in a zone which includes the Victorian spas of Llanwrtyd, Llangammarch and Builth with their associated mineral springs we have named it Rubus aquarum. Representative exsiccata: Brecs., v.c. 42, Llanwrtyd Wells, 14 July 1897, A. Ley (BIRM, NMW); Builth Wells, 9 August 1898, A. Ley (BIRM); Llangammarch Wells, 25 July 1907. A. Ley (BIRM, NMW); Gorwydd, 26 July 1907, A. Ley (BIRM); Llangammarch Wells, 1 August 1963, E. S. Edees (NMW), Glannau Wells, Builth, 9 August 1980, M. P. (herb. M.P.); Cwm Dyfnant, 21 August 1982, M.P. (herb. M.P.). The known distribution is shown in Fig. 2. A record for Rads., v.c. 43, from Llandrindod Wells, fide Watson (1958), has not been confirmed. FiGurE 2. Distribution of Rubus aquarum Newton & Porter FIVE BRAMBLES FROM WALES 193 Rubus gallofuscus Newton & Porter, sp. nov. (Series Micantes) Turio altiarcuatus in apricis atropurpureus obtuse angulatus faciebus planis modice pilosus, glandulis mediocribus brevibusque, aciculis nonnullis, aculeis mediocribus tenuibus rectis, patenti- bus vel declinatis e basi compressa vinaceis apice croceo, aculeolis interdum glanduliferis obsitus. Folia quinata + digitata superne strigosa inferne viridia velutina; foliolum terminale + longe petiolatum late obovatum vel subrotundum vel pentagonum basi subcordata vel emarginata breviter acutum margine interdum undulata aequaliter serratum. Inflorescentia intricata late pyramidata ad apicem singulatim foliosa; ramuli medii cymosi in summa parte divisi c. 7-flori. Rachis vix flexuosa dense pilosa tomentosa aculeis tenuibus declinatis, aciculis nonnullis, glandulis brevibus mediocribus crebris armata. Flores 1-5—2 cm diam; sepala griseoviridia tomentosa breviter aciculata glandulifera patentia vel fructum laxe amplectantia. Petala obovata vel elliptica fimbriata alba haud contigua; stamina stylos aequantia vel vix superantia; filamenta alba; anthera glabra; carpella hirsuta; receptaculum + glabrum. Fructus parvus sphaericus. Stem arching, deep purple in exposure, round or bluntly angled with flat sides, with numerous short or medium simple or tufted whitish hairs, few or numerous sessile or very short stalked glands and stellate hairs, occasional to frequent short and medium stalked glands and a few acicles, numerous short-medium prickles grading into pricklets some gland-tipped, largest prickles c. 7-8 mm long, 10-15 per 5 cm, straight, slender, patent to declining from a compressed base, hairy, reddish-purple with straw coloured tips. Young shoots tinged with bronze or purple. Leaves mostly digitate or subdigitate; leaflets 5, usually not contiguous, dark green and shortly strigose above, mid green and thickly and softly hairy below with short simple hairs, some stellate hairs and sparse to frequent sessile glands; terminal leaflet medium to long stalked, broadly obovate round or pentagonal, about 6 X 5cm, emarginate or slightly cordate at the base, usually abruptly narrowed at the apex to a short point (c. 1 cm); margin flat or undulate, serrate or slightly biserrate with fairly even rather short mucronate teeth of variable width; basal leaflets with stalks 4-7 mm; petioles longer than basal leaflets with about 10-15 falcate prickles (c. 3 mm), otherwise clothed and coloured as the stem. Flowering branch with 1-3 ternate leaves below with axillary peduncles of about the same length or shorter, then leafy to the apex with a series of diminishing simple leaves; inflorescence intricate when well developed, broadly pyramidal or cylindrical with a domed apex; middle peduncles branched above halfway, with about 7 flowers. Rachis slightly flexuose, coloured as stem, with frequent slender declining prickles (up to 4 mm), more densely hairy and felted than stems with patent short-medium simple or tufted hairs, frequent short to long stalked glands and some acicles. Pedicels with almost patent acicular prickles (up to 3 mm) otherwise clothed like the rachis but with shorter hairs and more numerous medium or long stalked glands. Flowers about 1-5-2 cm in diameter; sepals greyish-green, felted and with short spreading simple hairs, few to frequent short acicles and stalked glands, mainly reflexed in open flower and patent or clasping in fruit, those of the tip fruits developing long points; petals c. 10 xX 6 mm, obovate to elliptical, white with a greenish claw, downy on the margin and both surfaces, not contiguous; stamens about the same length or slightly longer than green styles, filaments white, anthers glabrous; young carpels hairy; receptacle glabrous or slightly hairy. Fruit round, small. Flowering in July and August. Hotorype: field bank, Penrhos, Brecs., v.c. 42, GR 22/802.113, 2 August 1988, M. Porter (NMW). This bramble appears to have been collected first by A. Ley in 1890 from Pont-nedd-fechan in Breconshire and later from various localities in surrounding counties. Several gatherings were seen by Rogers and usually determined as ““R. fuscus Wh. and N. forma” and the bramble appears as R. fuscus var. macrostachys from v.c. 42 in Rogers (1900). It is the Mid-Wales representative of the “Rubus fuscus group” which includes eight named regional species from southern and central Britain as well as two local unnamed plants, in addition to R. fuscus Weihe and R. insectifolius Lef. & Mueller (R. fuscus var. nutans Rogers). Representative exsiccata: Herefs., v.c. 36, Wormbridge, 24 August 1906, A. Ley (NMW). Glam., v.c. 41, Ystalyfera, July 1906, H. J. Riddelsdell (NMW); Resolven, 22 July 1984, M.P. (herb. M.P.); near Gurnos, 3 August 1988, M. P. (herb. M.P.). Brecs., v.c. 42, Pont-nedd-fechan, 30 July 1890 A. 194 A. NEWTON AND M. PORTER 1 2 3 4 Ficure 3. Distribution of Rubus gallofuscus Newton & Porter Ley (BIRM), Ystradgynlais, 11 July 1906, A. Ley (BIRM); Heol Senni, 7 August 1906, A. Ley (BIRM, NMW, OXF); Heol Senni, 19 August 1947, E. S. Edees (NMW); Nant Senni, 21 July 1973, A.N. (herb. A.N.); Nant Llech, 2 September 1976, M.P. (herb. M.P.); Penrhos, 1 August 1977, M.P. (herb. M.P.); Cwmbrynich, 6 September 1977, M.P. (herb. M.P.); Cwm Giedd, 25 August 1978, M.P. (herb. M.P.). Carms., v.c. 44, Cwm Twrch, 7 July 1899, A. Ley (BIRM); Cwm Twrch, 2 August 1988, M.P. (herb. M.P.). The known distribution is shown in Fig. 3. Rubus biloensis Newton & Porter, sp. nov. (Series Anisacanthi) Turio arcuatus tandem procumbens obtuse angulatus faciebus striatis planis vel leviter concavis, in umbrosis pallide viridis in apricis cinnabarinus parce pruinosus, leviter pilosus, aciculis nonnullis, glandulis numerosis inaequalibus, aculeis croceis pilosis brevibus vel mediis rectis vel raro curvatis e basi compressa declinatis plerumque ad angulos dispositis minoribus interdum glanduliferis armatus. Folia 3—5-nata pedata superne parce strigosa subplicata inferne capillis simplicibus interdum stellatis vestita. Foliolum terminale apice cuspidatum ellipticum vel subrotundum basi truncata vel subcordata aequaliter serratum dentibus majoribus vix prominentibus. Inflorescentia angusta rigida ad apicem haud foliosa superne racemosa pedunculis patentibus 1—3(-—5)-flori. Rachis valida + recta dense pilosa aculeis nonnullis declinatis pusillis glandulisque crebris obsita. Flores 2— 2:5 cm diam. Sepala griseoviridia tomentosa glandulis aciculisque ornata patentia vel fructum amplectantia. Petala haud contigua fimbriata obovata pallide rosea; stamina stylos virides superantia; filamenta alba tandem rubescentia. Anthera glabra. Fructus mature rubiginosus. Stem low-arching then prostrate, bluntly angled with flat or slightly concave sides, pale green in shade becoming orange-red in exposure, striate, slightly pruinose, with sparse to numerous medium or short simple or tufted hairs, occasional to frequent stalked glands of various lengths, sparse to numerous sessile glands and an occasional short acicle; prickles short to medium (3-7 mm) straight or rarely curved, declining from a compressed base, chiefly on the angles but not confined to them c.(10—)15—20 per 5 cm, yellow or orange-red with yellow tips, hairy, the smaller sometimes gland- ' FIVE BRAMBLES FROM WALES 195 tipped; young shoots bronze in exposure. Leaves 3-5 nate, pedate, leaflets imbricate or not, ultimately mid-green above, paler beneath; thinly strigose above with sparse subsessile glands at first, hairy beneath with sparse to numerous simple hairs and sometimes stellate hairs; terminal leaflet short stalked, broadly obovate or elliptical, but rarely ovate, emarginate to narrowly cordate at the base and with a cuspidate often curved tip (c. 1-5 cm); fairly evenly serrate with main teeth slightly prominent; upper surface of leaf often becoming convex and sometimes plicate or rugose; basal leaflets with short stalks (to 5 mm); petiole usually rather longer than basal leaflets with about 10-15 small slanting or falcate prickles (c. 3 mm), clothed like the stem but with more numerous spreading simple hairs, stellate hairs and stalked glands. Flowering branch narrow, with 1-3 basal peduncles usually shorter than their ternate leaves and 1-2 simple leaves with short axillary peduncles; not leafy to apex; upper part of inflorescence simply racemose with middle and upper peduncles almost patent up to 2 cm long and bearing 1—3(—5) flowers. Rachis coloured as the stem, fairly stout and almost straight with few or many declining prickles (to 3 mm), densely clothed with spreading simple hairs and stellate hairs and frequent short to long stalked glands; pedicels with almost patent acicular prickles (to 3 mm), clothed like the upper part of the rachis but with more numerous medium to long stalked glands. Flowers 2—2-:5 cm diameter; sepals greyish-green felted with medium or short spreading simple hairs and sparse to frequent short to medium stalked glands and acicles, short pointed, patent, erect to clasping in fruit; petals not contiguous, usually obovate, c. 14 x 8 mm, very pale pink with a greenish claw, downy on the back and margin; stamens slightly longer than the styles, filaments white, turning red after petal fall, anthers glabrous; styles green or pink-based; young carpels glabrous; receptacle hairy or glabrous; fruit coloured reddish-brown at early stage, with medium to large number of drupelets. Flowering in July and August. Although the inflorescence is remarkably constant in form, there is considerable variation in the frequency of stalked glands, acicles and hairs on the stem. Ho.otype: lane bank, Llanfillo, Brecs., v.c. 42, GR 32/117.333, 21 July 1988, M. Porter (NMW). In 1975 an unfamiliar bramble was noticed in Llanfillo churchyard, Breconshire, and later was found growing in considerable quantity around the nearby Iron Age hill fort. Specimens sent to A. N. revealed that it was nameless. The following year two sheets of the same bramble, collected by A. Ley at Nant Tresglen, Breconshire, in 1906, were found in his herbarium at BIRM. These had been identified, probably by W. M. Rogers, as R. longithyrsiger Lees var. botryeros Focke. A duplicate gathering was later seen at NMW. During the last few years the same bramble has been found in several localities in Brecs., v.c. 42, and the adjacent vice-counties of Glam., v.c. 41, Rads., v.c. 43, Carms., v.c. 44 and Cards., v.c. 46. Recently it was discovered in N. Devon, v.c. 4, during a B.S.B.I. field meeting based at South Molton. Although superficially similar to R. leyanus Rogers in some of its vegetative characters, its simple, narrow, columnar inflorescence with stout rachis bearing pale pink flowers is distinctive. It has been named after the Celtic Saint Bilo to whom Llanfillo church is dedicated. She is reputed to have been one of the twenty four saintly daughters of King Brychan who ruled over the small Welsh kingdom of Brycheiniog, later known as Breconshire, in the 5th Century A.D. Representative exsiccata: Brecs., v.c. 42, Nant Tresglen, 8 August 1906, A. Ley (BIRM, NMW); Llanfillo, 26 July 1978, M.P. (NMW)); Allt Fillo, 26 July 1978, M.P. (herb. M.P.); Nant Gwennol, 28 July 1982, M.P. (herb. M.P.). Rads., v.c. 43, Penmaenau, 5 August 1988, M.P. (herb. M.P.); Cwmbach, 8 September 1988, M.P. (herb. M.P.). Carms., v.c. 44, Nant Gwennol, 28 July 1982, M.P. (herb. M.P.); Dolgran, 3 August 1987, M.P. (herb. M.P.); Dolaucothi, 3 August 1987, M. P. (herb. M.P.); near Ynys Wen, 25 August 1988, M.P. (herb. M.P.). Cards., v.c. 46, Lampeter, 20 July 1978, M.P. (herb. M.P.). The known distribution is shown in Fig. 4. Rubus merlini Newton & Porter, sp. nov. (Series Hystrices) Turio arcuatus tandem procumbens obtuse angulatus faciebus planis, in apricis brunneo-roseus leviter pilosus glandulis mediocribus numerosis, aciculis glanduliferis nonnullis aculeisque tenuibus 196 A. NEWTON AND M. PORTER 1 2 3 4 Ficure 4. Distribution of Rubus biloensis Newton & Porter rectis e basi lata declinatis amplitudine variabili omnino obtectus. Folia pedata quinata superne parce strigosa glabrescentia inferne leviter pilosa. Foliolum terminale brevipetiolatum ovatum basi cordata sensim acuminatum undulatum profunde duplicato-dentatum. Inflorescentia late pyrami- data superne aphylla; ramuli medi 1—3(—5) flori pedunculis + patentibus. Rachis recta capillis densis glandulis crebris amplitudine variis, aculeolis aciculisque patentibus numerosis obsita. Flores stellati c. 2-5 cm diam; sepala patentia vel fructum laxe amplectantia tomentosa glandulifera aciculata. Petala alba anguste obovata margine glabra; stamina stylos pallide virides vix superantia; filamenta alba, carpella glabra, receptaculum pilosum; fructus oblongus. Stem medium-low arching then trailing, bluntly angled with flat sides, brownish-pink in exposure, slightly striate, thinly to moderately hairy with simple or clustered spreading hairs, few or numerous sessile glands and numerous short to medium stalked glands and few to many gland-tipped acicles. Prickles numerous on angles and faces, straight, slender, declining from a compressed base, ranging in size from slightly less than stem diameter down to c. 2 mm, about 10-15 of the large prickles per 5 cm. Young shoots golden or copper coloured. Leaves pedate; leaflets usually 5, contiguous or not, slightly strigose or glabrescent above, softly but sparsely hairy beneath. Terminal leaflet short- stalked, ovate, with cordate base and gradually acuminate apex, undulate margins usually distinctly jagged and sharply biserrate. Petiole about the same length or shorter than the basal leaflets, clothed as the stem except that the largest prickles (2-3 mm) are often falcate. Flowering branch broadly pyramidal, usually with 1-3 long ascending or divergent axillary branches more than half the length of the inflorescence subtended by ternate leaves at the base, 1—3 simple leaves above, not leafy to the apex; middle panicle branches with 1—3(—5) flowers on fairly long almost patent pedicels. Rachis straight, densely clothed with spreading simple or tufted hairs, short to long stalked glands, numerous patent subulate pricklets and acicles. Pedicels clothed as upper rachis but with more crowded pricklets and acicles and the hairs more adpressed so that the stalked glands appear more conspicuous. Flowers starry, about 2-5 cm diameter; sepals patent in open flower, patent or loosely clasping in fruit, long-pointed, densely clothed with simple and stellate hairs, short or medium stalked glands and frequent acicles; petals white c. 14 X 6 mm, narrowly obovate, sparsely hairy on the back but with glabrous margins; stamens slightly exceeding styles, filaments white, anthers glabrous; styles pale green, carpels glabrous; receptacle pilose. Fruit oblong of about 30 drupelets. FIVE BRAMBLES FROM WALES 197 Ho.otyPe: roadside hedgebank, Cilycwm, Carms., v.c. 44, GR 22/751.396, 14 July 1973, T. A. W. Davis (NMW). Isotyre: herb. A.N. This plant was collected by A. Ley at the end of the nineteenth century from several localities in Mid Wales (vide infra). The early gatherings were usually identified as Rubus viridis Kaltenb., and under that name the plant is included in Rogers (1900). It differs however in significant respects from this continental plant (for which the correct name is R. iuvenis van de Beek) as exhibited by Wirtgen’s authentic specimens in MANCH. R. merlini forms a conspicuous member of the mid-Welsh bramble community, being widespread and frequent in v.cc. 42-44 & 46. It is easily distinguished by the very prickly stem, bright green attenuate terminal leaflets and white starry flowers. As it is particularly prevalent in the deciduous woods of Carms., v.c. 44, it has been named after the local sage Merlin who is associated with this area in many legends and after whom Carmarthen is named. Representative exsiccata: Brecs., v.c. 42, Fannog, Nant Towy, 4 August 1897, A. Ley (BIRM); Dyffryn Crawnon, 27 June 1898, A. Ley (BIRM); Cwm Gwydderig, 8 August 1906, A. Ley (BIRM); Corryn Wood, Garth, 29 July 1977, M.P. (herb. M.P.); Nant Gwennol, 2 August 1977, M.P. (herb. M.P.); Llangoed, 15 July 1980, M.P. (herb. M.P.); Abergwesyn, 23 August 1980, M.P. (herb. M.P.). Myarth, Llangynidr, 17 August 1988, M.P. (herb. M.P.). Rads., v.c. 43, Cwm Elan, 2 August 1986, A. Ley (BIRM); Allt goch, 26 August 1892, A. Ley (BIRM, MANCH, NMW). Carms., v.c. 44, Pont Gwydderig, 5 August 1897, A. Ley (BIRM, NMW); Rhandirmwyn, 29 July 1962, T. A. W. Davis (NMW), Rhyd-y-groes, 28 July 1978, T. A. W. Davis (NMW), Abergorlech, 22 August 1988, M. P. (herb. M.P.). Cards., v.c. 46, Nant Doethi, 10 August 1897, A. Ley (BIRM). The known distribution is shown in Fig. 5. FicureE 5. Distribution of Rubus merlini Newton & Porter 198 A. NEWTON AND M. PORTER REFERENCES EpeEs, E. S. & Newron, A. (1988). Brambles of the British Isles. London. E.uis, R. G. (1983). Flowering plants of Wales. Cardiff. Newton, A. (1972). A Welsh bramble foray. Watsonia 9: 117-130. Newton, A. (1974). Five brambles from South Wales. Nature in Wales 14: 24-33. Newton, A. (1976). Brambles in Glamorgan and neighbouring areas. Trans. Cardiff Nat. Soc. 97: 44-50. Pryce, R. D. (1988). B.S.B.I. Rubus Meeting, Carmarthen, 1987. B.S.B.I. Welsh Bulletin 46: 21-30. RIDDELSDELL, H. J. (1907). A flora of Glamorganshire. J. Bot., Lond. 45: Suppl. Rocers, W. M. (1899). Radnorshire and Breconshire Rubi. J. Bot., Lond. 37: 193. Rocers, W. M. (1900). Handbook of British Rubi. London. Rocers, W. M. & Ley, A. (1906). New brambles from South Wales. J. Bot., Lond. 44: 58. SALTER, J. H. (1935). The flowering plants and ferns of Cardiganshire. Cardiff. Watson, W. C. R. (1950). Yb. bot. Soc. Br. Isl. 1950: 47. Watson, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge. (Accepted August 1989) Watsonia, 18, 199-213 (1990) 199 Short Notes HIERACIUM BRITANNICUM F. J. HANB. IN WALES This taxon was described by Hanbury (1892). The description is clear, and conveys an accurate image of the plant that at the present time grows commonly on limestone rocks in Derbyshire. The distribution given by Hanbury (1892) mentions three vice-counties: Derbys. (v.c. 57), Staffs. (v.c. 39), and Mid-W. Yorks. (v.c. 64), and a possible fourth, Fife (v.c. 85). It is apparent from Hanbury’s comments that the Yorkshire plant was slightly different. Within a few years Hanbury (1895) was able to record the plant from at least ten vice-counties on the strength of specimens in his own or Mr Backhouse’s herbarium. Unfortunately the details have not been published but it is reasonable to assume that specimens from some localities, particularly those in Wales, would have been assigned at a later date either to H. subbritannicum (A. Ley) Sell & West, first described as a variety of H. stenolepis in 1909, or H. britanniciforme Pugsley, described in 1941. The latter was first recognised as distinct by field botanists more than 40 years earlier. The next expert to publish a summary of distribution was Linton (1905) who added Westmorland (v.c. 69) and Caerns. (v.c. 49) to the vice-counties given by Hanbury (1892). Hanbury had personally confirmed the identifications for these additional vice-counties, which were published in J. Bot., Lond. 32: 304 (1894), and Rep. botl Soc. Exch. Club Br. Isl. 2: 17 (1901), although from the description given the latter record refers to specimens of H. britanniciforme. This is confirmed by a specimen in herb. E. S. Marshall at CGE. Hyde & Wade (1934) listed H. britannicum as occurring in two Welsh vice-counties, Brecs. (v.c. 42) and Caerns. (v.c. 49). It is clear from the opinions of Ley (1909) and Pugsley (1948) that the plants from Brecs. are correctly referred to the taxon now known as H. subbritannicum. The Caerns. record was based on specimens at the National Museum of Wales; examination of the specimens from Caerns. determined as H. britannicum now at NMW shows that they are H. britanniciforme, with the exception of two sheets, which remain undetermined due to their condition. The next expert to publish a statement about H. britannicum in Wales was Pugsley (1948) who on p. 117 stated when speaking of H. britanniciforme, ‘“‘Both species grow on different parts of the Great Orme, H. britannicum as a very dwarf form.” Sell & West in Perring (1968) mapped the distribution of H. britannicum and showed many localities, but only in Staffs. and Derbys. Following this expert opinion, Ellis (1983) omitted any mention of this taxon as a Welsh plant, despite the record in Pugsley (1948). On a visit to Llandudno on 13th and 14th May 1988, the writer had the good fortune to discover both species. A walk on the evening of the 13th led to the discovery of a small colony of H. britannicum on the limestone outcrops below Mynydd Pant. The following day H. britanniciforme was seen in quantity on the Great Orme, although H. britannicum was not observed. It is interesting to note that Griffith (1895) recorded H. britannicum f. from Great Orme’s Head and rocks above Bodafon, thus recording both areas where the taxa still occur. H. britannicum has also been discovered recently on one of the limestone outcrops between Llandudno and Derbyshire, paralleling the distribution of H. holophyllum and its variety dentulum. The former occurs in v.cc. 49 and 57, and the latter in v.cc. 36, 51, 57 and 64. H. britannicum is at present only known to occur in v.cc. 39, 49, 50 and 57. The Cambridge University Herbarium (CGE) was examined in an attempt to discover any specimens supporting the early literature records of H. britannicum in the Llandudno area. The following specimens in the H. britanniciforme folders were considered by the writer to be H. britannicum:— Caerns., v.c. 49: cliffs on Great Orme’s Head, July 1871, S. H. Bickham 5; rock near Little Orme’s Head, July 1893, J. E. Griffith; Great Orme’s Head, August 1894, J. E. Griffith (a similar plant from the same gathering ex herb. Wheldon, now at NMW,, is H. britanniciforme); near Glodaeth, lime rocks, 19th June 1901, A. Ley; Little Orme, lime rocks, 19th June 1901, A. Ley. (This sheet 200 SHORT NOTES has four specimens, three being H. britannicum, the remaining specimen possibly being H. britanniciforme although dark papillae are present on the styles.); south west side of the Great Orme’s Head, exposed limestone rocks, 9th July 1912, E. S. Marshall. (This sheet has four specimens, and it may be that the two small flowering plants are H. britanniciforme as are the four small specimens collected by W. A. Shoolbred, on the same day, and now at NMW). Denbighs., v.c. 50: GR. 23/832.797, Bryn Ewyn, Colwyn Bay, numerous, growing in cracks on south-facing limestone, altitude 350 ft., 15th June 1968, J. M. Brummit. The characters used to diagnose H. britanniciforme in the field are the spotting on the broad, thick leaves which are purple on the underside, together with the relatively short peduncles and the large pale heads; the paleness of the heads is due to floccum (stellate hairs en masse) on the phyllaries. In H. britannicum, in contrast, spotting is absent, leaves are narrower and thinner with large basal teeth, which, when the plant is large, are very large. The heads are much darker in appearance because few stellate hairs are present and the peduncles are generally longer. In H. britanniciforme the styles are pure yellow, whereas in H. britannicum they are yellow but with dark papillae present, giving an overall impression of a dingy off-yellow. Examination of plants growing in situ is the easiest way to appreciate the differences between the taxa. In herbarium material which has been poorly prepared it is difficult to separate the species. The spotting may have almost disappeared and large heads can shrink during drying if sufficient pressure is not maintained while the specimen is in the press. However, the simple hairs on the leaves are unaffected by drying and, in H. britannicum, are much thinner than in H. britanniciforme, where the hairs are nearly setose. The phyllaries, particularly of specimens collected late in the season, can look very similar, but in H. britannicum they are longer, thinner and tend to twist at the apex and have fewer glands or hairs on the section behind the apex, which is also often tinged faintly pink. H. britannicum is illustrated in Butcher (1961), from a specimen collected on Little Orme. The accompanying description indicates the identity to be correct. To conclude, H. britannicum occurs at three sites on limestone in two vice-counties (49 and 50) in North Wales. It has persisted unnoticed by nearly all botanical visitors for 117 years, and should be looked for at suitable localities, particularly in Flints. (v.c. 51). REFERENCES BuTcHErR, R. W. (1961). A new illustrated British Flora. London. Exuis, R. G. (1983). Welsh flowering plants. Cardiff. GriFFiTH, J. E. (1895). The flora of Anglesey & Carnarvonshire. Gangor. Hansury, F. J. (1892). Further notes on Hieracia new to Britain. J. Bot., Lond. 30: 366. Hansury, F. J. (1895). The London catalogue of British plants, part 1, 9th ed. London. Hype, H. A. & Wane, A. E. (1934). Welsh flowering plants. Cardiff. Ley, A. (1909). Brecon and West Yorkshire Hawkweeds. J. Bot., Lond. 47: 12. Linton, W. R. (1905). An account of the British Hieracia. London. PERRING, F. H., ed. (1968). Critical supplement to the Atlas of the British flora. London. Puas.ey, H. W. (1948). A prodromus of the British Hieracia. J. Linn. Soc. Lond. (Bot.) 54: 113-117. J. BEVAN 23 Priory Street, Cambridge, CB4 3QH CARDAMINE AMARA L.: ITS OCCURRENCE IN MONTANE HABITATS IN BRITAIN C. amara is usually considered to be a lowland species in Britain and Ireland although known to occur up to 450 m in Scotland (Clapham 1987). This altitudinal limit refers to Buchanan White’s Perthshire record: ‘‘ascends to 1500 feet in lowland Forth” (White 1898). During the past five years for the Flora of Cumbria survey, the author has found this species very locally up to 540 m in the northern Pennines in Cumberland (v.c. 70) and up to 488 m in the eastern Lake District in Westmorland (v.c. 69). As it appeared likely that the Perthshire (v.c. 87) record was sited in the Ochil Hills, a search was made in 1989 and the altitudinal limit of C. amara extended to 610 m. In the montane habitats of the northern Pennines and Lake District, C. amara is present in SHORT NOTES 201 bryophyte-dominated springs and flushes, and along the rills and streams issuing from them. Epilobium alsinifolium is an almost constant associate from 244 m upwards. Chrysosplenium oppositifolium and Cochlearia officinalis are usually present and Cardamine pratensis, Montia fontana, Stellaria alsine and Veronica beccabunga commonly so. The abundant bryophytes include Bryum pseudotriquetrum, Cratoneuron commutatum, C. filicinum and Philonotis fontana. Saxifraga stellaris is an associate at the highest Lake District site and Carex rostrata at one of the higher Pennine sites. Although Saxifraga aizoides occurs in some of the flush systems, it keeps to the bare gravelly areas from which C. amara is absent. In the Ochils, C. amara is locally common in similar spring and flush communities and on mossy rocks by burn sides, being associated with Epilobium alsinifolium from 228 m upwards. Saxifraga hypnoides is a not uncommon associate and Epilobium anagallidifolium with the rare moss Splachnum vasculosum occur at the highest site. Elsewhere, C. amara, associated with Epilobium alsinifolium, has been noted at 228 m in the Howgill Fells in West Yorks. (v.c. 65) (G. Halliday pers. comm.). However in the Southern Uplands of Scotland, where C. amara ascends to 312 m in Selkirks. (v.c. 79) and 358 m in Roxburghs. (v.c. 80) in similar base-rich flushes, E. alsinifolium is absent. There are obvious similarities between these British habitats and those described by Ellenberg (1988) from Central Europe, where C. amara is present in subalpine and alpine spring swamp communities up to over 2000 m. They are described under the Montio-Cardamineta and Montio- Cardametalia and have Epilobium alsinifolium and several other British higher plants present. The associated bryophytes are also similar. Intensive sheep grazing occurs at all the British sites so that plants are dwarfed and rarely flower, although they have been seen to do so at 457 m on a flushed Pennine ledge, inaccessible to sheep. The winter-green basal leaves can be recognised throughout the year but care is needed to distinguish them from some variants of Cardamine pratensis, which ascends to the highest flushes and is the commonest Cardamine in this habitat in Britain. C. amara has a patchy distribution in Britain, being very rare in Wales and the Scottish Highlands, so that it is unlikely to occur there as a montane species. It is absent from the Moorhouse national nature reserve in the northern Pennines, and so is undescribed from the plant communities there (Eddy, Welch & Rawes 1969), although it occurs close to the western boundaries. This very local and sporadic occurrence, mostly as a stunted and sterile variant, explains why its presence as a montane British species has been overlooked for so long. ACKNOWLEDGMENT I should like to thank Dr G. Halliday for his help in the preparation of this note. REFERENCES CrLapHaM, A. R. (1987). Cruciferae, in CLAPHAM, A. R., TutTin, T. G. & Moore, D. M. Flora of the British Isles, 3rd ed., pp. 92-3. Cambridge. Eppy, A., WetcH, D. & Rawes, M. (1969). The vegetation of the Moorhouse national nature reserve in the northern Pennines, England. Veget. Acta Geobot. 16(5—6): 239-284. ELLENBERG H. (1988). Vegetation ecology of Central Europe, 3rd ed., pp. 430-31. Cambridge. Wuite, B. W. (1898). The flora of Perthshire, p. 62. Edinburgh. R. W. M. CorNER Hawthorn Hill, 36 Wordsworth St., Penrith, Cumbria, CAll 7OZ THE DISTRIBUTION OF CAREX APPROPINQUATA SCHUMACHER (C. PARADOXA WILLD.) IN GREAT BRITAIN AND IRELAND Carex appropinquata Schumacher is widespread in the wetlands of central and northern Europe, including Scandinavia, and extends into Asia as far as the Urals. Its distribution is, however, more scattered than that of its close ally C. paniculata L., for reasons not yet fully substantiated. 202 SHORT NOTES Ellenberg (1988) cites a survey by E. Balatova-Tula¢kova in Czechoslovakia as evidence that C. appropinquata requires more acid conditions than does C. paniculata, a view already taken by Jermy & Tutin (1968) and carried over, for lack of clear evidence to the contrary, into Jermy, Chater & David (1982). The conclusion is nevertheless surprising for, while C. paniculata is tolerant of a variety of soils, the greatest concentrations of C. appropinquata in the British Isles are in the basic fens of East Anglia and County Westmeath. Observations in the field suggest that C. appropinquata is less tolerant of drought, shade and competition than is C. paniculata. When, as has frequently happened in East Anglia, a site dries out as a result of improved drainage, increased water extraction, or invasion by scrub, C. appropinquata is the first of the two to disappear. The presence ef this sedge in the British Isles was first established in 1841 when R. Spruce recorded it (as C. paradoxa Willdenow) from Heslington Fields near York. A year later D. Moore found it in Ireland at Ladestown near Lough Ennell, County Westmeath; and between then and the end of the century it was also recognised in a number of East Anglian fens as well as on canal sides where the counties of Middlesex, Hertfordshire and Buckinghamshire meet. A new chapter opened in 1928 when W. A. Sledge showed that a colony of sedge at Malham Tarn was this species (Turrill 1929), and in the last 25 years Dr R. W. M. Corner has found it in several places in the Scottish border counties, while the explorations of the late Miss E. Booth, C. Breen and Miss M. Scannell have doubled the total of Irish records. Carex appropinquata has also been reported from Pembrokeshire, but erroneously (Evans 1989). This sedge may still have been overlooked elsewhere in the British Isles, for it is not always easy to distinguish it from C. paniculata. Depauperate variants of the latter may come very close to C. appropinquata, and a hybrid (almost wholly sterile) generally appears wherever the two grow together. In addition a tussocky form of C. diandra known as C. pseudoparadoxa S. Gibson, with a branched inflorescence, may confuse the issue (Sledge 1937), and it was this taxon that was responsible for the false reports from Wales. The character usually cited as the main distinction between C. appropinquata and C. paniculata is the nature of the basal sheaths, which in C. paniculata are entire, red- to dark-brown and shiny, while in C. appropinquata they are black, matt, and eventually split into separate hair-like fibres; but the splitting hardly occurs until the sheath is dry and the plants are often growing in standing water. The leaves of C. appropinquata are a bright yellow-green and less than 2 mm broad, but those of a poor specimen of C. paniculata may be no wider than this. The glumes of C. appropinquata have a reddish tinge but this only becomes pronounced when the plant is dried. The safest guide is the shape of the utricle, more or less triangular with a broad serrated wing in C. paniculata, ovoid in C. appropinquata and abruptly narrowed into an unwinged beak. The utricle of C. diandra is narrowly winged, and the beak is split with the edges of the split overlapping. All recorded stations in the British Isles have been visited since 1980, by myself or by botanists who may be considered wholly reliable. The state of each colony so observed is indicated in the following list by the letters A = up to 20 plants, B = 21 to 100, C = 101 to 1000, D = over 1000. Where the sedge has not been refound, the date and authority for the most recent sightings are. given. The authenticity of specimens cited is confirmed by me. [W. Sussex, v.c. 13: 51/2.1, Henfield, 1939, J. E. Lousley (RNG). In Borrer’s garden and clearly one of the ‘foreign’ plants introduced there by him. ] Herts., v.c. 20: 51/0.9, Harefield, meadow near Copper Mills, 1885, J. Benbow (BM); [52/1.2, Hitchin, Oughton Head, 1921, J. E. Little (CGE), a puzzling sedge but certainly not C. appropinquata. | Middlesex, v.c. 21: 51/0.7, West Drayton, canal, 1873, J. L. Warren (BM, CGE); 51/0.8, Uxbridge Moor, 1910, C. B. Green (CGE); 51/0.9, Harefield, formerly abundant in several places from Springfield Lock towards Rickmansworth but finally destroyed by gravel-digging c. 1936 (Kent 1975). Bucks., v.c. 24: 51/0.8, near Denham, 1905, G. C. Druce (BM, CGE), destroyed by railway works c. 1925. W. Suffolk, v.c. 26: 52/7.7, Icklingham Poors Fen (B); Cavenham, ‘fen valley wood’ (= Ash Plantation), 1952, S. M. Walters (CGE); between Mildenhall and Eriswell, 1938, E. Nelmes (K), probably same as ‘Bombay Fen’, since destroyed; 52/7.8, Eriswell, Caudle Fen (Trist 1979), ploughed in the 1970s; Brandon, Fenhouse Heath, 1956, F. Rose, and Palmer’s Heath, 1975, A. SHORT NOTES 203 O. Chater & Mrs G. Crompton, now gone; 52/9.7, Market Weston Fen, last seen 1960 (Simpson 1982). E. Norfolk, v.c. 27: 62/4.9, between Haddiscoe and Somerleyton, 1955, F. Rose, exact site not traceable; 63/0.0, Hingham, Seamere (A), declining; 63/3.0, Aldercarr Fen (C), Wheatfen (C), and Parish Marsh (C); Strumpshaw (B); 63/3.1, Hoveton Great Broad (A); Hoveton Little Broad, 1975, F. Rose; Ranworth, west (C), east (D); Upton Broad, south (D), north (C); Woodbastwick Fen, three areas (A, B, C); Horning Fen, 1950, F. Rose; 63/3.2, Dilham Broad Fen (C); Barton Great Fen, 1902, C. E. Salmon (BM), may = Catfield Great Fen (C); Stalham, Wood Marsh, 1902, C. E. Salmon (BM); Longmoor Point, Middle Fen (C) and Sutton Broad Fen (D); 63/4.0, Acle Decoy Carr, 1955, F. Rose; 63/4.1, Thurne, Shallam Dyke, 1956,/. F. M. & M. J. Cannon (BM), now drained; 63/4.2, Hickling, 1888, A. H. Evans (BM); between Palling Wood and Horsey, 1956, T. G. Tutin, exact site not traceable. W. Norfolk, v.c. 28: 52/9.8, Middle Harling (A); Overa Heath, 1963 (Petch & Swann 1969), area since reclaimed; 52/9.9, Hockham, Cranberry Rough (B); Thompson Common (A); Stow Bedon Fen, 1926, T. J. Foggitt (BM); 53/6.1, Wormegay, Mow Fen (B); Shouldham Warren, east end, 1919, J. E. Little (CGE), extinct by 1943 (Petch & Swann 1969); 53/8.0, Great Cressingham, 1966, E. L. Swann, access now denied; 53/9.0, Scoulton Mere, before 1926, H. D. Hewitt (Petch & Swann 1969); 53/9.2, Guist Fen (C); 53/9.3, Sculthorpe Moor, 1985, C. P. Petch, one plant, formerly more plentiful; 62/0.9, Swangey Fen (A); Old Buckenham Fen, 1956, F. Rose. Cambs., v.c. 29: 52/5.7, Wicken Fen (B), formerly “abundant”, 1885, A. Fryer (BM). [Pembs., v.c. 45: 12/7.2, the records for Pwll Trefeiddan, Dowrog Common and Caerfarchell are all referable to C. diandra.| [Derbys., v.c. 57: 43/2.4, near Shirley Mill, error for C. paniculata (Linton 1903). | E. Yorks., v.c. 61: 44/6.4, Heslington Fields, 1846, W. W. Newbould (BM); Langwith Common, 1852, S. Thompson (BM), both sites drained by end of century; 54/0.4, Beverley, Pulfin Bog, 1968, R. W. David (A), now almost certainly lost under carpeting Phragmites and Glyceria; [54/ 0.5, of two specimens from Driffield, collected 1898 and 1903 by C. Waterfall and in BM determined by A. Bennett as C. appropinquata, the first is C. diandra and the second C. disticha; while a recent report from the same area is not supported by a specimen;] 54/1.4, Leven Canal, 1953, Miss M. E. Crackles (A), now gone; 54/1.7 or 2.7, Flamborough Head, 1912 (Robinson 1914), never confirmed. Mid-W. Yorks., v.c. 64: 34/8.6, Malham Tarn Fen, east (B), west (D); 44/5.4, between Heelaugh and Askham Richard (Lees 1888); Askham Bog (B). [N. W. Yorks or Co. Durham, v.c. 65 or 66: 35/9.2, unlocalised, 1947, N. Brownbridge, 10-km card at Biological Record Centre, not accepted by Perring & Walters (1962). ] Peebless., v.c. 78: [36/1.4, Medwen, a specimen collected 1979 by D. J. McCosh and at first thought to be C. appropinquata has now been determined as C. paniculata;| 36/2.3, Innerleithen, The Glen, 1858, Lyell (Balfour 1925), never confirmed. Selkirks., v.c. 79: 36/3.1, Clearburn Loch, one plant; Alemoor Back Loch, two places (A,B); Alemoor Loch, north (B). Roxburghs., v.c. 80: 36/3.1, Alemoor Loch, south (B); 36/4.1, Branxholme Wester Loch (B); 36/ 4.2, Dunhog Moss (B); 36/5.1, Adderstonlee Moss (B). [Mid Ebudes, v.c. 103: Mull, the record in Clapham, Tutin & Warburg (1952) is an error (Jermy & Crabbe 1978). | [N. Ebudes, v.c. 104: 17/1.5, Gunna, 1940 (Heslop-Harrison 1941) is almost certainly an error.] Co. Clare, v.c. H.9: 11/3.7, Dromoland (C). Co. Carlow, v.c. H.13: 21/8.8, Rahill Bog, 1973, Miss E. M. Booth, site destroyed before 1978. Offaly, v.c. H.18: 22/0.1, Lough Coura (B). Westmeath, v.c. H.23: 22/0.4, Killinure Lough, west (A); 22/1.4, Twy Bog (B); Lough Makeegan (C); 22/2.4, south-east of Lough Sawdy (C); 22/2.5, Lough Sawdy, west side (A); near Glencarry House (A); 22/2.6, Ballynacarrow, reported 1958 but specimen in TCD is C. diandra; 22/3.4, Lough Ennell, western shore (C); 22/3.5, Monroe (B); Mount Dalton Lake (C); south of Kenny (C); Royal Canal, Kilpatrick Bridge (A); 22/3.6, Lough Iron (C); Lough Owel, Bunbrosna (D); 22/4.4, Lough Ennell, eastern shore, two plants; Lough Ennell, Ladestown (C); 22/4.5, Lough Ennell, Bog of Linn (C); Tullaghan Bog (D); Lough Owel, south-eastern corner (C); Royal Canal, east of Mullingar (C); Scraw Bog (B); 22/4.6, Ballinafid Lough (C). 204 SHORT NOTES REFERENCES BaLFour, F. R. S. (1925). Botany, in Bucuan, J. W. A history of Peeblesshire 1: 338. Glasgow. CLAPHAM, A. R., Tutin, T. G. & WARBURG, E. F. (1952). Flora of the British Isles. Cambridge. ELLENBERG, H. (1988). Vegetation ecology of Central Europe, 4th edition, translated by G. K. Strutt. Cambridge. Evans, S. B. (1989). Carex appropinquata in Pembs., v.c. 45 -— a misidentification for C. diandra. B.S.B.I. News 53: 17-18. Hes.op-Harrison, J. W. (1941). The flora of the isles of Coll, Tiree and Gunna, Proc. Univ. Durham phil. Soc. 10: 302. Jermy, A. C., CHATER, A. O. & Davin, R. W. (1982). Sedges of the British Isles. London. Jermy, A. C. & Crass, J. A. (1978). The Island of Mull, a survey of its flora, p. 11.68. London. Jermy, A. C. & TutTin, T. G. (1968). British sedges. London. KENT, D. H. (1975). The historical flora of Middlesex. London. Lees, F. A. (1888). The flora of West Yorkshire. London. Linton, W. R. (1903). Flora of Derbyshire, p. 296. London. PERRING, F. H. & WALTERS, S. M., eds. (1962). Atlas of the British flora. London. Petcu, C. P. & Swann, E. L. (1969). Flora of Norfolk. Norwich. RoBInson, J. F. (1914). Botanical report. The Naturalist 1914: 32-33. Simpson, F. W. (1982). Simpson’s Flora of Suffolk. Ipswich. SLEDGE, W. A. (1937). Carex pseudoparadoxa S. Gibson. Rep. botl Soc. Exch. Club Br. Isl. 11: 358-361. TristT, P. J. O. (1979). An ecological flora of Breckland. Wakefield. TurrIiLL, W. B. (1929). Report of the distributor for 1928. Rep. botl Soc. Exch. Club Br. Isl. 8: 930-931. R. W. Davip 50 Highsett, Cambridge, CB2 INZ THE CUMBRIAN HERBARIUM OF W. H. YOUDALE Liverpool Museum (LIV) recently acquired W. H. Youdale’s herbarium of Cumbrian flowering plants and ferns. Youdale was a draper who lived in Cockermouth, Cumbria (Collins & Pound 1988). Little was hitherto known of Youdale’s botanical activities and, although he was a contemporary of W. Hodgson of Workington, the author of Flora of Cumberland, there is no reference to Youdale in this Flora, nor to his published records from Silloth (Youdale 1892). However, his herbarium specimens provide ample evidence that he undertook a thorough survey of the Cockermouth and Allonby areas of Cumberland (v.c. 70), with occasional forays further afield to St Bees, Sellafield, Honister Pass and Coniston. Youdale’s herbarium is well preserved and is mounted on standard-sized sheets. A short note appeared in B.S.B.I. News (Allen 1987) which gave the size of the collection as no more than 175 sheets; the correct number, excluding undetermined scraps and a small collection made by Miss McGlasson from Hyéres, S. France in December 1894, is 314 sheets. The collections were amassed over a period of 18 years. Youdale’s own gatherings cover the years 1891 to 1908 (excluding 1901 and 1905). His collaborators were as follows: T. Brown, 1892, 1894; J. P. Dalton, September 1891; James Dobbins, 1892, 1893; Rev. Hilderic Friend, 1894-96; Mr Postgate, July 1902; J. Scott, August 1907; W. West jnr, May, June & August 1891; F. Yeomans, June 1901; George Yeomans, 1891-93, 1895, 1901, 1904, 1908; L. M. Youdale, August & September 1893; Mrs W. H. Youdale, July 1892. It is possible that the last two collectors listed are the same person. Of the eleven or twelve collectors represented in the herbarium, only two, Rev. Hilderic Friend and William West jnr, are listed in Kent & Allen (1984). William West’s specimens were collected from Caernarvonshire, Hertfordshire and Surrey. The Rev. Hilderic Friend was a Methodist minister who was allotted the Cockermouth circuit for three years around 1895 (Friend 1943), and he botanised widely in Britain. His collections occur in a number of public herbaria (Kent & Allen 1984). With these exceptions, the botanists whose collections are represented in Youdale’s herbarium confined their activities to western Cumberland, an area for which few collections exist from this period. The relevant tetrads, and those visited by Youdale himself, are shown in Fig. 1. From a total of 314 named and localised specimens, 246 were collected by Youdale alone; a small SHORT NOTES 205 Vg | |S ERA ar fap eR 1 3 5 = Ficure 1. Distribution of localities of Cumbrian specimens represented i in the W. H. Youdale herbarium, shown as tetrads. number was collected jointly by Youdale and Friend. The majority of specimens are of widespread native species. There are, however, a small number of aliens present, some of which were collected from around the docks in Silloth (GR 35/12.52). An example is Hesperis matronalis L. Other records from Silloth Docks include two species now extinct in v.c. 70: Agrostemma githago L. and Arabis glabra (L.) Bernh. The latter species was also recorded by Friend (1895, 1896) as Turritis glabra L. from near Cockermouth, where he claimed it was native. There are a number of records of non-native species from ‘“‘wood behind South Lodge, Cockermouth, T. Brown” which are the first records for v.c. 70. These are all species which could be expected to have persisted in, or escaped from, gardens: Centaurea montana L., Euphorbia lathyrus L., Laburnum anagyroides Medic., Ruta graveolens L. and Salvia argentea L., the latter two not having been subsequently recorded in the county. Turgenia latifolia (L.) Hoffm. from “‘Stackyard, Dubb Mill, Allonby, August 1894, W. H. Youdale”’ is also the first record for v.c. 70, the only other record being from Carlisle in 1905 where it was found by T. S. Johnstone. Another first record is of Prunus lusitanica L., collected by W. H. Youdale from a plantation at Holmwood, Cockermouth in June 1894. Youdale’s record of Centaurium littorale (D. Turner) Gilmour is, surprisingly, from an inland 206 SHORT NOTES site, “near the summit of the Hay, Cockermouth, September 16, 1891, alt. 700 ft’’. Atriplex glabriuscula Edmondston from “‘sea shore near Dubb Mill, Allonby, August 1894, W. H. Youdale”’ is the first v.c. 70 voucher specimen for this scarce seaside plant. Salvia verticillata L. was recorded for v.c. 70 by Youdale from Skinburness in July 1896 and from Silloth Docks in August 1907. It still occurs at Silloth Docks, its only extant locality in the vice-county. Among the notable records published in Youdale’s only known botanical article (Youdale 1892) was Crithmum maritimum L. from the coast south of Silloth. This is the only Cumberland record from north of St Bees Head, but there is no voucher specimen in his herbarium. & REFERENCES ALLEN, D. E. (1987). A local herbarium for sale. B.S.B.I. News 47: 32. Couns, J. & Pounp, E. (1988). Natural history, science & medicine: Catalogue no. 1090, item no. 6433, Maggs Bros. Ltd. London. FRIEND, H. (1895). Turritis glabra, etc., in the Lake District. The Naturalist, August 1895: 238. FRIEND, H. (1896). Tower Cress near Cockermouth. The Naturalist, August 1896: 256. FRIEND, J. W. (1943). Hilderic Friend (1852-1940). [Obituary]. N.W. Naturalist 18: 117-119. Hopecson, W. (1899). Flora of Cumberland. Carlisle. KENT, D. H. & ALLEN, D. E. (1984). British and Irish herbaria. London. YouDALE, W. H. (1892). Silloth in August. Sci. Gossip 28: 14-15. H. HoFMANN, J. R. EDMONDSON & G. HALLIDAY Liverpool Museum, William Brown St., Liverpool, L3 8EN ALOPECURUS BULBOSUS GOUAN IN DORSET Alopecurus bulbosus Gouan is relatively easy to see for two or three weeks from mid-May when it is flowering, but very hard to find later in the season. FitzGerald (1989) showed that it still survives at many sites in south-eastern England where it had not been recorded for many years. Inspired by knowledge of her fieldwork, I searched for the plant in Dorset (v.c. 9) in the summer of 1989. The results were similarly encouraging, showing that, while the grass has certainly been lost at some sites because of urban expansion, agricultural improvement and reduced salinity, much of the apparent decline is because the plant is not actually looked for. Another reason for the search was to look for the hybrid Alopecurus X plettkei Mattfeld (A. bulbosus X geniculatus), reported from as far west as Christchurch Harbour (Trist & Wilkinson 1989). In both of the Dorset sites where the hybrid was found it was growing both a little further from the sea and a little higher above HWMOT in a less open community. The results of the searches are given below. In addition, all the likely salt-marsh areas of the Fleet were investigated, but with no success other than at the new site at Rodden. Though it may seem rash to say so, there are few other likely sites in the county, although it is odd that there are no other records from the southern shores of Poole Harbour. The following list details the Dorset sites searched for the occurrence of Alopecurus bulbosus in 1989 in grid reference order from the west. (The names in brackets refer to field records collected and held at the Dorset Environmental Record Office in Dorchester, with the exception of those of Mansel-Pleydell referred to below.) Seatown, GR 30/42.91 (H. J. M. Bowen 1950). Not refound. Site now a car park. West Bay, GR 30/463.905 (R. Good 1936). Many thousands of plants over meadow apparently now used only for a summer fair. By far the best site in the county (herb. D.P.). Burton Bradstock, GR 30/476.897 (R. Payne 1981). In small quantity amongst caravans behind shingle bar (herb. D.P.). Burton Mere, GR 30/509.878 (Graveson 1956). Many hundreds of plants over 0-5 km but all specimens examined were the hybrid A. X plettkei (herb. D.P.). West Bexington, GR 30/528.867 & 523.871 (R. McGibbon 1985 — this is Mansel-Pleydell’s “Swyre”’ site). In small quantity to the east of the extensive reedbed, and in greater quantity to the west (herb. D.P.). SHORT NOTES 207 Abbotsbury, GR 30/572.842 & 568.842 (R. Walls 1986). In abundance in open field to north-west of Swannery. The next field to the west produced only the hybrid — this field was much more overgrown with richer vegetation (both herb. D.P.). Rodden Hive, GR 30/604.823. A new site. Abundant both on edge of Fleet and at the seaward end of a newly ploughed field with no other vegetation (herb. D.P.). Radipole, GR 30/67.79 (H. J. M. Bowen 1963). Not refound. The salinity is dropping as a result of sluices, although Carex divisa still present. All of the other old sites in Weymouth are long since built over. Lodmoor, GR 30/688.814 (H. J. M. Bowen 1955). Refound in one small area, but as access to this R.S.P.B. reserve is very difficult there may well be more. Piddle Marshes, GR 30/93.88 (D. Ranwell 1965). Not visited, access is very difficult. Redcliffe, GR 30/939.873 (R. Good 1936, etc.). Only a few plants were found in one area. The rest of the grazing marshes have been steadily improved. Swineham, GR 30/943.879 (D. Ranwell 1965). A few plants. Keysworth, GR 30/947.897 (R. McGibbon 1985). A very few plants in one small area. It is odd that the vast areas of grassland on the north-western shore of Poole Harbour produces only this one patch. The Moors, Arne, GR 30/94.89 (D. Ranwell 1965). Ranwell described this in 1965 (Card Index at I.T.E., Furzebrook) as ‘‘Optimum development over several areas, almost pure sward locally”’. As mentioned above the grazing marshes have been improved and although there are still patches of Juncus maritimus with a poor grass flora, there was no sign of any A. bulbosus. Poole, GR 40/01.90 (Mansel-Pleydell 1895). Marsh near Railway Station. Presumably long since built over. Swanage, GR 40/02.79 (Mansel-Pleydell 1895). Presumably long since built over. There is little to be added to FitzGerald’s (1989) excellent notes on associations. Suffice it to say that Carex divisa was found in most of the western sites, but Ranunculus baudotii only at the hybrid site at Burton Mere. The other associates were much as described by her. The Rodden site (colonizing a ploughed field) was most interesting: the site abutted closed Phragmites marsh on the other side of the stream to the main colony, and it seemed as though saline turf had been ploughed. However, the site must have been too saline for the new ley to germinate, as opposed to the rest of the field inland, and the result was that there was no vegetation other than A. bulbosus, which is scattered over an area of 10 m”. Its fate is awaited with interest. ACKNOWLEDGMENTS I would like to thank Robin Walls and Anne Horsfall for assistance; P. J. O. Trist for identifying all the specimens that are now in herb. D.P.; C. D. Preston for comments and advice and above all Rosemary FitzGerald for inspiration and encouragement. REFERENCES FitzGERALD, R. (1989). ‘Lost and Found’ — Alopecurus bulbosus Gouan in S.E. England. Watsonia 17: 425-428. MANSEL-PLEYDELL, J. C. (1895). Flora of Dorset. Dorchester. Trist, P. J. O. & WiLkinson, M. J. (1989). Alopecurus X plettkei Mattfeld in Britain. Watsonia 17: 301-307. D. PEARMAN The Old Rectory, Frome St. Quintin, Dorchester, Dorset, DT2 OHF THE ORIGIN AND TAXONOMY OF SPARTINA X NEYRAUTII FOUCAUD The salt marsh grass Spartina X townsendii H. & J. Groves is well known as the hybrid progenitor of Spartina anglica Hubbard, the ‘classic’ example of a recently evolved allopolyploid species. Marchant (1967, 1968) concluded that the available historical, morphological, and cytological 208 SHORT NOTES evidence on the status of S. x townsendii is entirely consistent with it being a hybrid between S. maritima (Curtis) Fernald, a native of England, and S. alterniflora Loiseleur, a species introduced into Southampton Water from the United States in the early 1800s. The hybridisation appears to have occurred at Hythe, Hampshire, presumably just before the first discovery of S. x townsendii in 1870. The nature of the origin has recently been confirmed beyond all reasonable doubt by the use of isozyme markers (Raybould 1989; Raybould et al., in press.). What appears to be a similar event has been recorded from the coasts of south-western France and northern Spain. S. maritima is a native of the area, and S. alterniflora was introduced by shipping (presumably independently of the English introduction); the first record is from 1806 near the mouth of the River Bidassoa (Hubbard, Grimes & Marchant 1978). In 1892 Neyraut collected plants at Hendaye (Mobberley 1956), which he believed to be identical to S. X townsendii (Chevalier 1923). However, Foucaud (1984) described the variants found by Neyraut as a new species, S. neyrautii Foucaud. Since the discovery of S. neyrautii there has been doubt over its true status (see Marchant (1977) for a full discussion of the arguments), the most generally accepted conclusion being that the taxon is a hybrid between S. maritima and S. alternifiora, but that it can be distinguished from British S. x townsendii on morphological grounds. One possible explanation for this difference is that S. x townsendii and S. X neyrautii are reciprocal hybrids (Arber 1934). To investigate the matter further, a search was made for S. X neyrautii in 1970 (Hubbard, Grimes & Marchant 1978). Apparently the only remaining site for S. X neyrautii was in a small area of salt marsh adjacent to San Sebastian airport. Marchant (1977) compared the morphology of these plants with S. x townsendii from England and concluded that S. x neyrautii was “not referable to S. X townsendii’”. However, for each of the vegetative characters considered, the range given for each taxon overlaps in every instance. Marchant also presented cytological data from the two taxa. They both have the same AP. ESea. GDE se OF EL. NED NEE: PGI SkDH SS FiGurE 1. Common phenotype of Spartina X townsendii and S. X neyrautii for eleven isozyme systems. Abbreviations: AP — acid phosphatase, EST — esterase, GDH — glutamate dehydrogenase, GOT — glutamate oxalacetate transaminase, IDH — isocitrate dehydrogenase, MDH - malate dehydrogenase, ME - malic enzyme, PER — peroxidase, PGI — phosphoglucose isomerase, SkDH — shikimic acid dehydrogenase, SOD — superoxide dismutase. SHORT NOTES 209 chromosome number of 2n=62 and very similar chromosome pairing behaviour, although S. x neyrautii has a slightly higher frequency of multivalents. These data strongly support the idea that, like S. X townsendii, S. X neyrautii is a hybrid between S. maritima and S. alterniflora. To confirm this hypothesis we obtained isozyme phenotypes from a clone of S. x neyrautii which had been collected in Spain by Hubbard et al. and has since been kept in cultivation first at the Coastal Ecology Research Station in Norwich and subsequently at the John Innes Institute. We have previously shown (Raybould et al., in press) that twelve clones of S. x townsendii from the Hythe population when examined for eleven isozyme systems showed no variation and phenotypes which are entirely consistent with S. X townsendii being a hybrid between S. maritima and S. alterniflora. The isozyme phenotypes of S. < neyrautii for these eleven systems were obtained using the same methodology and were found to be completely identical with those of S. X townsendii. These systems resolved into over 70 bands and may well represent the products of at least 50 loci (Fig. 1). This extensive evidence confirms the nature of S. < neyrautii as a hybrid between S. maritima and S. alterniflora and also shows that the actual parents of the two hybrids were identical with respect to these isozyme systems. This result raises three points. First, it at least suggests that S. maritima may be highly uniform genetically throughout its whole range. It has previously been shown that this species has very low levels of genetic variability in Britain (Raybould 1989). Second, the two introductions of S. alterniflora into Europe appear also to be very similar genetically. Third, S. x townsendii and S. x neyrautii are synonyms because they have the same parentage; the former has priority. ACKNOWLEDGMENTS We should like to thank Dr A. Smith of the John Innes Institute for her gift of the Spartina x neyrautii clone used in this study. A.F.R. gratefully acknowledges the financial support of an N.E.R.C. (C.A.S.E.) studentship. REFERENCES ArBER, A. (1934). The Gramineae. Cambridge. CHEVALIER, A. (1923). Note sur les Spartina de la flore frangais. Bull. Soc. bot. Fr. 70: 54-63. Foucaup, M. (1894). Un Spartina nouveau. Ann. Soc. Nat. Sci. Rochelle 31: 8. HusBarD, J. C. E., Grimes, B. H. & Marcuanrt, C. J. (1978). Some observations on the ecology and taxonomy of Spartina X neyrautii and Spartina alterniflora growing in France and Spain and comparison with Spartina X townsendii and Spartina anglica. Doc. Phytosociol., New Ser. 2: 273-282. MarcuanT, C. J. (1967). Evolution in Spartina (Gramineae). I. History and morphology of the genus in Britain. Bot. J. Linn. Soc. 60: 1-24. MarcuantT, C. J. (1968). Evolution in Spartina (Gramineae). II. Chromosomes, basic relationships and the problems of the S. X townsendii agg. Bot. J. Linn. Soc. 60: 381-409. Marcuant, C. J. (1977). Hybrid characteristics in Spartina X neyrautii Fouc., a taxon rediscovered in northern Spain. Bot. J. Linn. Soc. 74: 289-296. MosseErLEY, D. (1956). Taxonomy and distribution of the genus Spartina. Iowa St. Coll. J. Sci. 30: 471-574. RAYBOULD, A. F. (1989). The population genetics of Spartina anglica C. E. Hubbard. Ph.D. thesis, University of Birmingham. RAYBOULD, A. F., GRAY, A. J., LAWRENCE, M. J. & MARSHALL, D. F. The evolution of Spartina anglica C. E. Hubbard (Gramineae): Origin and genetic variation. Biol. J. Linn. Soc. (in press). A. F. RAyBouLp, A. J. Gray, M. J. LAWRENCE & D. F. MARSHALL School of Biological Sciences, University of Birmingham, Birmingham, B15 2TT RUMEX ACETOSA L. SUBSP. BIFORMIS (LANGE) VALDES-BERMEJO & CASTROVIEJO IN BRITAIN Many years ago I described and illustrated a coastal cliff ecotype of Rumex acetosa L., based on the following specimen in the Cambridge University herbarium (CGE): Cornwall, Zennor, 20 July 210 SHORT NOTES 1839, C. C. Babington (Rechinger 1961). I wrote: “‘a very similar if not identical plant observed by myself in July 1959 in similar habitats along the north coast of Spain, Cabo Penas near Santander . it looks very distinctive, especially when alive, but I still hesitate to give it taxonomic recognition as my field experience is limited to one locality in northern Spain and as I have seen only one good herbarium specimen from Britain.” Since then, the Spanish plant in question has been identified by Valdés-Bermejo & Castroviejo (1977) as R. acetosa subsp. biformis (Lange) Valdés-Bermejo & Castroviejo. There are beautiful full-page illustrations of this taxon by Lange (1861, 1867). When recently visiting north-western Spain I was fortunate to observe and collect subsp. biformis at the following localities in the province of La Coruna: Cabo Finisterre, littoral siliceous rocks, 7.7.1988, K. H. Rechinger 64750 (W); Cabo Toriniana, littoral siliceous rocks, 17.7.1988, K. H. Rechinger 64784 (W); Cabo Vilano, 5km NW Carmarinas, granitic rocks, 18.7.1988, K. H. Rechinger 64842 (W). As can be seen from the type locality quoted below, my Spanish specimens come from the classical area. Here follows a description of my plants from the ditio classica. Rumex acetosa L. subsp. biformis (Lange) Valdés-Bermejo & Castroviejo in Anales Inst. Bot. A. J. Cavanilles 34: 326 (1977). R. biformis Lange in Index Sem. Hort. Haun. 26 (1857). Tyre: In rupibus maritimis prope oppidum La Coruna Galiciae, 10. vii. 1852, Lange. Stems (10—)15—30 cm high, usually several ascending from an indurate branching rootstock; base sometimes horizontally elongated. Basal leaves thick, fleshy and shiny when alive, thick leathery when dried, (20—)30-40(-50) x (8-)15—20(-30) mm, ovate-hastate, with very short, acute, sometimes obtuse and reduced basal lobe; petiole as long as or up to 1-5 times as long as the blade. Stem leaves few (1—)2+4, oblong-hastate, up to 4 times as long as broad, with very short acutish or obtuse basal lobes, short petiolate to sessile. Inflorescence short and dense, with few undivided erect or slightly divergent branches, only the lower ones sometimes remote. Valves 4-5 x 3-5-5 mm. In all the Spanish localities of the litoral subspecies, the populations appeared to be completely isolated from any inland populations of R. acetosa. No transitional variants have been observed. It might be worthwhile for British botanists to revisit the old Cornish locality and to look for new ones along the south-western coasts of Britain. I have no doubts about the identity of the Cornish plant with the northern Spanish R. acetosa subsp. biformis, which evidently is an Atlantic geographical race. Its distribution is comparable to that of R. rupestris Le Gall, a relative of R. conglomeratus Murr. (section Rumex). REFERENCES LANGE, J. M. C. (1861). Vidensk. Meddel. Dansk Naturk. Foren. Kjébenhavn 1861: 48-49. LANGE, J. M. C. (1867). Descriptio iconibus illustrata plantarum novarum vel minus cognitarum praecipue e Flora Hispania, t.24. Copenhagen. RECHINGER, K. H. (1961). Notes on Rumex acetosa L. in the British Isles. Watsonia 5: 64-66 & pl. 4,5. VALDE£s-BERMEJO, E. & CastrovigEso, S. (1977). Notas cariosistematicas sobre Flora Espanola: 2. Anales Inst. Bot. A. J. Cavanilles 34: 325-334. K. H. RECHINGER A-1130 Wien, Beckgasse 22 A NEW HYBRID BINOMIAL IN MIMULUS L. As was shown by Roberts (1964), the hybrid between M. guttatus DC. and M. luteus L. s.1. is widespread in Britain and frequently confused with its parents. It occurs in extensive, clonal SHORT NOTES 211 populations independently of its parents and clearly merits its own binomial epithet. Although the present author is currently preparing a full account of Mimulus in Britain, there is a need for early publication of a name for this hybrid. The view is taken here that the ‘luteus’ parent is M. luteus var. rivularis Lindley, though there are good reasons for considering this taxon as not being conspecific with the typical M. luteus, which does not occur in Britain. In view of the fundamental contribution made by R. H. Roberts to the understanding of naturalised Mimulus taxa, and since he published an account of this particular hybrid, it is appropriate that he be commemorated by the choice of epithet. Mimulus X robertsii Silverside, hybr. nov. Ho.orypus: Hethpool, College Burn, Northumberland, v.c. 68, GR 36/895.280, in hill stream, 10th August 1989, Silverside 1989/159 (E). Hybrida ex M. guttatus DC. et M. luteus var. rivularis Lindley orta. Planta sterilis. Corollae flavae, palatis rubro-guttatis, lobis frequenter rubro-maculatis. Calyces basibus sparse hispidulis. Pedicelli sparse hispiduli, infimi calycibus 1—3plo longiores. Folia ovata vel ovato-rhombea apicibus acutis vel subacutis, dentata, saepe dentibus acutis, patentibus vel subantrorsis, et obscuris alternantibus. Bracteae late ovatae, dentatae. Stolones crassi breves, foliis mediocribus. A sterile hybrid, variably intermediate between the parents. Corollas yellow, palates red-spotted, lobes commonly unmarked, as in M. guttatus, or else blotched with red on the labellum or on all five lobes. Pedicels and calyx-bases minutely and usually sparsely hispid. Lower pedicels up to three times as long as their calyces at maturity. Leaves ovate or ovate-rhomboid with acute or subacute apices, dentate, often with acute, patent to subantrorse teeth alternating with shallow, indistinct teeth. Bracts broadly ovate, dentate. Stolons short and thick, with the leaves not noticably different in size from other cauline leaves. M. guttatus normally differs from superficially similar clones of the hybrid in its more rounded, crenately toothed leaves, its small, suborbicular, entire-margined upper bracts, its more elongated inflorescences, the horizontally held labella of newly-opened flowers and in its conspicuously inflated fruiting calyces. This last character may, however, be seen in some clones of the hybrid, though no seed is matured. The long, slender, small-leaved stolons produced in some abundance by M. guttatus in the late summer and autumn have not yet been observed in any clone of its hybrids. M. luteus var. rivularis, a very local plant of hill streams in north-eastern England and southern and central Scotland, is a smaller, neater, mat-forming plant, with usually paler, more darkly blotched corollas, few-flowered inflorescences, pedicels often more than five times the length of their fruiting calyces and with leaf-teeth often more than twice as long as wide and somewhat twisted. The pedicels and calyces are normally entirely glabrous but in hot weather and in greenhouse cultivation they become minutely glandular-puberulent. The type of material of M. x robertsii is from a clone with blotches on all five corolla-lobes and with rather blunt leaf-teeth somewhat approaching those of M. guttatus. M. X robertsii is widespread in northern and western Britain and is often the only taxon present along upland river systems. It is only very locally established in the south-east and is apparently rare, or under-recorded, in Ireland. Mimulus ‘A. T. Johnson’ is a horticultural clone of the hybrid, frequently sold for bog and rock gardens, collected by Mr Johnson from a boggy stream in the Welsh hills before 1935 (Thomas 1980). The above description must be taken to exclude certain plants with large, very broad leaves and with very heavily blotched, broader corollas. These plants, locally established in a few areas of Scotland, are hybrids involving M. variegatus J. St-Hil. (Silverside 1981). Although M. xX robertsii is described above as sterile, Parker (1975) reported a Mimulus population from the Lleyn peninsula in N. Wales with pollen fertility exceeding 70% and which on cytological grounds he regarded as a variant of this hybrid. CHOICE OF TYPE MATERIAL The hybrid of M. cupreus Dombrain and M. luteus var. rivularis (i.e. M. x maculosus T. Moore) is at least partially fertile and can be further crossed with M. guttatus to produce a sterile triple hybrid. 212 SHORT NOTES Roberts (1968) found that some artificially produced clones of this hybrid were indistinguishable from what is here called M. X robertsii. The present author has carried out similar crosses and has also produced plants that would be taken to be M. xX robertsii, though most such triple hybrids are instantly recognisable. It follows that type material for M. xX robertsii has had to be carefully selected. The College Valley, Northumberland, is a site where M. luteus var. rivularis occurs, along with at least three different clones of M. x robertsii. The occurrence of multiple clones of M. x robertsii is a phenomenon that is also linked with the presence of M. luteus var. rivularis in other localities, and it appears highly probable that they represent spontaneous local hybridisation. Living material of M. luteus var. rivularis collected in the College Valley by J. E. Halfhide and the author in 1975 was subsequently used for much experimental work and at least two generations of plants were obtained through self-fertilisation. No segregation of cupreus-characters was observed. Artificial hybridisations using this material and M. guttatus produced a range of products including one closely similar to the College Valley clone that has been selected as the source of the type material. REFERENCES PARKER, P. F. (1975). Mimulus in Great Britain — a cytotaxonomic note. New Phytol. 74: 155-160. Roserts, R. H. (1964). Mimulus hybrids in Britain. Watsonia 6: 70-75. Roserts, R. H. (1968). The hybrids of Mimulus cupreus. Watsonia 6: 371-376. SILVERSIDE, A. J. (1981). Mimulus L., in Wiccinton, M. J. & GRAHAM, G. G., eds. Guide to the identification of some of the more difficult vascular plant species. Nature Conservancy Council England Field Unit Occasional Paper No. 1: 81-84. Tuomas, G. S. (1980). A. T. Johnson. The Garden 105: 484-488. A. J. SILVERSIDE Department of Biology, Paisley College of Technology, Paisley, Renfrewshire, PAI 2BE NEW NAMES AND COMBINATIONS IN THE BRITISH FLORA The six following names are required for forthcoming floristic publications: 1. Ulmus minor Miller subsp. sarniensis (C. Schneider) Stace, comb. et stat. nov. U. glabra f. sarniensis C. Schneider, Ill. Handb. Laubholzk. 1: 220 (1904). This new combination was made invalidly by Stace (1989), the basionym given there being a nomen nudum. 2. Mentha x villosa Hudson var. nicholsoniana (Strail) R. Harley, comb. nov. M. nicholsoniana Strail in Rep. botl Soc. Exch. Club Br. Isl. 1: 186 (1888). M. x villosa Hudson is the correct name for the hybrid combination M. spicata L. x M. suaveolens Ehrh., of which var. nicholsoniana is a distinctive pubescent variant with lanceolate- oblong leaves with a more or less acuminate apex and teeth that are not or scarcely folded under. 3. X Tripleurothemis Stace, nothogenus nov. Hybridae inter Tripleurospermum Schultz Bip. et Anthemis L. This intergeneric combination is the only British one known to me that does not already have a nothogeneric name. 4. x Tripleurothemis maleolens (P. Fourn.) Stace, comb. nov. x Anthemimatricaria maleolens P. Fourn., Fl. Compl. Plaine Frang. 274 (1928). This appears to be the correct binomial for Tripleurospermum inodorum (L.) Schultz-Bip. x Anthemis cotula L., the only combination so far recorded in Britain in the nothogenus x Tripleurothemis. SHORT NOTES 213 5. Veronica reptans Kent, nom. nov. pro Veronica repens Clarion ex DC. in Lam. & DC., Fl. Fran¢., 3rd ed., 3: 727 (1805); non V. repens Gilib., Fl. Lit. Inch. 1: 108 (1781), nom. illegit. 6. Schoenoplectus < kuekenthalianus (Junge) Kent, comb. nov. Scirpus X scheuchzeri Bruegger in Jahresber. Naturf. Ges. Graubiindens 23/24: 119 (1878-1880), non Vitman, Summa pl. 1: 150 (1790). S. X kuekenthalianus Junge in Jahrb. Hamburg Wiss. Anst. 22, Beih. 3: 73 (1905). Schoenoplectus X scheuchzeri Palla ex Janchen, Cat. fi. austr. 1(4): 752 (1960). This new combination is required for the hybrid Schoenoplectus tabernaemontani X S. triqueter. ACKNOWLEDGMENTS I am grateful to D. H. Kent for pointing out that my earlier U/mus combination was invalid and for other help, and to R. M. Harley and D. H. Kent for publishing their contributions in this note. REFERENCE Stace, C. A. (1989). New combinations in the British and Irish Flora. Watsonia 17: 442-444. C. A. STACE Department of Botany, University of Leicester, Leicester, LEI 7RH Watsonia, 18, 214-228 (1990) Plant Records Records for publication must be submitted to the appropriate Vice-county Recorder (see Vice-county Recorders (1988)), and not the Editors. The records must normally be of species, hybrids or subspecies of native or naturalized alien plants belonging to one or more of the following categories: 1st or 2nd v.c. record; 1st post-1930 v.c. record; only extant v.c. locality, or 2nd such locality; a record of an extension of range by more than 100 km. Such records will also be accepted for the major islands in v.cc. 102-104 and 110. Only 1st records can be accepted for Rubus, Hieracium and hybrids. Records for subdivisions of vice-counties will not be treated separately; they must therefore be records for the vice-county as a whole. Records of Taraxacum are now being dealt with separately; by Dr A. J. Richards, and will be published at a later date. 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)). All records are field records unless otherwise stated. With the exception of collectors’ initials, herbarium abbreviations are those used in British and Irish herbaria by D. H. Kent & D. E. Allen (1984). Records from the following vice-counties are included in the text below: 2, 4-8, 11, 12, 14, 16, 21, 25, 26, 29, 33, 35, 37-39, 41, 43-47, 50, 52-54, 64, 67, 68, 72, 73, 75-77, 79-81, 83, 93, 94, 96-99, 103, 108, H35, H36, H40. The following signs are used: * before the record: to indicate a new vice-county record. + before the species number: to indicate that the plant is not a native species of the British Isles. + before 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. 1/4. LYCOPODIUM CLAVATUM L. *52, Anglesey: Parys Mountain, GR 23/44.90. D. F. Evans, 1989, NMW, det. G. Hutchinson. 1/5. DIPHASIASTRUM COMPLANATUM Subsp. ISSLERI (Rouy) Jermy 96, Easterness: Strath Nethy, GR 38/0.0. Amongst Calluna. D. J. Tennant, 1981, E, det. A. C. Jermy. 2nd record. 4/1. EQUISETUM HYEMALE L. 39, Staffs.: North-east of Upper Arley, GR 32/77.81. Stream banks in deciduous woodland. R. Maskew, 1989, herb. B.R. Fowler, det. J. M. Camus. 1st record since 1863. 4/6 X 10. EQUISETUM PALUSTRE L. X E. TELMATEIA Ehrh. *52, Anglesey: Traeth Lligwy, GR 23/49.87. Stabilised dune sand. R. H. Roberts, 1989. *72, Kirkcudbrights.: Tarras Water, GR 35/39.83. Wooded, marshy slope. M. E. R. Martin & O. M. Stewart, 1989, E, det. C. N. Page. 19/1. CYSTOPTERIS FRAGILIS (L.) Bernh. 7, N. Wilts.: South Wraxall, GR 32/82.63. Stonework of bridge. J. Presland, 1987. 1st record since 1941. 21/2. DRYOPTERIS AFFINIS (Lowe) Fraser-Jenkins 53, S. Lincs.: Nocton Wood, GR 53/08.63. Primary woodland. N. Hards, 1984. 2nd record. 21/2 aff. DRYOPTERIS AFFINIS (Lowe) Fraser-Jenkins subsp. AFFINIS 75, Ayrs.: Troon, GR 26/ 3.3. T. Wise, n.d. but c.1900, GL, det. C. R. Fraser-Jenkins. 2nd record. 21/2 bor. DrYOPTERIS AFFINIS subsp. BORRERI (Newman) Fraser-Jenkins 75, Ayrs.: Kelburn Glen, GR 26/22.56. A. R. Church, 1989. 2nd record. 21/2 cam. DRYOPTERIS AFFINIS subsp. CAMBRENSIS Fraser-Jenkins *41, Glam.: Bwich y Clawdd, GR 21/9.9. C. R. Fraser-Jenkins, 1988. 75, Ayrs.: Kelburn Glen, GR 26/22.56. A. R. Church, 1989. 2nd record. 21/6. DRYOPTERIS CARTHUSIANA (Vill.) H. P. Fuchs 93, N. Aberdeen: Horntowie, GR 38/ 48.46. Moss. D. Welch, 1989, ABD. 2nd record. (24/5. GYMNOCARPIUM ROBERTIANUM (Hoffm.) Newman 96, Easterness: Delete record pub- lished in Watsonia 17: 464 (1989), specimen is G. dryopteris (L.) Newman, det. A. C. Jermy.] 214 PLANT RECORDS 215 26/1. PILULARIA GLOBULIFERA L. *53, S. Lincs.: Swanholme, GR 43/94.68. Old sand and gravel pit. I. Weston, 1985. Whisby, GR 43/92.67. Recently abandoned sand pits. N. F. Stewart, 1987. 1st and 2nd records. 75, Ayrs.: Loch Maberry, GR 25/28.75. Loch margin. H. A. Lang, 1984. Only extant locality. +27/1. AZOLLA FILICULOIDES Lam. 26, W. Suffolk: Whatfield, GR 62/02.46. Village pond. E. M. Hyde, 1988. 2nd record. *H36, Tyrone: Lough Neagh south of Farsnagh Point, GR 23/ 96.39. P. Corbett and R. S. Weyl, 1988. 29/1b. OPHIOGLOSSUM AZORICUM C. Presl *11, S. Hants.: Warwick Slade, GR 41/26.06. A. J. & B. Rackham, 1984, BM. Beaulieu Old Airfield, GR 41/35.00. R. P. Bowman, 1987, herb. R.P.B. 1st and 2nd records, both det. A. C. Jermy & A. M. Paul. 46/5. RANUNCULUS ARVENSIS L. 7, N. Wilts.: Near Corston, GR 31/91.82. Weed in nursery. J. Hall, 1985. Only extant locality. 46/20. RANUNCULUS CIRCINATUS Sibth. 81, Berwicks.: Kettleshiel Burn, GR 36/69.51. Gravelly burn. M. E. Braithwaite, 1989, herb. M.E.B. 2nd extant locality. 46/22a. RANUNCULUS AQUATILIS L. 94, Banffs.: Portsoy, GR 38/59.64. Ditch. J. Edelsten, 1987, E, det. S. D. Webster. 1st post-1930 record. 46/22b. RANUNCULUS PELTATUS Schrank 7, N. Wilts.: Oaksey, GR 31/98.92. Pond. M. A. R. & C. Kitchen, 1987, det. D. Green. Only extant locality. +51/off. PAEONIA OFFICINALIS L. *77, Lanarks.: Shiels, GR 26/53.66. Grassy bank. P. Macpherson, 1985, herb. P.M. Still present in 1989. +53/agg. BERBERIS AGGREGATA C. Schneider *12, N. Hants.: Stonerwood Park, Steep, GR 41/ 73.25. Hanger above Ridge Farm. J. D. Fryer, 1985, herb. J.D.F., det. J. M. Mullin. +54/1. MAHONIA AQUIFOLIUM (Pursh) Nutt. 94, Banffs.: Forglen, GR 38/69.52. Neglected garden. T. C. G. Rich, 1987. 2nd record. 57/1. CERATOPHYLLUM DEMERSUM L. 83, Midlothian: Dunsapie Loch, GR 36/28.73. Eutrophic lake. D. R. McKean, 1988, E. Only extant locality. *H36, Tyrone: Ballagh Lough, GR 23/ 50.50. Lake. R. S. Weyl, 1982. 65/3. CORYDALIS CLAVICULATA (L.) DC. *77, Lanarks.: Deanside, Glasgow, GR 26/52.65. Heath remnant. P. Macpherson, 1989, herb. P.M. 66/6b. FUMARIA MURALIS subsp. BORAEI (Jordan) Pugsley 38, Warks.: Sherbourne, GR 42/ 25.62. Waste ground. A. Newton & J. C. Bowra, 1987, WAR, det. M. G. Daker. 1st record since 1872. +67/3. BRASSICA RAPA L. 94, Banffs.: 6 km N. of Aberchirder, GR 38/62.58. Roadside verge. J. Edelsten, 1987, E, det. T. C. G. Rich. 2nd record. +67/jun. BRASSICA JUNCEA (L.) Czern. *38, Warks.: Leamington Spa, GR 42/33.65. Allot- ment. J. W. Partridge, 1986, WAR, det. A. L. Grenfell. Digbeth, Birmingham, GR 42/08.85. Roadside. J. W. Partridge, 1988, WAR, det. T. C. G. Rich. 1st and 2nd records. +67/tou. BRASSICA TOURNEFORTIT Gouan *38, Warks.: Coventry, GR 42/32.80. Allotment. Leamington Spa, GR 42/32.63. Allotment. Both J. W. Partridge, 1989, herb. T.C.G. Rich, det. T.C.G.R. 1st and 2nd records. [+70/2. SINAPIS ALBA L. 93, N. Aberdeen: Delete record published in Watsonia 16: 144 (1987), specimen at ABD is Raphanus raphanistrum L.| +71/1. HIRSCHFELDIA INCANA (L.) Lagréze-Fossat 38, Warks.: Nechell’s Green, Birmingham, GR 42/08.87. Waste ground. J. W. Partridge, 1988, WAR, det. J. C. Bowra. 2nd record. +74/3. RAPHANUS SATIVUS L. 50, Denbs.: Near Tregeiro, GR 33/17.33. Shingley river bank. J. A. Green, 1989, NMW. 2nd record. 216 PLANT RECORDS 75/1. CRAMBE MARITIMA L. 45, Pembs.: Broomhill Burrows, GR 12/88.00. Shingle. P. Rhind, 1989. Single plant. Only extant locality. 79/6. LEPIDIUM LATIFOLIUM L. 738, Warks.: Birmingham Inland Port, GR 42/09.87. Railway sidings. J. W. Partridge, 1989, WAR, det. T. C. G. Rich. 2nd record. 83/1. IBERIS AMARA L. 12, N. Hants.: Porton Ranges, GR 41/24.37. Bare chalk of rabbit scrapings. D. Graiff, 1988. Only extant locality. 97/4 X 1. CARDAMINE FLEXUOSA With. X C. PRATENSIS L. *38, Warks.: Balsall Street, GR 42/ 22.76. Swampy meadow. J. W. Partridge, 1989, herb. T. C. G. Rich, det. T.C.G.R. 98/2. BARBAREA‘STRICTA Andrz. *25, E. Suffolk: Oulton Broad, GR 62/50.92. Mud dredged from river. J. M. Tusting, 1989, herb. T.C.G. Rich, det. T.C.G.R. 102/5 x 4. RorIpPA PALUSTRIS (L.) Besser X R. AMPHIBIA (L.) Besser *38, Warks.: Stratford upon Avon, GR 42/20.54. Mud in reed bed. J. W. Partridge, 1989, herb. T.C.G. Rich, det. el oh ce |e 102/76 X 3. RORIPPA AUSTRIACA (Crantz)Besser X R. SYLVESTRIS (L.)Besser *98, Main Argyll: Loch a’Mhuilin, GR 17/85.29. Waste ground. B. H. Thompson, 1987 and 1989, herb. B.H.T., det. T. C. G. Rich. 1st Scottish and 2nd Bnitish record. *RHUS TYPHINA L. 41, Glam.: Lower Swansea Valley, GR 21/6.9. Cycle trackway. G. Hutchinson, 1989, NMW. 2nd record. 115/6. HYPERICUM MACULATUM subsp. OBTUSIUSCULUM (Tourlet) Hayek 25, E. Suffolk: Beccles, GR 62/43.92. By disused railway line. P. G. Lawson & F. W. Simpson, 1987. Only extant locality. 115/6 X 5. HyPERICUM MACULATUM Crantz X H. PERFORATUM L. *38, Warks.: Leek Wootton, GR 43/30.69. Rough grass. J. W. Partridge, 1989, WAR, det. N. K. B. Robson. "G7; S. Northumb.: Near Briarwood, GR 35/79.62. Roadside. G. A. & M. Swan, 1989, herb. G.A.S., det. N. K. B. Robson. *68, Cheviot: North bank of R. Coquet near Rothbury, GR 46/05.01. River bank. G. A. Swan, 1989, herb. G.A.S., det. N. K. B. Robson. 133/1b. STELLARIA NEMORUM subsp. GLOCHIDISPERMA Murb. 7*45, Pembs.: Withy Bush, Haverfordwest, GR 12/95.16. Weed in flower bed. S. B. Evans, 1989. Probably imported with forestry bark mulch. 7141/6. ARENARIA BALEARICA L. *73, Kirkcudbrights.: Below Shambellie House, New Abbey, GR 25/96.66. Wall. O. M. Stewart, 1989. +153/1. AMARANTHUS RETROFLEXUS L. 4.N. Devon: East Putford, GR 21/38.19. Arable field. W.H. Tucker, 1989. 2nd record. +156/7. ATRIPLEX HALIMUS L. *5_S. Somerset: Blue Anchor, GR 31/00.44. Behind sea wall. Lady R. FitzGerald, 1989. 156/lon. ATRIPLEX LONGIPES Drejer *35, Mons.: R. Wye below Chepstow, GR 31/53.93. River bank. M. A. R. & C. Kitchen, 1988, det. J. R. Akeroyd. 156/pra. ATRIPLEX PRAECOX Hilphers *108, W. Sutherland: Badnabay, GR 29/22.46. Muddy track to salt marsh. E. Norman, 1988, BM. 160/nit. SALICORNIA NITENS P. W. Ball & Tutin *68, Cheviot: Alnmouth, GR 46/24.09. Upper salt-marsh. G. A. Swan, 1989. Holy Island, GR 46/10.43. Upper salt-marsh. G. A. Swan & M. Swan, 1989. Ist and 2nd records, both det. I. K. Ferguson. 162/2. TILIA CORDATA Miller *8,S. Wilts.: Whiteparish Common, GR 41/24.22. Bank. D. W. Soden, 1986, det. P. M. Woodruffe. +168/2. GERANIUM MACRORRHIZUM L. *39, Staffs.: Ettinghall, GR 32/92.95. Waste mine site. C. B. Westall, 1988. a en _ PLANT RECORDS 217 +170/6. OXALIS ARTICULATA Savigny *75, Ayrs.: Ardrossan Castle Rock, GR 26/23.43. Long grass. A. Rutherford & A. McG. Stirling, 1989. +170/exi. OXALIS ExILIS A. Cunn. *38, Warks.: Leamington Spa, GR 42/32.65. J. W. Partridge, 1988, WAR, det. M. F. Watson. Stretton-on-Fosse, GR 42/22.38. Footpath. J. C. Bowra, 1989, WAR, det. P. J. Copson. Ist and 2nd records. 81, Berwicks.: Longformacus, GR 36/ 69.57. Pavement. M. E. Braithwaite, 1989, herb. M.E.B. 2nd record. +171/3. IMPATIENS PARVIFLORA DC. *35, Mons.: Twyn Lane, Glascoed, GR 32/33.01. Lane banks. R. Fraser, 1989. 185/1. GENISTA TINCTORIA L. 45, Pembs.: Moory Farm north of Redberth, GR 22/07.05. Molinia — dominated heathy pasture. H. Harries & R. Elliot, 1989, det. S. B. Evans. 2nd record. 185/2. GENISTA ANGLICA L. 80, Roxburghs.: Brotherstone Hill, GR 36/61.36. Calluna heath. R. W. M. Corner, 1989, herb. R.W.M.C. 1st record since 1890. 190/1 x +2. MEDICAGO FALCATA L. X M. SATIVA L. *77, Lanarks.: Govan, GR 26/59.63. Waste ground by road. P. Macpherson, 1988, herb. P.M., det. P. J. O. Trist. 190/4. MEDICAGO MINIMA (L.) Bartal. +*12, N. Hants.: Blackmoor Fruit Farm, GR 41/76.32. Arable weed. Lady A. Brewis, 1988. Introduced with shoddy but established for at least 14 years. 192/19. TRIFOLIUM FRAGIFERUM L. *39, Staffs.: Stafford Common, GR 33/92.25. Short turf in pasture. S. Lawley, 1989, herb. B. R. Fowler. +193/1 pol. ANTHYLLIS VULNERARIA subsp. POLYPHYLLA (DC.) Nyman *29, Cambs.: Near Burwell, GR 52/57.65. Bank of old railway line. P. D. Sell, 1966, CGE, det. J. R. Akeroyd. +206/7. VICIA VILLOSA subsp. VARIA (Host) Corb. 6, N. Somerset: Odd Down, Bath, GR 31/ 74.62. Waste ground. D. E. Green, 1988, herb. E. J. Clement, det. E.J.C. 1st record since 1917. 206/12. VICIA LUTEA L. +7, N. Wilts.: Wanborough Plain, GR 41/22.80. Roadside verge. J. Newton, 1988, herb. D. Green. Only extant locality. 207/6. LATHYRUS SYLVESTRIS L. 43, Rads.: Burfa Bank, GR 32/28.60. Scrub on wooded hillside. D. R. Humphreys, 1989. Only extant locality. +208/1. PHyYSOCARPUS OPULIFOLIUS (L.) Maxim. *H36, Tyrone: Glenlark, GR 23/57.87. Riverside. J. Harron, 1987. Well established. 1st Irish record. +209/alb. X 2. SPIRAEA ALBA Duroi X S. DouGLASI Hooker *73, Kirkcudbrights.: South of Corse, GR 25/67.76. O. M. Stewart, 1980. *77, Lanarks.: Cathkin, GR 26/62.58. Roadside. P. Macpherson, 1989, herb. P.M., det. A. J. Silverside. 210/1. FILIPENDULA VULGARIS Moench *75, Ayrs.: Knockdaw Hill, Colmonell, GR 25/16.88. Amongst heather on serpentine. L. McTeague, 1987, E, det. A. McG. Stirling. 1st native record from W. Scotland. *211/S. RUBUS PARVIFLORUS Nutt. *98, Main Argyll: R. Add, Kilmichael Glassary, GR 16/ 86.94. Island in river. M. G. B. Hughes & B. H. Thompson, 1989, det. D. R. McKean. 4211/7. RUBUS PHOENICOLASIUS Maxim. *5,S. Somerset: Leighland Chapel, Old Cleeve, GR 31/03.36. Lane side. C. J. Giddens, 1988. 211/11/14. RUBUS CONJUNGENS (Bab.) Rogers *4, N. Devon: Braunton Burrows, GR 21/ 45.34. Dunes. B.S.B.I. Meeting, 1989, det. A. Newton. 211/11/27. RuBUS TUBERCULATUS Bab. *4, N. Devon: Croyde Bay, GR 21/43.39. Pathside. B.S.B.I. Meeting, 1989, det. A. Newton. 211/11/32. RUBUS BRITANNICUS Rogers *4, N. Devon: West Molland, GR 21/79.28. Forestry track. B.S.B.I. Meeting, 1989, det. A. Newton. 211/11/116. RuBus AccLiviraTuM W. C. R. Watson *4,N. Devon: Heasley Mill, GR 21/73.32. Rough field. B.S.B.I. Meeting, 1989, det. A. Newton. 218 PLANT RECORDS 211/11/123. RUBUS CARDIOPHYLLUS P. J. Mueller & Lefévre *75, Ayrs.: Finnarts Bay, Loch Ryan, GR 25/05.72. A. McG. Stirling, 1972, E, det. A. Newton. 211/11/133. RUBUS ROSSENSIS Newton *4, N. Devon: Hares Down, GR 21/84.20. Moorland edge. B.S.B.I. Meeting, 1989, det. A. Newton. 211/11/183. RUBUS DREJERI G. Jensen *75, Ayrs.: Grassyards, Mauchline, GR 26/51.27. Planted shelter belt. A. Newton & A. McG. Stirling, 1987, E, det. A. Newton. 211/11/249. RUBUS MORGANWGENSIS W. C. Barton & Riddelsd. *4, N. Devon: Odam Moor, GR 21/74.18. Forestry plantation. B.S.B.I. Meeting, 1989, det. A. Newton. 211/11/380. RUBUS AEQUALIDENS Newton *4, N. Devon: Huntsham Wood, GR 31/00.18. Forestry entrance. L. J. Margetts, 1985, det. A. Newton. 211/11/bar. RuUBUS BARTONII Newton *4, N. Devon: Hobby Drive, Clovelly, GR 21/33.23. Woodland. A. Newton, 1989. 211/lli/lam. RUBUS LAMBURNENSIS Rilstone *4, N. Devon: Hockworthy, GR 31/02.19. Roadside. L. J. Margetts, 1988, det. A. Newton. 4212/8. POTENTILLA NORVEGICA L. 41, Glam.: Mountain Ash, GR 31/0.9. Reclaimed land. J. P. Curtis et al., 1987, NMW, det. R. G. Ellis. 2nd extant locality. 212/13. POTENTILLA ERECTA subsp. STRICTISSIMA (Zimm.) A. Richards *H40, Co. London- derry: Grange Park Wood, GR 24/74.29. Grass heath over peat. D. S. Lambert, 1989, det. A. J. Richards. 212/14. POTENTILLA ANGLICA Laicharding 7, N. Wilts.: Prickmoor Wood, GR 31/95.65. Pasture by wood. D. Green, 1987, det. B. Harold. Only extant locality. *79, Selkirks.: Wood Street, Galashiels, GR 36/47.37. J. A. Murray, 1982, herb. R.W.M. Corner, det. B. Harold. 220/3/2. ALCHEMILLA FILICAULIS subsp. VESTITA (Buser) M. E. Bradshaw *46, Cards.: Cwm Berwyn, GR 22/72.78. Sheep-grazed slope with boulders. A. O. Chater, 1988, NMW, det. S. M. Walters. 225/11. ROSA TOMENTOSA Sm. *39, Staffs.: Kinver Edge, GR 32/83.82. Border of conifer plantation. C. B. Westall, 1989. Rudyard Lake, GR 33/94.61. Disused railway. D. J. Tinston, 1989, herb. B.R. Fowler. lst and 2nd records, both det. A. L. Primavesi. 225/13 X 11. RosA MOLLIS Sm. X R. TOMENTOSA Sm. *39, Staffs.: Bateman’s Dingle, Upper Arley, GR 32/77.80. Scrub on woodland edge. R. Maskew, 1989, det. A. L. Primavesi. +225/mul. ROSA MULTIFLORA Thunb. *39, Staffs.: Planks Lane, Wombourne, GR 32/86.92. Road verge. C. B. Westall, 1988. Known here for 20 years. 226/+2 X 1. PRUNUS DOMESTICA L. X P. SPINOSA L. *5,S. Somerset: Muchelney Ham, GR 31/ 43.23. Hedge. P..&1..Green, 1978; det. A. Cideshie: +227/2. COTONEASTER SIMONSII Baker 77, Lanarks.: Shiels, GR 26/52.66. Waste ground. P. Macpherson, 1985. 2nd record. +227/3. COTONEASTER HORIZONTALIS Decne 77, Lanarks.: Cambuslang, GR 26/65.60. Shrubby ground. P. Macpherson, 1986. 2nd record of established population. *79, Selkirks.: St Mary’s Loch, GR 36/24.23. Rocky bank. R. W. M. Corner, 1989, herb. R.W.M.C. +227/4. COTONEASTER MICROPHYLLUS Wallich ex Lindley 39, Staffs.: Caldon Low, GR 43/ 08.48. Face of lime quarry. B. R. Fowler, 1988. 1st post-1930 record. +227/bul. COTONEASTER BULLATUS Boiss. 77, Lanarks.: Drumoyne, Glasgow, GR 26/54.64. Roadside. P. Macpherson, 1986, herb. P.M., det. J. R. Palmer. 2nd record. +227/die. COTONEASTER DIELSIANUS E. Pritzel ex Diels *12, N. Hants.: Wheatham Hill, GR 41/74.27. Chalky bank. J. D. Fryer, 1985, herb. J.D.F., det. J. R. Palmer & C. A. Stace. a A Lanarks.: Yorkhill, Glasgow, GR 26/55.65. Wood. P. Macpherson, 1985. Drumoyne, Glasgow, PLANT RECORDS 219 GR 26/54.65. Roadside. P. Macpherson, 1986. Both herb. P.M., det. J. R. Palmer. 1st and 2nd records. +227/div. COTONEASTER DIVARICATUS Redh. & Wils. *29, Cambs.: Near Chrishall Grange, GR 52/43.43. Shelterbelt of Pinus nigra. D. E. Coombe, 1989, CGE. *77, Lanarks.: Meadowside, Glasgow, GR 26/55.66. Waste ground. P. Macpherson, 1986, herb. P. M., det. J. R. Palmer. +227/fra. COTONEASTER FRANCHETII Boiss. *77, Lanarks.: Hutchestontown, GR 26/59.64. Along old wall. P. Macpherson, 1986, herb. P. M., det. J. R. Palmer. +227/san. COTONEASTER SANGUINEUS Y1Ii *77, Lanarks.: Cambuslang, GR 26/64.60. Shrubby waste ground. P. Macpherson, 1987, herb. P.M., det. J. R. Palmer. +227/wat. COTONEASTER X WATERERI Exell *12, N. Hants.: Ashford Hanger, Steep, GR 41/ 73.26. Clearing in chalk hanger. J. D. Fryer, 1985, herb. J.D.F., det. J. R. Palmer & C. A. Stace. *77, Lanarks.: Bellahouston, GR 26/64.60. Old railway line. P. Macpherson, 1987, herb. P.M., det. J. R. Palmer. 232/7. SORBUS TORMINALIS (L.) Crantz 47, Monts.: R. Banwy near New Bridge, Meifod, GR 33/13.10. Steep, wooded river bank. P. M. Benoit & M. Wainwright, 1989, NMW. Ist post-1930 record. +235/3. SEDUM SPURIUM Bieb. *39, Staffs.: Cellarhead, GR 33/96.47. Disused sand quarry. I. J. Hopkins, 1988. +235/4. SEDUM DASYPHYLLUM L. 41, Glam.: Rhoose, GR 31/0.6. Disused quarry. J. P. Curtis, 1989. Only extant locality. 237/1. CRASSULA TILLAEA Lester-Garland *2, E. Cornwall: Par, GR 20/08.53. Areas of sandy gravel in caravan site above beach. R. M. Belringer & T. E. Griffiths, 1988, det. Lady R. FitzGerald & R. J. Murphy. +237/hel. CRASSULA HELMSII (T. Kirk) Cockayne 46, Cards.: Plas Gogerddan, GR 22/63.83. Pond. A. Jones, 1989. 2nd record. *47, Monts.: Llangadfan, GR 33/02.13. Mud by man-made pool. M. Wainwright, 1989, det. F. H. Dawson. *53, S. Lincs.: Swanholme, GR 43/94.68. Old sand and gravel pit. T. Smith, 1985. *54, N. Lincs.: Double Drain, Hirst Priory, GR 44/77.09. Large drain. H. E. Stace, 1986. +239/7. SAXIFRAGA CYMBALARIA L. 35, Mons.: St Maughan’s, GR 32/4.1. Garden weed. P. C. Hall, 1980. 2nd record. 239/8. SAXIFRAGA TRIDACTYLITES L. *75, Ayrs.: Nobel Works, Ardeer, GR 26/28.40. Sandy trackside. A. McG. Stirling, 1988, GL. +240/1. TELLIMA GRANDIFLORA (Pursh) Douglas ex Lindley 80, Roxburghs.: R. Tweed below Littledean Tower, GR 36/63.31. River bank. R. W. M. Corner, 1989, herb. R.W.M.C. 2nd record. 246/2. RIBES SPICATUM Robson 67, S. Northumb.: R. East Allen near Wooley Scar, GR 35/ 83.54. River bank. G. A. & M. Swan, 1974, herb. G.A.S. 1st post-1930 record. *68, Cheviot: Denwick, GR 46/21.14. Disused limestone quarry. G. A. Swan & R. S. G. Thompson, 1974, herb. G.A.S. 1st native record, but probably now extinct here. North bank of R. Coquet, GR 46/09.00. River bank. G. A. Swan, 1989, herb. G.A.S. 2nd record. 251/1. DAPHNE MEZEREUM L. +*43, Rads.: Dolyhir, GR 32/24.58. Old limestone quarry. I. D. Soane & R. G. Woods, 1989. 44, Carms.: Coedydd Carmel S.S.S.I., GR 22/59.16. Limestone woodland. R. N. Stringer, 1989. 2nd record. 251/2. DAPHNE LAUREOLA L. +*43, Rads.: Dolyhir, GR 32/24.58. Roadside verge. I. D. Soane & R. G. Woods, 1989. 2nd record. 254/7. EPILOBIUM TETRAGONUM L. subsp. TETRAGONUM *67, S. Northumb.: Near Anick, GR 35/95.65. Margin of pond. G. A. Swan, 1989, herb. G.A.S., det. T. D. Pennington. 220 PLANT RECORDS 254/9 xX 5. EPILOBIUM OBSCURUM Schreber X E. ROSEUM Schreber *67, S. Northumb.: Morpeth, GR 45/18.86. Garden weed. G. A. Swan, 1988, herb. G.A.S., det. T. D. Pennington. *256/2. OENOTHERA ERYTHROSEPALA Borbas *80, Roxburghs.: Allan Water, Galashiels, GR 36/51.36. Refuse tip. C. O. Badenoch, 1987, E, det. D. R. McKean. +256/3. OENOTHERA STRICTA Ledeb. ex Link 38, Warks.: Leamington Spa, GR 42/33.65. Allotment. J. W. Partridge, 1989, WAR, det. J. C. Bowra. 1st post-1930 record. +256/fal. OENOTHERA FALLAX Renner *25, E. Suffolk: Cotton, GR 62/07.66. Roadside verge. R. Addington, 1989, det. J. C. Bowra. 258/1. CIRCAEA LUTETIANA L. 94, Banffs.: Forglen, GR 38/69.52. Woodland. J. Edelsten, 1987, E, det. P. M. Benoit. 2nd record. 261/1. HIPPURIS VULGARIS L. +45, Pembs.: Blaencilgoed quarry, Ludchurch, GR 22/15.10. Flooded quarry. S. B. Evans & J. W. Donovan, 1989. 2nd record. 262/5. CALLITRICHE HERMAPHRODITICA L. *53, S. Lincs.: Teale’s Pits, Whisby, GR 43/91.67. Old gravel pits. I. Weston, 1985, det. N. T. H. Holmes. Swanholme, GR 43/94.68. Old sand and gravel pits. H. Drewett & T. Smith, 1986. 1st and 2nd records. 262/6. CALLITRICHE TRUNCATA Guss. *53, S. Lincs.: Towthorpe Hollow Pond, Belton House, GR 43/92.38. Pond. K. Hearn, 1984. 277/2. TORILIS ARVENSIS (Hudson) Link 11, S. Hants.: Upper Hamble Country Park, Botley, GR 41/50.11. Arable field. M. Southam, 1987. Only extant locality. 300/2. OENANTHE PIMPINELLOIDES L. 7, N. Wilts.: Melksham Without, GR 31/93.62. Pasture. D. Green, 1988, herb. D. G. 2nd record. +309/3. PEUCEDANUM OSTRUTHIUM (L.) Koch 79, Selkirks.: Ettrick Water, Oakwood Mill Farm, GR 36/44.26. River bank. R. W. M. Corner, 1989, herb. R.W.M.C. 2nd extant locality. 311/42 X 1. HERACLEUM MANTEGAZZIANUM Sommier & Levier X H. SPHONDYLIUM L. ise Ayrs.: Maidenhead Bay, GR 26/21.08. Sandy ground. A. McG. Stirling, 1987. *99, Dunbarton: Kirkmichael, Helensburgh, GR 26/30.83. Waste ground near stream. A. Rutherford, 1989. 319/7. EUPHORBIA PLATYPHYLLOS L. 38, Warks.: Walton, GR 42/29.52. Edge of field. H. A. Roberts, 1988, WAR, det. J. C. Bowra. 1st record since 1891. 319/8. EUPHORBIA SERRULATA Thuill. +*5, S. Somerset: Curry Rivel, GR 31/41.27. Disused railway line. P. Green, 1988, det. A. C. Leslie. +319/15 X wal. EUPHORBIA ESULA L. X E. WALDSTEINI (Sojak) A. Radcliffe-Smith *39, Staffs.: Coseley, GR 32/94.93. Waste ground. W. A. Thompson, 1987, herb. B.R. Fowler, det. A. Ratcliffe-Smith. 320/1/3. POLYGONUM RURIVAGUM Jordan ex Boreau 7, N. Wilts.: Biddistone, GR 31/87.72. Arable field. D. Green, 1988. Only extant locality. *46, Cards.: Llwynysgaw, Felin-wynt, GR 22/21.52. Beanfield. A. O. Chater, 1989, NMW, det. J. R. Akeroyd. Dominant over c.4 acres. *73, Kirkcudbrights.: Southerness, GR 25/97.54. Sandy ground. O. M. Stewart, 1984, E, det. B. T. Styles. 320/1/bor. POLYGONUM BOREALE (Lange) Small *103, Mid Ebudes: Aird Mor, Balephetrish Bay, Tiree, GR 07/99.47. Upper, sandy beach. C. D. Preston & N. F. Stewart, 1989, CGE, det. J. R. Akeroyd. +320/19 « 20. REYNOUTRIA JAPONICA Houtt. X R. SACHALINENSIS (Friedrich Schmidt Petrop.) Nakai *11, S. Hants.: Sarisbury, GR 41/50.07. Under trees by fence. R. M. Veall, 1988, herb. R.P. Bowman, det. J. P. Bailey 325/1/2. RUMEX ACETOSELLA subsp. ANGIOCARPUS Murb. 46, Cards.: R.A.E. site, Aberporth, GR 22/24.52. Disturbed ground. J. R. Akeroyd, 1989. 2nd record. PLANT RECORDS 221 325/8. RUMEX LONGIFOLIUS DC. 39, Staffs.: 2 km S. of Onecote, GR 43/04.53. Roadside verge. I. W. Brown 1989, det. J. R. Akeroyd. 1st post-1930 record. 325/11 X 13. RUMEXx crIspUS L. X R. PULCHER L. *12, N. Hants.: Chilbolton Common, GR 41/39.40. J. Fryer, 1988, herb. A. Brewis, det. J. R. Akeroyd. 325/12 X 13. RUMEX OBTUSIFOLIUS L. X R. PULCHER L. *12, N. Hants.: Chilbolton Common, GR 41/39.40. J. Fryer, 1988, herb. A. Brewis, det. J. R. Akeroyd. 325/15 X 13. RUMEX CONGLOMERATUS Murray X R. PULCHER L. *12, N. Hants.: Chilbolton Common, GR 41/39.40. J. Fryer, 1988, herb. A. Brewis, det. J. R. Akeroyd. 325/15 X 17. RUMEX CONGLOMERATUS Murray X R. PALUSTRIS Sm. *16, W. Kent: Shorne Marshes, GR 51/69.74. Edge of grazing marsh dyke. A. C. Leslie, 1986, MNE, det. J. R. Akeroyd. 325/17. RUMEX PALUSTRIS L. +*39, Staffs.: Old Town Hill, Rowley Regis, GR 32/95.86. Demolition site. B. Westwood, 1989, herb. B.R. Fowler, det. J. R. Akeroyd. 343/13 xX 12. SALIX AURITA L. X S. CINEREA L. *94, Banffs.: Hillhead of Mountblairy, GR 38/ 68.52. Wet woodland. J. Edelsten, 1987, E, det. R. D. Meikle. 359/3b. PYROLA ROTUNDIFOLIA subsp. MARITIMA (Kenyon) E. F. Warburg *50, Denbs.: Minera, GR 33/26.51. Dense scrub over wet limestone quarry waste. G. Spencer, 1989, NMW, det. R. G. Ellis. 362/1. MONOTROPA HYPOPITYS L. 39, Staffs.: Sandy Slade, GR 43/00.13. Pine plantation. J. Brough, 1989, herb. B.R. Fowler. 2nd post-1930 record. 372/4. ANAGALLIS MINIMA (L.) E. H. L. Krause *H36, Tyrone: Curran’s Glen, GR 23/35.83. Damp, sandy ground near lake. I. McNeill, 1987, BEL. 382/1. CENTAURIUM PULCHELLUM (Swartz) Druce *68, Cheviot: Bamburgh Links, GR 46/ 19.34. Fixed sand dune. G. A. Swan, 1988. Budle, GR 46/16.36. Grassy flush near sea. G. A. Swan, 1989, both herb. G.A.S., det. F. Ubsdell. Ist and 2nd confirmed records. 383/1. BLACKSTONIA PERFOLIATA (L.) Hudson +*67, S. Northumb.: West Hartford Farm, GR 45/25.78. Central reservation of dual carriageway. H. Watson, 1987, herb. G.A. Swan. Ist confirmed record. 385/1. GENTIANELLA CAMPESTRIS (L.) BOrner 12, N. Hants.: Highclere Park, GR 41/45.60. Old herb-rich grassland. A. Byfield, 1986, det. F. Rose. 1st record for 75 years. 385/4. GENTIANELLA ANGLICA (Pugsley) E. F. Warburg *5,S. Somerset: West Hatch, GR 31/ 27.21. Limestone grassland. M. Davidson, 1989. 387/1. NYMPHOIDES PELTATA (S. G. Gmelin) O. Kuntze +*80, Roxburghs.: Bowden Moor Reservoir, GR 36/54.31. Margin of reservoir. R. W. M. Corner, 1986, herb. R.W.M.C. +PHACELIA TANACETIFOLIA Bentham *12, N. Hants.: Berry Head Farm, GR 41/77.28. Grass weed. F. Finucane, 1973, det. J. E. Lousley. Kingsley, GR 41/78.38. Abundant garden weed. A. Macey, 1982, still present in 1988, det. F. Rose. 1st and 2nd records. 392/1. SYMPHYTUM OFFICINALE L. *79, Selkirks.: S. side of R. Ettrick below Annelshope, GR 36/30.16. Carex acuta dominated fen. R. W. M. Corner, 1988, herb. R.W.M.C., det. F. H. Perring. 392/6. SYMPHYTUM TUBEROSUM L. +*8, S. Wilts.: Berwick St Leonard, GR 31/92.33. Roadside verge. Fonthill Bishop, GR 31/93.33. Shaded, damp waste ground. Both V. Hopkinson, 1987, det. A. M. Hutchison. ist and 2nd records. +392/7. SYMPHYTUM IBIRICUM Steven *8, S. Wilts.: West Ashton, GR 31/88.55. J. Swanbor- ough, 1973. Fonthill Gifford, GR 31/93.31. Roadside verge. J. Hindley, 1987, det. A. M. Hutchison. 1st and 2nd records. 67, S. Northumb.: Halton, GR 35/99.67. By churchyard wall. G. A. Swan, 1987, herb. G.A.S. 2nd record. 222 PLANT RECORDS 400/8 umb. MyosoTis ARVENSIS (L.) Hill subsp. UMBRATA (Rouy) O. Schwarz *G7;),S. Northumb.: Hag Wood near Whitley Chapel, GR 35/91.57. Edge of wood. G. A. & M. Swan, 1989. *68, Cheviot: Warkworth, GR 46/25.06. G. A. Swan, 1989. Hazely Hill, GR 36/95.40. Woodland. G. A. & M. Swan, 1989. 1st and 2nd records. All herb. G.A.S., det. D. Welch. 400/10 glo. MyosoTIS RAMOSISSIMA Rochel subsp. GLOBULARIS (Samp.) Grau *68, Cheviot: Ross Links, GR 46/13.39. Sand dune. G. A. Swan, 1989, herb. G.A.S., det. D. Welch. 407/1. CUSCUTA EUROPAEA L. 12, N. Hants.: Ashford Hill, GR 41/56.62. Nettles on stream- bank. P. Brough, 1988. 1st record since 1928. 7409/1. LycrUM BARBARUM L. *75, Ayrs.: Shalloch park, Girvan, GR 25/18.96. Roadside verge. A. Rutherford, 1989. Bennane Lea, GR 25/09.85. Sandy roadside near sea. A. McG. Stirling & A. Rutherford, 1989. 1st and 2nd records. +409/2. LycIUM CHINENSE Miller *12, N. Hants.: Shipton Bellinger, GR 41/23.44. Roadside hedge. R. P. Bowman, 1975, herb. R.P.B., det. F. Rose. *77, Lanarks.: Westburn, GR 26/ 65.60. Hedge. P. Macpherson, 1989, herb. P.M. +413/4. SOLANUM SARRACHOIDES Sendtner *7, N. Wilts.: Bromham, GR 31/96.64. Market gardens. D. Green & R. Randall, 1986. 416/7 X 1. VERBASCUM NIGRUM L. X V. THAPSUS L. *14, E. Sussex: Crawley, GR 51/28.37. Railway embankment. A. G. Hoare, 1989, det. I. K. Ferguson. 420/3 x 4. LINARIA REPENS (L.) Miller X L. VULGARIS Miller *99, Dunbarton: Bearsden Station, GR 26/54.71. Stony ground, with L. repens. A. McG. Stirling, 1989. 7425/3. MIMULUS MOSCHATUS Douglas ex Lindley *77, Lanarks.: Westburn near Glasgow, GR 26/65.60. Dug-out sandy area. P. & A. C. Macpherson, 1989, herb. P.M. 430/20 hed. VERONICA HEDERIFOLIA L. subsp. HEDERIFOLIA 80, Roxburghs.: St Boswell’s, GR 36/58.30. Shaded earthy bank. R. W. M. Corner, 1989, herb. R.W.M.C. 2nd record. *99 Dunbarton: Overtoun, GR 26/41.75. Lane-side. A. McG. Stirling, 1985, herb. A.McG.S. 430/20 luc. VERONICA HEDERIFOLIA subsp. LUCORUM (Klett & Richter) Hartl *75, AyIs.: Barassie, GR 26/32.33. Garden weed on sandy soil. A. McG. Stirling, 1989. *99, Dunbarton: Helensburgh, GR 26/30.82. Base of wall by path. A. McG. Stirling, 1989. Montrose Street, Helensburgh, GR 26/29.82. A. Rutherford, 1989, herb. A. McG. Stirling, det. A.McG.S. Ist and 2nd records. 430/22. VERONICA POLITA Fries 94, Banffs.: Portsoy, GR 38/58.65. Garden weed. J. Edelsten, 1987, E. 2nd record. 433/2 cal. RHINANTHUS MINOR L. subsp. CALCAREUS Wilmott 7, N. Wilts.: Aston Keynes, GR 41/0.9. Limestone gravel. D. Green, 1988, det. D. J. Hambler. 2nd extant locality. 435/1/1 X 13. EUPHRASIA MICRANTHA Reichenb. X E. NEMoROSA (Pers.) Wallr. *12, N. Hants.: Blackbushe, Yateley, GR 41/81.59. Heathland on old airfield. F. Rose & C. R. Hall, 1988, herb. R.P. Bowman, det. A. J. Silverside. 435/1/2. EUPHRASIA SCOTTICA Wettst. *35, Mons.: Twmbarlwn, GR 31/2.9. Wet grassland on mountain slopes. T. G. Evans, 1986, herb. T.G.E., det. A. J. Silverside. 435/1/5. EUPHRASIA FOULAENSIS Townsend ex Wettst. *98, Main Argyll: Cullipool, Luing Island, GR 17/73.12. Grassland by sea. P. F. Yeo, 1988, CGE. 435/1/15. EUPHRASIA CONFUSA Pugsley *25, E. Suffolk: Between Sizewell and Thorpeness, GR 62/47.61. At top of beach, below heathland. E. M. Hyde, 1985, IPS, det. A. J. Silverside. 435/1/15 X 2. EUPHRASIA CONFUSA Pugsley X E. scoTTica Wettst. *77, Lanarks.: Carnwell, GR 36/0.4. Heath. P. Macpherson, 1988, herb. P.M., det. A. J. Silverside. 435/1/17. EUPHRASIA ARCTICA subsp. BOREALIS (Townsend) Yeo *12, N. Hants.: Winnal Moors, Winchester, GR 41/48.30. Grass path along stream. R. P. Bowman, 1985. Bransbury PLANT RECORDS 223 Common, GR 41/41.41. Damp, alluvial fen grassland. R. P. Bowman, 1988. Both herb. R.P.B., det. A. J. Silverside. 1st and 2nd records. 435/1/17 < 13. EUPHRASIA ARCTICA subsp. BOREALIS (Townsend) Yeo X E. NEMOROSA (Pers.) Wallr. *12, N. Hants.: Bransbury Common, GR 41/41.41. Fen meadow. F. Rose, 1988, herb. R.P. Bowman, det. A. J. Silverside. 437/1. PARENTUCELLIA VISCOSA (L.) Caruel 8, S. Wilts.: Scotchel Green, Pewsey, GR 41/ 16.60. Rough grassland. R. Dauntsey, 1988, det. A. M. Hutchison. 2nd record. 1439/2. LATHRAEA CLANDESTINA L. 4, N. Devon: Stowford, GR 20/40.87. Extensive patches in wet woodland. J. Heath, 1989. 2nd record. *35, Mons.: Lydart, GR 32/50.09. Old green lane. S. Harper, 1987, herb. T.G. Evans. 1st Welsh record. 440/3. OROBANCHE RAPUM-GENISTAE Thuill. 12, N. Hants.: Silchester Common, GR 41/62.62. P. Brough, 1987. 2nd record. 440/10. OROBANCHE HEDERAE Duby **12, N. Hants.: Alresford, GR 41/58.32. On Hedera under beech. A. A. Butcher, 1989, det. F. J. Rumsey as forma monochroma G. Beck. 455/4. SALVIA VERBENACA L. 52, Anglesey: Penmon Priory, GR 23/62.80. Limestone outcrop. M. R. Davies, 1989, det. R. H. Roberts. 1st post-1930 record. 459/3. STACHYS ARVENSIS (L.) L. 7, N. Wilts.: Sandy Lane, GR 31/96.67. Arable land. D. Green, 1985. 2nd extant locality. +461/1 arg. LAMIASTRUM GALEOBDOLON subsp. ARGENTATUM (Smejkal) Stace *67, S. North- umb.: Near Ponteland, GR 45/15.73. Edge of small wood. G. A. & M. Swan, 1976, herb. G.A.S. Hart Burn near Rothley Lodge, GR 45/04.87. Shady bank. G. A. & M. Swan, 1984. Ist and 2nd records. *79, Selkirks.: Island at confluence of Ettrick and Yarrow Waters, GR 36/44.27. Riverbank. D. J. Methven, 1988, herb. R.W.M. Corner, det. A. McG. Stirling. 1st localised record. 462/3. LAMIUM HYBRIDUM Vill. 11, S. Hants.: Gardners Lane, Romsey, GR 41/33.19. Winter wheat. P. J. Wilson, 1987. Only extant locality. 472/1 int. PLANTAGO MAJOR subsp. INTERMEDIA (DC.) Arcangeli *67,S. Northumb.: Morpeth, GR 45/18.86. Garden weed on clay soil. G. A. Swan, 1988. *68, Cheviot: Near Whiteadder Water, GR 36/95.52. Waste ground. G. A. Swan, 1989. Both herb. G.A.S., det. J. R. Akeroyd. 475/2. CAMPANULA TRACHELIUM L. +73, Kirkcudbrights.: South-west of Laudlaugh Bridge, GR 25/79.71. Roadside ditch. O. M. Stewart, 1989, E. 1st post-1930 record. 7475/4. CAMPANULA LACTIFLORA Bieb. *79, Selkirks.: Sunderland Hall, GR 36/48.31. Steep bank above river. R. W. M. Corner, 1986, E, det. D. R. McKean. 475/8. CAMPANULA PATULA L. 39, Staffs.: Upper Arley, GR 32/75.80. Grazed woodland. M. Smith, 1987. 1st post-1930 record. 479/1. JASIONE MONTANA L. 38, Warks.: Middleton, GR 42/19.98. Waste ground. M. J. Senior, 1989, WAR, det. J. C. Bowra. 1st record since 1874 record in this area. 53.9. lines.: Doddington Road, Lincoln, GR 43/94.67. Waste ground. V. Pennell, 1984. 2nd record. 485/3. GALIUM MOLLUGO L. *79, Selkirks.: S. side of Gala Water, Galashiels, GR 36/47.37. Old railway track. J. A. Murray, 1986, herb. R.W.M. Corner. 485/9. GALIUM DEBILE Desv. *8, S. Wilts.: North Charlton water meadows, GR 41/17.24. Margin of drainage ditch. N. L. Chadwick, 1984, det. J. Ounsted. Teffont, GR 31/99.31. Dry pond. V. Hopkinson, 1988, det. A. M. Hutchison. 1st and 2nd records. +489/1. SYMPHORICARPOS ALBUS (L.). S. F. Blake var. LAEvIGATUS (Fernald) S. F. Blake *99, Dunbarton: Helensburgh, GR 26/30.82. Woods. A. Rutherford, c. 1979. 1st record of species. 7491/1. LONICERA XYLOSTEUM L. 50, Denbs.: Loggerheads near Mold, GR 33/19.62. B.Ing, 1988. 2nd record. 224 PLANT RECORDS 4491/4. LONICERA CAPRIFOLIUM L. *79, Selkirks.: Philiphaugh, Yarrow, GR 36/44.27. Wood- land. A. J. Smith, 1988, herb. R.W.M. Corner. [+491/cae. LONICERA CAERULEA L. 57, Derbys.: Delete record published in Watsonia 16: 192 (1986), plant is L. involucrata (Richardson) Banks ex Sprengel, det. C. A. Stace, and is not naturalized. | 494/2. VALERIANELLA CARINATA Loisel. 7, N. Wilts.: Cotswold Water Park, GR 41/01.94. Dumped soil. S. C. Holland, 1988. Only extant locality. 497/2. DIPSACUS PILOSUS L. 41, Glam.: East Aberthaw, GR 31/0.6. Streamside. J. P. Curtis, 1987. 2nd record, 506/2 < 1. SENEcIO AQguaTicus Hill x S. JACOBAEA L. *38, Warks.: Hampton Lucy, GR 42/ 25.55. Churchyard. J. W. Partridge, 1988, WAR, det. C. Jeffrey. +506/9. SENECIO TANGUTICUS Maxim. *76, Renfrews.: Cowglen, Glasgow, GR 26/54.60. Roadside. E. L. S. Lindsay & A. C. Macpherson, 1981, herb. P. Macpherson, det. M. McC. Webster. Still present. +506/13. SENECIO FLUVIATILIS Wallr. 67, S. Northumb.: Elson Burn, GR 35/93.92. Stream- bank. G. A. & M. Swan, 1989, herb. G.A.S. 2nd record, 1st this century. +509/4. PETASITES FRAGRANS (Vill.) C. Presl 77, Lanarks.: Thorntonhall, GR 26/59.55. Path through wood. P. Macpherson, 1989. 2nd record. +518/3. SOLIDAGO GIGANTEA Aiton *26, W. Suffolk: Elmswell, GR 62/00.66. Open glade in wood. R. Addington, 1989, det. E. M. Hyde. *39, Staffs.: Weston Sprink, GR 33/93.43. Scrub in old pasture. Berry Hill, GR 33/90.46. Waste ground. Both I. J. Hopkins, 1988. 1st and 2nd records. 4519/7 X 6. ASTER LAEVIS L. X A. NOVI-BELGII L. *39, Staffs.: Wombourne, GR 32/87.92. Roadside verge. C. B. Westall, 1987, det. P. F. Yeo. 526/3. ANTHEMIS ARVENSIS L. 33, E. Gloucs.: Aldsworth, GR 42/18.11. Edge of arable field. R. J. Cooper & S. C. Holland, 1989. Only extant locality. 4534/1. COTULA CORONOPIFOLIA L. *39, Staffs.: Drummond Road, Stafford, GR 33/92.24. Slightly saline marsh. R. Hill, 1989, herb. B.R. Fowler. 539/3. CARDUUS NUTANS L. +*103, Mid Ebudes: West of A’ Chroic, Coll, GR 17/22.62. Roadside by sea. C. D. Preston & N. F. Stewart, 1989. 558/1/94. HIERACIUM DURICEPS F. J. Hanb. *93, N. Aberdeen: Cabrach, GR 38/36.21. Rock ledge. D. Welch, 1987, herb. D. W., det. P. D. Sell. 558/1/193. HIERACIUM PLACEROPHYLLOIDES Pugsley *73, Kirkcudbrights.: Kirkpatrick Durham, GR 25/76.76. Grassy laneside bank. J. Bevan, 1986, herb. J.B. 558/1/196. HIERACIUM SPARSIFOLIUM Lindeb. *75, Ayrs.: Stinchar Bridge, Straiton, GR 26/ 39.95. Rocks in stream. A. McG. Stirling, 1989. 558/1/210. H1ERACIUM STRICTIFORME (Zahn) Roffey *93, N. Aberdeen: Cabrach, GR 38/38.26. Rock ledge in gorge. D. Welch, 1988, ABD, det. P. D. Sell. 558/1/211. HtERACIUM RETICULATUM (Lindeb.) Lindeb. *93, N. Aberdeen: Ravenscraig, GR 48/09.48. Walls of ruined castle. D. Welch, 1988, ABD, det. P. D. Sell. 558/1/221. HIERACIUM RIGENS Jordan *35, Mons.: Slade Wood, Rogiet, GR 31/45.89. Pathside in wood. T. G. Evans, 1986, herb. T.G.E., det. J. Bevan. 558/1/222. HIERACIUM SALTICOLA (Sudre) Sell & C. West +*35, Mons.: Newport Docks, GR 31/31.86. Rail ballast. T. G. Evans, 1980, herb. T.G.E., det. J. Bevan. PLANT RECORDS 225 558/2/1 eur. PILOSELLA OFFICINARUM subsp. EURONOTA (Naegeli & Peter) Sell & C. West n1S), Ayrs.: Muck Water, Dalmellington, GR 26/48.05. Stream bank. A. McG. Stirling, 1989, herb. A.McG.S. 558/2/1 off. PILOSELLA OFFICINARUM F. W. Schultz & Schultz Bip. subsp. OFFICINARUM 75; Ayrs.: Muck Water, Dalmellington, GR 26/48.05. Stream bank. A. McG. Stirling, 1989, herb. A.McG.S. 559/5. CREPIS BIENNIS L. +2, E. Cornwall: Stratton, GR 21/22.06. Grassy bank by road. L. J. Margetts, 1989, herb. L.J.M. 1st post-1930 record. 561/1. BALDELLIA RANUNCULOIDES (L.) Parl. 21, Middlesex: Glebelands Wood, East Finch- ley, GR 51/26.91. Edge of shallow pond. D. Bevan, 1988, herb. D.B. Only extant locality. 567/1. HyDROCHARIS MORSUS-RANAE L. 67, S. Northumb.: Near Healey Wood, GR 45/23.83. Pond. N. Scott, 1986, herb. G.A. Swan. 2nd record, Ist this century. 7570/3. ELODEA NUTTALLII (Planchon) St. John 26, W. Suffolk: Great Ashfield, GR 62/00.67. Pond. R. Addington, 1989, det. M. N. Sanford. 2nd record *46, Cards.: Rhos Pil-bach, Plwmp, GR 22/36.56. Pond in pasture. Afon Aeron 1 km E. of Talsarn, GR 22/55.56. River. Both A. O. Chater, 1989, NMW, det. D. A. Simpson. Ist and 2nd records. 67, S. Northumb.: Queen Elizabeth Country Park, GR 45/28.89. Artificial lake. G. A. Swan, 1989, herb. G.A.S., det. D. A. Simpson. 2nd record. *H36, Tyrone: Lough Neagh S. of Stanniards Point, GR 23/96.77. Lough Neagh, Ardboe Point, GR 23/96.76. Both D. A. Simpson & J. A. N. Parnell, 1984. Ist and 2nd records. 577/6. POTAMOGETON GRAMINEUS L. 68, Cheviot: Nelly’s Moss Lakes, GR 46/08.02. G. A. Swan, 1989. 2nd record. 577/7 X 8. POTAMOGETON ALPINUS Balbis X P. PRAELONGUS Wulfen *H35, W. Donegal: Lough between Lough Goragh and Carryblagh, GR 24/23.42. C. D. Preston & N. F. Stewart, 1989, CGE. 1st Irish record. 577/11. POTAMOGETON FRIESII Rupr. 7, N. Wilts.: Kennet and Avon Canal, Allington, GR 41/ 07.62. J. Lovell, 1987, det. D. Green. 2nd extant locality. *H35, W. Donegal: Melmore Lough, GR 24/12.43. Coastal lake. C. D. Preston et al., 1989, CGE. 577/12. POTAMOGETON RUTILUS Wolfg. 103, Mid Ebudes: Loch Ballyhaugh, Coll, GR 17/ 17.58. Machair loch. C. D. Preston & N. F. Stewart, 1989, CGE. ist Coll and 2nd vice-county record. *108, W. Sutherland: Loch Awe, GR 29/24.15. E. Charter, 1988, NCCE, det. N. T. H. Holmes & C. D. Preston. 577/16. POTAMOGETON TRICHOIDES Cham. & Schlecht. 2, E. Cornwall: Bude Canal, Bude, GR 21/21.04. Canal. C. D. Preston & N. F. Stewart, 1989, CGE. Only extant locality. 54, N. Lincs.: Snakeholme Drain, Bardney, GR 53/10.71. T. C. G. Rich & I. Weston, 1987, CGE, det. C. D. Preston. Barton and Barrow Clay Pits, GR 54/03.22. Disused clay pit. C. D. Preston, H. E. Stace & N. F. Stewart, 1987, CGE. 1st and 2nd post-1930 records. 577/19 X 9. POTAMOGETON CRISPUS L. X P. PERFOLIATUS L. *73, Kirkcudbrights.: Carlingwark Loch, GR 25/76.61. Shallow water near edge of loch. C. D. Preston & O. M. Stewart, 1989, CGE. 577/19 X 11. POTAMOGETON crispus L. X P. FRIESII Rupr. *73, Kirkcudbrights.: Carlingwark Loch, GR 25/76.61. Shallow water near edge of loch. O. M. Stewart, 1989, E, det. C. D. Preston. Ist Scottish record. 577/20. POTAMOGETON FILIFORMIS Pers. 80, Roxburghs.: Lady Moss, GR 36/52.30. Pond. R. W.M. Corner, 1989, CGE, det. C. D. Preston. 2nd record. 581/1. NAJAS FLEXILIS (Willd.) Rostk. & W. L. E. Schmidt *97, Westerness: Loch a’ Bhada Dharaich, GR 17/69.94. Water 1m deep. A. E. Newton & A. J. Shine, 1977, CGE, det. A. C. Jermy. 103, Mid Ebudes: Loch an t-Sagairt, Coll, GR 17/25.60. C. D. Preston & N. F. Stewart, 1989, CGE. 2nd Coll record. 226 PLANT RECORDS 591/1b. ASPARAGUS OFFICINALIS subsp. PROSTRATUS (Dumort.) Corb. *2, E. Cornwall: Rocky Valley near Tintagel, GR 20/07.89. Steep, rocky slope. R. E. N. & C. J. Smith, 1988, det. L. J. Margetts. +593/1. LILIUM MARTAGON L. *99, Dunbarton: Near Bearsden Station, GR 26/54.71. Wooded waste ground. A.McG. Stirling et al., 1989. 600/42 xX 1. HYACINTHOIDES HISPANICA (Miller) Rothm. X H. Non-scripta (L.) Chouard ex Rothm. *39, Staffs.: Malthouse Lane, Caverswall, GR 33/94.45. Roadside verge. I. J. Hopkins, 1987. +605/2. JUNCUS TENUIS Willd. *7, N. Wilts.: Winsley, GR 31/80.61. Roadside. J. Presland, 1987, herb. D. Gréen. 605/4. JUNCUS COMPRESSUS Jacq. 5, S. Somerset: Durleigh Reservoir, GR 31/26.36. Barer areas just above high-water mark. P. Green, 1989. 2nd extant locality. 39, Staffs.: Drummond Road, Stafford, GR 33/92.24. Marsh. R. Hill, 1989, herb. B.R. Fowler. 1st post-1930 record. 605/amb. JUNCUS AMBIGUUS GuSs. *108, W. Sutherland: Clachtoll, GR 29/04.27. Damp, sandy ground at head of beach. Badnabay, GR 29/22.46. Muddy track leading to salt marsh. Both E. Norman, 1988, det. T. A. Cope. 1st and 2nd records. +607/7. ALLIUM CARINATUM L. *77, Lanarks.: Bothwell, GR 26/69.58. Bank of R. Clyde. J. R. S. Lyth, 1989, GL. 607/8. ALLIUM SCHOENOPRASUM L. +*77, Lanarks.: Meadowside, Glasgow, GR 26/55.66. Roadside. P. Macpherson, 1989, herb. P.M. +*80, Roxburghs.: Rule Water below Bedrule, GR 36/59.17. Rocky edge of river. R. W. M. Corner, 1989. +607/nig. ALLIUM NIGRUM L. *5,S. Somerset: Staple Plain, West Quantoxhead, GR 31/11.41. Rough pasture. C. J. Giddens, 1989, det. A. L. Grenfell. 4616/1. IRIS SPURIA L. *6, N. Somerset: Sand Point, GR 31/32.66. Limestone scrub. A. L. Grenfell, c.1950. Still present in 1988. +616/sib. IRIS SIBIRICA L. *83, Midlothian: Musselburgh, Inveresk, GR 36/35.71. Rough grass by railway. D. R. McKean, 1988, E. 625/4. EPIPACTIS LEPTOCHILA (Godfrey) Godfrey 8, S. Wilts.: Hound Wood, Winterslow, GR 41/22.30. Edge of wood. M. N. Jenkinson, 1984, det. R. P. Bowman. 2nd record. 625/6. EPIPACTIS PHYLLANTHES G. E. Sm. *6, N. Somerset: Leigh Woods, GR 31/55.74. Woodland by towpath. P. & I. Green, 1988, det. A. J. Richards. 635/1. COELOGLOSSUM VIRIDE (L.) Hartman 79, Selkirks.: Selkirk Golf Course, GR 36/48.28. Grassland. M. J. Roger, 1986. 1st record since 1883. 643/1 x 3a. DACTYLORHIZA FUCHSII (Druce) So6 Xx D. INCARNATA (L.) S06 subsp. INCAR- NATA *64, Mid-W. Yorks.: Upper Wharfedale, GR 34/97.67. Wet, basic grassland. F. Horsman, 1989. 643/1 x 3b. DAcTYLORHIZA FUCHSII (Druce) S06 X D. INCARNATA subsp. PULCHELLA (Druce) S06 *64, Mid-W. Yorks.: Upper Wharfedale, GR 34/97.67. Wet, basic grassland. F. Horsman, 1989. 643/3b x 4. DACTYLORHIZA INCARNATA subsp. PULCHELLA (Druce) So6 X D. PRAETERMISSA (Druce) So6 *46, Cards.: Ynys-las dunes, GR 22/60.93. Dune slack. F. Horsman, 1989. 643/3b x 6cam. DACTYLORHIZA INCARNATA Subsp. PULCHELLA (Druce) S06 X D. MAJALIS subsp. CAMBRENSIS (R. H. Roberts) R. H. Roberts *46, Cards.: Ynys-las dunes, GR 22/60.93. Dune slack. F. Horsman, 1989. 643/5. DACTYLORHIZA PURPURELLA (T. & T. A. Stephenson) So6 *43, Rads.: Lane Farm, Rhosgoch, GR 32/18.46. Marshy field. H. L. J. Drewett, 1989. 2nd record. PLANT RECORDS 227 SERAPIAS PARVIFLORA Parl. *2, E. Cornwall: Locality withheld. P. Cobbing, 1989. See BSBI News no. 52, p. 11 (1989). 1st British record. +646/1. ACORUS CALAMUS L. *8, S. Wilts.: Kennet & Avon Canal, Devizes, GR 31/98.61. Canal bank. D. Green, 1986. 1647/1. CALLA PALUSTRIS L. *39, Staffs.: Stockwell Heath, GR 43/05.21. Shallow village pond. B. R. Fowler, 1988. 650/2. LEMNA TRISULCA L. 73, Kirkcudbrights.: Lochrutton Loch, GR 25/89.72. C. D. Preston & O. M. Stewart, 1989, CGE. 2nd record. +650/min. LEMNA MINUSCULA Herter *38, Warks.: Solihull, GR 42/16.81. Pond. J. W. Partridge, 1989, WAR, det. A. C. Leslie. 654/3. ERIOPHORUM LATIFOLIUM Hoppe 44, Carms.: Cae Maesyffynnon near Port Aber, GR 22/77.20. Wet, flushed grassland. I. K. Morgan, 1989. 2nd record, 1st since 1908 record in this area. 50, Denbs.: Near Graianrhyd, GR 33/20.54. Marsh. J. A. Green, 1989, NMW. 2nd record. 657/2. BLYSMUS RUFUS (Hudson) Link *44, Carms.: Ffos Fach Marsh, Bynea, GR 21/55.99. Top of saltmarsh. N. C. C. Field Survey Team, 1988. 658/1. CyPERUS LONGUS L. +25, E. Suffolk: The Grove, Felixstowe, GR 62/30.35. Edge of pond. B. Matthews, 1989, det. M. N. Sanford. 2nd record. 663/10. CAREX VIRIDULA Michx subsp. vIRIDULA *39, Staffs.: Chasewater Reservoir, GR 43/ 03.08. Margin of reservoir. J. P. Martin, 1988, herb. B.R. Fowler, det. A. O. Chater. 663/16 < 17. CAREX ROSTRATA Stokes X C. VESICARIA L. *46, Cards.: Dolaugwyrddon-isaf, Lampeter, GR 22/55.46. Recently created artificial pond. A. O. Chater, 1989, NMW, det. A. O.C., R. W. David & A. C. Jermy. 663/22. CAREX PENDULA Hudson +80, Roxburghs.: Ettrick Water above Lindean Mill, GR 36/ 47.31. Open conifer plantation near river. E. Middleton, 1988, herb. R.W.M. Corner. 2nd record. 663/25. CAREX TOMENTOSA L. 7, N. Wilts.: South of Pike Corner, Minety, GR 41/03.92. Ridge and furrow pasture. L. Wild, 1988, herb. D. Green, det. R. W. David. 2nd extant locality. 663/28. CAREX LIMOSA L. *50, Denbs.: Hiraethog Moor, GR 23/93.55. Soligenous flush. P. Day, 1989, NMW, det. A. O. Chater. 81, Berwicks.: Brotherstone Hill, GR 36/61.35. Peat bog. R. W. M. Corner, 1989. 1st post-1930 record. 663/47 < 46. CAREX ACUTA L. X C. ELATA All. *6, N. Somerset: Axbridge, GR 31/43.53. Ditch. D. Pearman, 1983, herb. D.P., det. A. O. Chater & A. C. Jermy. [663/55. CAREX APPROPINQUATA Schumacher 45, Pembs.: Delete records, which are misidenti- fications of C. diandra Schrank, cf BSBI News no. 53, pp. 17-18, 1989.] 663/69. CAREX ELONGATA L. *47, Monts.: Llangyniew, GR 33/12.09. Salix swamp. W. Metcalfe & M. Wainwright, 1989, NMW, det. A. O. Chater & R. W. David. 663/72 < 54. CAREX CURTA Good. X C. PANICULATA L. *45, Pembs.: Dowrog Common, GR 12/77.26. R. W. David & S. B. Evans, 1989, NMW, det. A. O. Chater & R.W.D. 2nd Welsh and 3rd British record. 667/1 alt. MOLINIA CAERULEA (L.) Moench subsp. ALTIssiIMA (Link) Domin 35, Mons.: Llangedfedd Reservoir, GR 31/3.9. Wet heath. T. G. Evans, 1987, herb. T.G.E. 2nd record. 669/2. GLYCERIA PLICATA (Fries) Fries 2, E. Cornwall: Kingsmill Lake, Landulph, GR 20/ 43.61. Marsh. R. W. Gould, 1989, herb. R.J. Murphy, det. L. J. Margetts & R. J. M. 1st post-1930 record. 670/6b. FESTUCA NIGRESCENS Lam. *77, Lanarks.: Between Glasgow and Renfrew, GR 26/ 51.67. Bare, grassy bank. P. Macpherson, 1986, herb. P.M., det. P. J. O. Trist. 228 PLANT RECORDS 670/9. FESTUCA FILIFORMIS Pourret *75, Ayrs.: Upper Noddesdale, GR 26/26.68. Dry heathy ground. A.McG. Stirling, 1989, herb. A. McG. S. 670/12. FESTUCA GUESTFALICA Boenn. ex Reichenb. 54, N. Lincs.: Seacroft Golf Course, Skegness, GR 53/56.60. I. Weston, 1982, det. P. J. O. Trist. 2nd record. 674/1. DESMAZERIA RIGIDA (L.) Tutin 47, Monts.: Big Forest, Maesmawr, GR 33/15.10. Forest road. M. Wainwright, 1989, det. P. M. Benoit. 2nd record. 677/1. CATABROSA AQUATICA (L.) Beauv. *76, Renfrews.: Lunderston Bay, GR 26/20.73. Streamside on sandy shore. A. McG. Stirling, 1989. 80, Roxburghs.: Primside Bog, GR 36/ 80.26. Fen and ditch. R. W. M. Corner, 1989, herb. R.W.M.C. 2nd extant locality. 681/2. MELICA NUTANS L. 43, Rads.: North of Llanbadarn Fynydd, GR 32/08.79. Wooded roadside bank. R. G. Woods, 1989. 2nd record. +683/4. BROMUS INERMIS Leysser *5, S. Somerset: Williton, GR 31/08.41. Grassy road bank. Lady R. FitzGerald, 1989. 683/5. BROMUS STERILIS L. +93, N. Aberdeen: St Combs, GR 48/04.63. Set-aside field. D. Weich, 1989. 2nd extant locality. 683/8. BROMUS ARVENSIS L. 33, E. Gloucs.: Southfields Farm, Hardwicke, GR 32/77.10. Set- aside arable field. S. C. Holland & P. Wilson, 1989, herb. S.C.H., det. D. M. Barling. Only extant locality. 4683/19. BROMUS CARINATUS Hooker & Arnott *5, S. Somerset: Henstridge, GR 31/72.21. Green lane. I. Green, 1989. *12, N. Hants.: Greywell, GR 41/71.51. C. Hora, 1986, RNG. Near King John’s Castle, GR 41/72.51. Edge of Basingstoke Canal. C. Hora, 1989, herb. Lady A. Brewis, det. A. B. 1st and 2nd records. +687/jub. HORDEUM JUBATUM L. 72, Dumfriess.: A701 road, Moffat, GR 36/04.13. Roadside verge. H. J. Noltie & A. G. Miller, 1988. 2nd record. 688/1. HORDELYMUS EUROPAEUS (L.) C. O. Harz 53, S. Lincs.: Skellingthorpe Wood, GR 43/ 90.72. Ancient wood. F. Aungier, 1981. 2nd extant locality. 694/1 bul. ARRHENATHERUM ELATIUS subsp. BULBOSUM (Willd.) Schibler & Martens *67, S. Northumb.: Coalburn Farm, GR 45/20.83. Edge of cultivated field. G. A. Swan, 1989. 1st localized record. *68, Cheviot: Near Red Burn, GR 46/15.20. Cultivated field. G. A. Swan, 1989. 1st localized record. +697/3. AIRA CARYOPHYLLEA subsp. MULTICULMIS (Dumort.) Bonnier & Layens *67, S. Northumb.: Near Deadwater, GR 35/61.95. Forest track. G. A. & M. Swan, 1989, herb. G.A.S., det. O. M. Stewart. *68, Cheviot: Near Woodbridge, GR 36/95.33. Sandy gravel near roadside. Thrunton Forest, GR 46/07.08. Side of forest road. Both G. A. & M. Swan, 1989, herb. G.A.S. Ist and 2nd records. 701/2b. AGROSTIS VINEALIS Schreber *43, Rads.: Harley Dingle, GR 32/18.64. Damp, shaded locality. R. G. Woods, 1989. 702/1. APERA SPICA-VENTI (L.) Beauv. +7, N. Wilts.: Biddestone, GR 31/87.73. Cornfield. D. Green, 1988. 2nd record. +8, S. Wilts.: Grovely Wood, GR 31/06.33. Margin of barley field. D. Simpson-Green, 1984, det. P. J. O. Trist. Fiddington Sands, GR 41/01.54. Edge of arable field. R. Fussell, 1984, det. A. Hutchison. Ist and 2nd records. 708/3 * 2. ALOPECURUS GENICULATUS L. X A. PRATENSIS L. *14, E. Sussex: Southease, GR 51/ 42.04. Tidal river meadows. E. Norman, 1985, det. P. J. O. Trist. *29, Cambs.: R. Granta, Linton, GR 52/56.46. River bank. P. J. O. Trist, 1982, herb. P.J.O.T. 709/1. MILIUM EFFUSUM L. 81, Berwicks.: North Cleugh, Penmanshiel Wood, GR 36/79.68. Oakwood. M. E. Braithwaite, 1989, herb. M.E.B. 1st post-1930 record. +720/1. SETARIA ITALICA (L.) Beauv. *39, Staffs.: Knypersley Reservoir, GR 33/89.54. Amongst sandstone blocks. I. W. Brown, 1989, det. T. A. Cope. Watsonia, 18, 229-237 (1990) Zoe Book Reviews Flore de la Suisse et des territoires limitrophes. D. Aeschimann & H. M. Burdet. Pp. liv + 597, with 17 pp. of figures. Editions du Griffon, Neuchatel. 1989. Price SF48 (ISBN 2-88006-503-7). The appearance of an entirely new Swiss flora is a rare but welcome event. With the subheading “‘le nouveau Binz’’, the title of this pocket flora pays homage to A. Binz’s School and Excursion flora of 1920, a French translation of which first appeared in 1941. Three later editions of this work appeared, the most recent in 1976. A Pocket Atlas of the Swiss flora by Thommen, Becherer & Antonietti was published in 1983, but the new Flore de la Suisse is the first entirely new flora to appear since Hess, Landolt & Hirzel’s Flora der Schweiz of 1967-72. Dichotomous keys to families, genera and species are provided but there are no familial or generic descriptions. Limited synonymy is provided at the level of species and subspecies, but without literature citations. French vernacular names are given for genera, and translations of specific epithets are provided. For each species, longevity (categories are annual, biennial, perennial, shrub and tree), life-form (Raunkizr’s categories are used together with ‘lianes’) and height are stated, followed by a very brief descriptive text which omits characters used in the keys. Phenology (taking account of altitude, which rather blurs the picture) and ecology (altitudinal zone, habitat category and substrate) are followed by a summary of the plant’s distribution, frequency and status within the area covered by the flora. This includes not only the whole of Switzerland but also the adjacent areas of Savoy, the entire Jura massif, southernmost Germany, Vorarlberg and westernmost Tirol together with the alpine portions of N. Italy. A four-figure number provides a cross-reference to Thommen et al.’s Pocket Atlas illustrations. Subspecies are described, and hybrids listed, in small type. Considerable progress has been made in clarifying the taxonomic treatment of the Swiss and alpine floras in recent decades. The authors state in the introduction that they have followed the nomenclature of Greuter et al.’s Med Checklist for the parts so far published, together with Flora Europaea and certain recent parts of Hegi which deal with the Asteraceae. This is a workable compromise which users of this flora will find easy to follow. The only other flora which has appeared post-Flora Europaea and covers part of its area is Pignatti’s Flora d’Italia, whose lack of portability rivals that of Hess et al. The book has been given a robust plastic binding, and the use of millimetre-ruled graph paper for the end papers is a thoughtful feature which should be widely copied by publishers of pocket fioras. Although one must refer to other works for whole-plant illustrations, the 343 line drawings form a useful crib for features mentioned in the keys which would otherwise require lengthy descriptions, such as the appendices of the involucral bracts of species of Centaurea. For users like myself whose knowledge of French descriptive terminology is limited, there is a very useful glossary; this makes frequent reference to the figures and contains a few un-numbered illustrations of leaf shapes, inflorescence categories and a diagram showing the parts of an orchid flower. Being able to reach a much wider readership than the ‘heavyweight’ floras of the region, this admirable excursion flora upholds the tradition of quality which one associates with the publications of the Conservatoire et Jardin botaniques of the City of Geneva, under whose auspices it appears, and it will give much pleasure to its users. J. R. EDMONDSON Vegetation of the Soviet polar deserts. V. D. Aleksandrova. Pp. xii + 228. Translated by D. Love. Studies in Polar Research; Cambridge University Press. 1988. Price £30 (ISBN 0—-521-32998-1). The living tundra. Y. 1. Chernov. Pp. xiv + 213. Translated by D. Love. Studies in Polar Research; Cambridge University Press. 1985, paperback edition 1988. Price (paperback) £12-95 (ISBN 0-521- 35754-3). 230 BOOK REVIEWS Aleksandrova’s account deals with the most northerly part of the Russian Arctic, where the mean temperature of the warmest month fails to exceed 2°C. It covers the western and relatively oceanic Zemlya Frantsa-Josifa archipelago and the northern part of Novaya Zemlya and the more eastern and continental Severnaya Zemlya archipelago. Polar deserts are characterised by a very sparse vegetation, often intimately related to patterned ground. The dominant plants are lichens and, to a lesser extent, mosses. The 90 or so species of flowering plants mentioned by the author play a very subordinate role. By comparison with the tundra zone to the south, sedges and gamopetalous species are few and regional differentiation of the flora is weak. This is essentially a reference book, much of it being devoted to a detailed consideration of 58 vegetation relevés from Zemlya Frantsa-Josifa. Of particular interest is the remarkably detailed and painstaking systematic account of the seasonal development of the fiowering plants on this archipelago. The living tundra is a really excellent, vivid and absorbing ecological account of the Russian tundra. It is written by an acknowledged and evident expert who is clearly alive to the importance of this still largely untouched ecosystem and also its extreme fragility. Despite dealing exclusively with the Russian tundra — and for that reason alone it is invaluable — the book is essential reading for anyone interested in the Arctic. I myself found particularly interesting and revealing the chapters on tundra landscapes and on snow as an all-pervading ecological factor. The author intersperses animal and plant examples with consummate ease and this familiarity is especially evident and valuable in the long chapter on the diverse relationships between organisms in which he convinces us of the importance of entomophily for many of the flowering plants as well as commenting on the partiality of salmon for drowning lemmings! Together, these two complementary books provide the first comprehensive account of the Russian Arctic readily available in the west. Both authors have been well served by Dr Léve’s readable but unobtrusive translation and helpful introductory notes. G. HALLIDAY Comparative plant ecology. A functional approach to common British species. J. P. Grime, J. G. Hodgson & R. Hunt. Pp. ix + 742, with numerous text figures and tables. Unwin Hyman, London, 1988. Price £85 (ISBN 0-04-581028-1). The heart of this volume is the ecological accounts of British species, each occupying two facing pages, and jointly comprising over three quarters of the book. The majority of the data has been gathered by the authors and their eleven collaborators from the Unit of Comparative Plant Ecology, University of Sheffield, and supported by the Natural Environment Research Council. This collaboration, over 25 years, has resulted in directly comparable data for 281 species, and is summarised, with data for a further 221 species, in a succeeding chapter. The accounts are preceded by four introductory chapters. Study of these is fundamental to understanding the ecological background and theory implicit in the accounts. Data are of three types and are described in detail in Chapter 2. Firstly there are field records, mainly gathered from a vegetation survey covering some 3000 km* surrounding Sheffield and encompassing the range of habitats across the lowland — upland boundary of the Pennines and a wide range of bedrocks and associated soils. The authors point out that this sample area is representative of a much wider area of Britain. These data have been used to produce for each species a habitat diagram with 32 categories, corresponding to single land use/habitat categories; from the same data set are derived histograms showing altitude range, occurrence by slope angle, aspect, bare soil, hydrology and soil pH, together with tables of abundance in each of seven major habitat types. The second group of data come from standardised experiments, providing germina- tion data, occurrence in a soil seed bank, relative growth rate and, unusually for ecological work, 2C DNA amounts. There is also a wide range of information taken from the literature, including phenology, chromosome number and floral dispersal mechanisms. All these data would be sufficient justification for an autecological flora, yet the introductory chapters make clear that the driving forces for this work have been “‘plant strategy concepts BOOK REVIEWS 231 developed at Sheffield, which have been used to provide a unifying conceptual frame-work”’. Chapter 1 sets out the history of such concepts and explains the C-S-R model 1 (competitor — stress tolerator — ruderal) developed by Grime in his book Plant strategies and vegetation processes (1979), and depicted here as a ‘triangular ordination’ diagram, set symbolically in the centre of one page of each species account. The three sides of the triangle, originally referring to the relative importance of competition, stress and disturbance are, however, discarded for derived measurements which have been calculated on the basis of a circle of 91 hexagons superimposed on a symbolic triangle. Non-professional ecologists will find this heavy going and it may well have been more successfu! to have maintained the simpler triangular representations of Grime’s earlier work. A more serious concern is the treatment of trees and shrubs. Of only twelve included in the main accounts (which exclude species such as Corylus avellana, Ilex aquifolium, Prunus spinosa and all Rosa species), Betula and Quercus only appear as genus accounts, while the single Salix account includes both S. caprea and S. cinerea. The chief reason for this treatment is the use throughout the vegetation survey of a 1m? quadrat which only included seedlings and small saplings. In view of the importance of trees and shrubs in colonising and dominating unmanaged lowlands, such a treatment is disappointing, even though balanced to some extent by inclusion of published data. The autecological value of this book is obvious. Yet for the authors this is only a starting point. Chapter 4 deals with ‘the interpretation and use of autoecological accounts’. Proposals range from a suggestion that family history (in an evolutionary sense) will emerge as an important influence on the present characteristics and contemporary ecology of individual species, to a substantial discussion of the relationship of DNA amounts to ecological characteristics. Also included are discussions of phenology, regenerative strategies and germination. One will not agree with everything that is said here. We have given you the bricks and tools, together with some plans, say the authors. It is up to you to build, make other bricks and other plans. This book will be used for these purposes for many years to come. T. T. ELKINGTON Somerset ferns, a field guide. Pat Hill-Cottingham. Pp. 84, with many text-figures. Somerset Archaeological and Natural History Society, Taunton. 1989. Price £2-50, post paid £3 (ISBN 0- 902152-15-7). This attractively produced booklet is a comprehensive guide to the ferns (not the horsetails nor club- mosses) of the county. With an excellent cover illustration in colour of Oreopteris limbosperma and printed on good quality paper which makes it a pleasure to handle, it should stimulate the local study of ferns. It is strictly for use in Somerset, excluding all species not currently occurring in the county. It seems a pity that it excludes Adiantum capillus-veneris, of which several long-established colonies occur in Somerset. After a very readable and well illustrated introduction on the life-cycle of ferns there are dichotomous and multi-access keys to identification, followed by detailed descriptions of all the species covered, in alphabetical sequence of genera and of species within genera. Each species is illustrated by reduced photocopies of actual fronds: these are mostly very satisfactory, though the frond chosen for Dryopteris carthusiana is atypical and fails to show the distinctively narrow parallel-sided shape which contrasts with the more triangular frond of D. dilatata; and the blobs depicting Azolla are not very helpful, though in this case there is an enormously enlarged line- drawing as well. The book concludes with useful sections on the conservation, propagation and cultivation of ferns, a very ample glossary (was it necessary to define ‘triangular’?), lists of synonyms and a bibliography. Unfortunately the book is marred by numerous misprints, and even the errata slip enclosed fails to correct inter alia the dichotomous key, in which errors remain in couplets 7 and 16. Although the two sub-species of Asplenium trichomanes are mentioned (both occurring in Somerset), they are not differentiated. The Polypodium hybrids are dealt with in some detail, but there is no mention of Dryopteris or Polystichum hybrids, some of which certainly occur in the county. A note on the Dryopteris affinis complex curiously tells us which is the commonest subspecies in Scotland, but makes no reference to Somerset. 232 BOOK REVIEWS There is a useful map of the county showing the vice-county boundary, although the accounts of distribution of each species are not consistently on a v.c. basis. Since the synonymy in Appendix 2 is intended to help the reader “‘to identify descriptions in old books” it should surely have included the specific name montana for O. limbosperma, the name used in many older fern books. However, these are all minor criticisms; the text is sound and the book well worth its modest price for local botanists. R. M. PAYNE A Flora of Wensleydale. A centennial review of the plants of the Dale. D. Millward. Pp. 180, with 2 maps. Yoredale Natural History Society. 1988. Price £3 (ISBN 0-9514287-0-5). This neat, compact flora marks the centenary of the publication in 1888 of John Percival’s original flora in The Naturalist. The meat of the flora comprises a 118 page listing of the species present, with simple qualitative assessments of the present status of all species of flowering plant and fern according to the categories ‘abundant’, ‘common’, ‘occasional’, ‘rare’, ‘very rare’ and ‘no recent record’ (i.e. since 1985). In the assessment, individual sites are rarely mentioned, grid square references are not listed, nor are the sources of modern records. One of the better entries is for Primula farinosa (Bird’s-eye Primrose) — “Frequency — occasional. In damp pastures and meadows on limestone or glacial till and in calcareous flushes, from 150m at the river in Aysgarth to 570m at Jeffrey Pot. Percival recorded the species as “abundant in many places in the valley”’ but it is now very rare below 250m due to draining, as at Rabbit Warren, Carperby”. Such an entry enables a broadbrush understanding of the ‘then and now’ status of selected species. The flora would have been greatly enhanced by the presentation of a summary table of those species whose distributions have been drastically reduced, extinctions, and extension of ranges due to more detailed recording. A pleasing feature is the inclusion of geological and geomorphological maps of the Dale and the appropriate cross-referencing of species distributions related, in particular, to glacial drift and limestone. For example, the entry for Coeloglossum viride (Frog Orchid) is informative in ecological terms — “most sites are on glacial till but it has been recorded directly over the limestone, all are below 250m”. The main body of text is followed by a succinct series of essays under the general heading of ‘Environmental Setting’, comprising ‘geological foundation’, ‘climate’, ‘soil’, ‘vegetation’, ‘geomor- phology and vegetation’, ‘changes over the century’ and ‘past and present land use and conservation’. The nub of what could have been an important contribution to an understanding of changes in and conservation of the flora, inherent in the titles of the latter two sections, is covered in less than three pages of text. Nevertheless, the book provides another timely reminder that our flora is continually under threat and that only by bold initiatives of synthesis such as this can we begin to make realistic assessments of what has happened, is happening and what needs to be done in the future. D. W. SHIMWELL Developmental biology of fern gametophytes. V. Raghavan. Pp. vit+361. Cambridge University Press, Cambridge. 1989. Price £40 (ISBN 0-521—33022-X). This is a well written, thoroughly researched book. The title was wisely chosen, as it is a good indication of the content. The emphasis reflects the author’s recent interest in the area of spore germination, but morphogenesis, vegetative growth and regeneration, sexuality, and departures from conventional life cycles are well covered. The author focuses on the physiological, cytological and biochemical background of each topic under discussion. This makes for a well rounded text, but one which seldom removes fern gametophytes from the laboratory (and which may therefore have limited appeal for many B.S.B.I. members). “‘Natural reproductive biology of ferns” covers only a page and a half, and the ecology of gametophytes gets only an occasional mention (but the author did not set out to write for the field biologist). I thought this a pity, considering, for example, the BOOK REVIEWS 233 recent studies that show at least some gametophytes to be considerably more resistant to environmental extremes (and indeed, more widespread) than their sporophytes. The book does help, however, to explode the still widely accepted myth that ferns are self-fertilising, often highly polyploid plants. The author’s enthusiasm for his subject carries the reader fairly painlessly through the more complex biochemical processes crucial to gametophyte functioning, and makes the morphological feats gametophytes achieve sound fascinating. There is a wealth of up-to-date, well-summarised data on each topic covered, and although not lavishly illustrated, the book does make points clearly and simply. There are very few typographical errors, and my only regret as far as production is concerned is the lack of a glossary. If you don’t already know what a periplasmodium is, you will not find out from this volume, but if you want a comprehensive set of references to work on the developmental biology of fern gametophytes covering the last forty years, this is the book for you. I very much enjoyed reading this book, and will find it an invaluable resource for the forseeable future. I can only hope that it finds a market large enough to repay the author’s considerable efforts outside those fortunate enough to be asked to review it. E. SHEFFIELD Colour identification guide to the grasses, sedges, rushes and ferns of the British Isles and north- western Europe. F. Rose. Pp. 239, with 62 colour plates and numerous text figures. Viking, London. 1989. Price £35 (ISBN 0-670-80688-9). This book purports to complement the same author’s The wild flower key (1981), covering the four groups that that book omitted (Gramineae, Cyperaceae, Juncaceae and vascular cryptogams), but it is much more expansive in both text and illustrations, and has a much larger, hardback only, format (24-5 x 19-5 cm). It is very rare nowadays to find a field guide by a good naturalist that does anything like justice to the author’s knowledge and experience, and Viking are to be congratulated on giving _ Francis Rose something like enough space to express himself adequately. There is of course a price to pay for this, and although I think £35 is too much in comparison with some other recent plant guides, it is a good buy if you can afford it and in many respects it is the best book of its kind covering _ these groups. The emphasis of coverage is on the British Isles, but extends on the continent from the Loire valley to Denmark and south-east to the Jura. The introductory material includes very helpful accounts of the morphology of the groups, and of the features important for identification. The keys are generally excellent and clear, being both user-friendly and, equally importantly, plant-friendly, avoiding the use of underground parts (although the author forgets his exemplary principles in one place in the Carex key and uses rhizome-width as one of the characters). In addition to the main keys there are seven keys to the commoner grasses of various habitats using vegetative characters; this is one of the best sections in the book, and is the sort of item that in the days of reader-friendly publishing would have been additionally printed as a booklet and tucked into a pocket on the back cover so it could be used separately in the field. The 62 colour plates illustrate 350 of the 420 species in the book. They face the corresponding pages of description and are by Laura Mason, Claire Dalby and R. B. Davis (it is not stated who did which). The six plates of non-Carex Cyperaceae, containing numerous detailed drawings as well as the habit paintings, are in a class of their own and quite exceptionally informative and attractive. The grass and rush plates are good, and it is not easy to represent these plants in water-colour. The Carex plates are rather less satisfactory, and it is not always easy to guess which species is being depicted without checking the names. There are some annoying inconsistencies, for example in the C. muricata group where the short ligule is drawn for three of the taxa but the diagnostic long ligule of C. spicata is omitted (though mentioned in the text), and in C. distans where the leaf-apex is shown in detail but not that of C. hostiana which is again diagnostic in this group. There are a few _ Surprising mistakes in these plates, such as C. pilulifera being shown with a sheath to its lowest bract, C. bigelowii being shown with a strongly veined utricle, and C. demissa being shown with 234 BOOK REVIEWS proportionally shorter leaves than C. lepidocarpa. The fern plates are variably successful, those of the filmy ferns being rather misleading, but others, such as those of the clubmosses and horsetails, are good. The descriptions in all the groups are full of useful and diagnostic detail. My chief doubt about unreservedly recommending the book to all B.S.B.I. members, though, lies in the treatment of the rarer and more critical species. Altogether 70 species described are left out of the colour plates, although details of some are shown in drawings in the text. The Cyperaceae are very thoroughly covered in both text and plates, even such very rare plants as Carex microglochin and C. muricata subsp. muricata being treated in full. But in the vascular cryptogams Isoetes hystrix for example, is unillustrated, J. echinospora has only its spores depicted, and such widespread and distinctive- looking ferns as-Polypodium interjectum and P. cambricum are unillustrated, as well as unkeyed. In Juncus the segregates of J. bufonius are unillustrated, although keyed. The grasses are similarly poorly served with such species as Festuca vivipara, Poa angustifolia and P. subcaerulea left out of the plates. This book’s direct competitor, R. Fitter, A. Fitter & A. Farrer, Collins guide to the grasses, sedges, rushes and ferns of Britain and northern Europe (1984), covers critical taxa such as these much more fully, serves a much wider geographical area, is pocket-sized, and now sells for less than a quarter the price, but in other respects does not reach quite the same standard. The book under review is generally so good that it has a lot to offer even the knowledgeable botanist, although he or she will be just the person most frustrated by its failure to help adequately with the more critical species. Nevertheless, buy it for the diagnostic detail in the descriptions and keys, and for | plates 24-29. A. O. CHATER Norges ville blomster — fra bier og blomster til fro og frukt. [Norway’s wild flowers — from bees and flowers to seeds and fruit.] Edited by L. Ryvarde, Oslo. R. Berg & K. Fegri, pp. 240. H. Aschehoug & Co. (W. Nygaard), Oslo. 1989. Price not stated (ISBN 82-03—-15976-1). This book is the first in a planned series on Norwegian plant life edited by Leif Ryvarden. The aim of this volume is to provide the general Norwegian reader with an introduction to the pollination and dispersal biology of flowering plants. Our honorary member Knut Fegri, Emeritus Professor of Botany at the University of Bergen, has written the pollination section (139 pp.), whereas Rolf Berg, Professor of Botany at the University of Oslo has prepared the dispersal section (98 pp.). Both sections are profusely illustrated with many superb photographs contributed by at least 29 photographers and collated by Even Tj@rve. Although primarily about the Norwegian flora, examples illustrating particular pollination or dispersal mechanisms are discussed with reference to tropical and desert plants. This is an excellent book, clearly and concisely written but unfortunately for many B.S.B.I. members only available in Norwegian. The photographs are superb, and many must have been specially commissioned to illustrate particular biological features, especially in the dispersal section. I know of no other book that illustrates so effectively and so elegantly the diversity and beauty of different pollination and dispersal mechanisms. Michael Proctor and Peter Yeo’s New Naturalist The Pollination of Flowers (1973) covers pollination biology in more detail than here, but there is no comparable and readable book on dispersal biology. It is a pity that Latin plant names are used so infrequently in the Norwegian text, as it is tedious to have to use a Norwegian flora to discover what plant is being discussed or illustrated. It would be wonderful if this book were translated into English. It would then be the first — semipopular, up-to-date, and readable book on these important, fascinating, but often ignored — topics in plant biology. Congratulations are due to Rolf Berg, Knut Fegri, Leif Ryvarden, and Even Tjgrve for producing such a fine book. H. J. B. Birks BOOK REVIEWS 235 Ted Ellis: the people’s naturalist. E. Stone. Pp. 157. Jarrold Colour Publications, Norwich. 1988. Price £3-95 (ISBN 0-7117-0436-8). This little book is a biography of Ted Ellis (E. A. Ellis), the famous Norfolk naturalist. It is good to see such an excellent biography of Ted. I have known him personally for nearly 50 years, and my wife and I have many happy memories of staying with him and his family many years ago. All those who knew him remember him as a delightful man of great gentleness, simplicity and humour. Though the author never met him, Ted’s personality comes through quite remarkably in this book. Eugene Stone based his work largely on interviews with Ted’s wife Phyllis, and many others who knew him well. The result is a remarkably authentic biography. Ted was a unique figure among British naturalists. He developed (and acted upon) the concept of conservation as a youth, at a time when few had even thought about the subject. He put his ideas into practice over many years, particularly at his Wheatfen terrain. Wheatfen has now been bought as a permanent Nature Reserve, and is managed by the Norfolk Trust. His main contributions to serious scientific research come through very well in this book. They were his (often pioneering) studies of many of the smaller fungi, particularly those that cause diseases in plants or in invertebrates. However, we are told clearly that Ted was a most comprehensive naturalist, with a deep and extensive knowledge of nearly all groups of plants and animals, and of their ecology, in a way that has become very rare in these days of specialisation. Such breadth and depth will probably not be seen again in one person. Eugene Stone also tells us much of his life-long contributions to Norwich Museum, in particular the creation of the wonderful dioramas of Norfolk habitats. Such vivid portrayals have never been equalled elsewhere, and have been models which many others have tried to imitate with varied success. He tells also what a wonderful teacher Ted was. He was to my knowledge particularly brilliant with children, and was an excellent radio and television broadcas- ter. He also makes it clear how essential Phyllis was in all of Ted’s life and work; as he says, ‘‘any story of Ted’s life is also her story’’. I have only spotted one small error, in the caption to the lower photograph on p. 77: the man on the right is A. E. (not E. A.) Ellis. This charming little book is a splendid read, and I would recommend all naturalists to obtain it; to read it is an inspiration. F. ROSE The European garden flora. A manual for the identification of plants cultivated in Europe, both out- of-doors and under glass. Volume III, Dicotyledons (Part 1). Edited by S. M. Walters, J. C. M. Alexander et al. Cambridge University Press, Cambridge. 1989. Pp xv+474. Price £65 (ISBN 0- 521-36171-0). There are many books on the horticultural aspects of garden plants, whereas their taxonomy is a somewhat neglected field. The European garden flora, which is planned to comprise six volumes, is intended to be an identification manual for plants cultivated outdoors as well as under glass, and thus fills a long-felt gap in the literature. Volume III deals with 49 families of dicotyledonous angiosperms (Casuarinaceae to Aristolochia- ceae). It is a surprise to see how many families contribute to the garden flora of Europe (there are for example members of the Olacaceae, Didiereaceae and Canellaceae). According to the main aim of the flora, much effort has been devoted to the contstruction of keys for families, genera and species. These are, indeed, well elaborated. There are — as is usual in floras — strictly dichotomous formal keys which will guide the user reliably to the name of a given taxon. In some medium-sized genera also so-called informal keys are presented, being a guide to some species of the genus on the basis of characters more easily recognised. In the Cactaceae also a key to groups based only on vegetative characters is given. In the genus Berberis, some special comments are made for using the key because the species are very difficult to identify. There is a key to the double- flowered variants of the Caryophyllaceae. 236 BOOK REVIEWS The most voluminous families in this volume are the Cactaceae followed by Ranunculaceae and Aizoaceae. The largest genus is Berberis with 103 species, followed by Salix with 79, Mammillaria with 70, and Quercus with 57 species respectively. The arrangement of the genera and species is not alphabetical, but is in taxonomic order so that the descriptions of closely related taxa can be easily compared. In the full-length descriptions of families, genera and species, a terminology is used which will be easily understood also by the layman (there is also a glossary for the inevitable technical terms). Horticultural information is given in a very brief manner only. Literature is presented for each family, and genus and illustrations are cited for each species. Synonyms are given only to a limited extent. English as well as any other vernacular names are lacking. There are remarks on geographical distribution, flowering time, hardiness, infraspecific taxa and hybrids. Except for four diagrams illustrating terms defined in the glossary, the 42 line drawings (or silhouettes) show diagnostic details of critical genera. Certainly the nine figures of Salix leaves, five figures of Quercus leaves, four figures of Ficus leaves and the inflorescence types in Berberis, etc., are extremely useful for determinations within these difficult groups, but this results in a very unequal distribution of figures throughout the text, many families not being represented in figures at all. This seems to be a shortcoming of the book. The Flora, modelled in outer appearance, size and structure after Flora Europaea, is well produced. There are only some minor misprints in the citations of German journals. Because the selection of species included in this Flora has been very extensive, the book will show its usefulness also outside of Europe. The editors and authors are to be congratulated for tackling such an enormous and difficult task as this Flora. The European garden flora, being a major event in the taxonomy of garden plants, will be extremely useful for laymen as well as for professional botanists, and will furthermore act as a stimulus in this field. The following volumes are eagerly awaited, and it is to be hoped that they will come out in the near future. J. SCHULTZE-MOTEL Morphology of flowers and inflorescences. F. Weberling, translated by R. J. Pankhurst. Pp. xx + 405, with 193 text-figures. Cambridge University Press, Cambridge. 1989. Price £55 (ISBN 0-521- 251346). The comparative study of plant morphology has reached its most elaborate form in Germany, where the fundamental researches of Hofmeister in the middle of last century laid the foundations for the work of Goebel and his pupil Troll. The language used by these workers has always been ‘difficult’, even for native German speakers; and so it is not surprising that their works have been slow in gaining acceptance outside the German-speaking world. This is a pity, because Troll succeeded in producing an extremely detailed and logically developed system of terminology that, correctly used, could much improve the accuracy and comparability of taxonomic descriptions in Floras and monographs. Now one of Troll’s students, Focko Weberling, has condensed and updated the Trollian concepts of floral and inflorescence comparative morphology in a readable and convenient form. Richard Pankhurst’s English version is not merely a translation. The originally extensive bibliography has been considerably enlarged, mostly with recent references, and Professors Weberling and Miller- Doblies have compiled a glossary of morphological terms, because their meaning in German sometimes differs from that in English. The glossary is indeed a help in understanding the frequently complex arguments, although some definitions, e.g. of acrotonal and basitonal, are inadequate. Taken on its own terms, this is a marvellous book. It provides, for example, a guide to the comparative study of inflorescences by Troll and his school, when nearly all taxonomists (with the notable exceptions of Laurie Johnston and Barbara Briggs in Sydney) are still at the stage of: open versus closed, cyme versus raceme, thyrse versus panicle. Its own terms, however, rule out any theory based on floral vasculature. Weberling would appear to subscribe to Goebel’s doctrine that vascular bundles arise where they are needed, so there is no question of their being conservative and therefore indicative of ancestral states. He follows the classical Euanthium theory, where the main floral organs are regarded as modified leaves — sterile BOOK REVIEWS 237 (sepals, petals) or fertile (stamens, carpels), and dismisses what he terms Pseudanthium theories (e.g. the Telome, Gonophyll and Anthocorm theories of Zimmerman, Melville and Meeuse, respectively). The Telome theory is not even mentioned, and Melville’s work merely “strikes [him] as rather complicated”. Weberling does, however, invoke vascular structure where it suits his argument, e.g. in the interpretation of the inflorescence of some Cordia species, but not in the interpretation of the inferior ovary. The short discussion of heterostyly makes no mention of the work of Bob Ornduff, and the term ‘aril’ does not appear at all. In short, this book provides an entry to, and a summary of, the work of the Trollian school of morphology and its modern manifestations, with regard to the flower and inflorescence. As such, it is an essential source for morphologists and for Flora writers who try to make their descriptions really comparative. The translation reads very well, in general, although not all the ‘Germanicisms’ have been eliminated. The price, however, may prevent its becoming a university text-book, which it should be. N. K. B. Rosson Watsonia, 18, 238-246 (1990) Obituaries EVELYN MARY BOOTH (1897—1988) The acknowledged contribution of amateurs to British and Irish botany has particular relevance to the life of Evelyn Booth. Coming from an Irish Ascendancy background, she was born in Co. Wicklow, later settling at Bunclody, where Co. Carlow borders Co. Wexford, and here she spent most of her life. Before World War II her interests were centered round horses and pursuits like fishing, though she was always a gardener, and this made it possible for her to extend her interest to wild plants when chance changed her life in the early years of the war. She was recalled from nursing in Essex to care for her ailing mother in Ireland, and at this time met Edith Rawlins, a dedicated and determined member of the Wild Flower Society, who was tutoring families in Evelyn’s home county of Carlow, and in neighbouring Co. Wexford. They began to botanise together, Evelyn’s considerable energies were engaged, and as her life-long friend and neighbour Daphne Hall-Dare has said, “thereafter, wherever she went, her head was down in search of some treasure’. From this time, in the 1940s, to her death in December 1988, Evelyn Booth never stopped contributing to botany. The early work during the war, hunting and ticking “‘on slag heaps, bogs and mountains”’, bicycling to see rarities with her cousin Daisy Barton (also a botanist) and with Miss Rawlins, developed into a most thorough piece of county recording and research. This resulted in a major contribution to the B.S.B.I. mapping scheme in the 1960s, when she covered the whole of Counties Carlow and Wexford, and culminated in the useful and approachable Flora of Co. Carlow which was published in 1979. This has the distinction of being the first Irish county Flora to be written by a woman. The final stages of its production were helped by Maura Scannell, herself a major contributor to modern Irish floristic knowledge, notably by her co-authorship of The flora of Connemara and the Burren (1983) with Professor D. A. Webb. Evelyn Booth also published notes on finds from various parts of Ireland in the Jrish Naturalists Journal. A complete bibliography of her work by Dr E. Charles Nelson (himself a good friend and gardening colleague of Evelyn) will appear in Glasra 1 (n.s.), published by the National Botanical Gardens, Glasnevin. It is difficult to single out any one quality which made Evelyn Booth the remarkable character she was. Her place in the lives of countless friends, botanical and otherwise, could never be neatly labelled. A visit to Lucy’s Wood, her house above Bunclody and the River Slaney, always involved a whole range of pleasures and interests: her most delightful garden (that of a true plantswoman), her sometimes wonderfully salty humour, tranquil rooms full of books and flowers where she filled visitors with good food and inspiration. A typical memory comes from David Webb, of a botanical meeting for serious work on publications. Of course lunch was provided — an extremely good salmon. Appreciative comments only brought the most casual reply from Evelyn, “Oh I thought you might enjoy one, so I got up early and caught it’’. My own long-term memories centre on her courtesy. Even when I was but a child ‘woof’, and later as a very shy amateur, she always treated me as if there was no doubt that I was a real botanist too. I owe her much of the confidence that has caused me to remain one, and I am sure that others share this debt. More recently, during her last years, I have had the benefit of her indomitable spirit and humour. Suffering a broken hip, uncertain eyesight, and what she complained of as “dicey puff’, there was never any withdrawal from plants or friends. How inspiring it was to have a lady of 89 commenting in very modern language on having ripped an arm on barbed wire while making one of her regular seed collections for the National Botanic Garden, and at the same time showing quite youthful excitement about a proposed Hammarbya hunt, and berating the insensitivity of her garden help, whom she called ‘‘the veg-brutal’’. When I was asked to write a tribute to Evelyn, only affection and my own Co. Carlow background seemed to justify my involvement. But during the year since her death, a personal jigsaw has been completed. The pieces in the puzzle have been Miss Rawlins, fanning Evelyn’s enthusiasm for wild plants, and coincidentally teaching me Latin names as I learned to talk at the house in Borris where she was tutoring my aunt; the counties of Carlow and Wexford which involved so much of Evelyn’s 238 OBITUARIES 239 life, and gave me so much of my early experience of botany; and Evelyn herself, who kept me in touch with Irish plantlife at times when I was far away. In 1989 I became, together with John Akeroyd, B.S.B.I. recorder for Co. Wexford, and the chance this gives me to use and build on Evelyn’s work seems to make sense of the gratitude I feel for her existence as both friend and colleague. The power of the amateur tradition, which Evelyn Booth with her lively dedication embodied so finely, must never be underestimated. The continuity, the intricate web of communica- tion and encouragement, can often make valuable sense and records out of apparent trivia. Some two years before her death, when I was gathering biographical information on Miss Rawlins, Evelyn sent me the prophetic present of a tatty old notebook which turned out to be Miss Rawlins’ checklist for Co. Wexford. This is proving a rare and valuable addition to the county records, and I will always smile when I think of Evelyn’s prescience, and remember the utterly botanical phrase she used when she wrote to say she wanted me to have the notebook “because no-one will know what it is when I’m over’. R. FirzGERALD JOHN CAMPBELL GARDINER (1905—1989) The death on 4 September 1989 of Jack Gardiner, as he was known with much affection to many of the older members of the Botanical Society of the British Isles, breaks a link with the past in the development of the Society. Although the Society was formed in the middle years of the 19th century, it almost ceased to exist during the period of World War II. A reconstituted society had been formed shortly after the end of the war and was much in need of a period of stability. When a minor crisis arose in 1958 with the resignation of the Honorary Treasurer, a review was made of the then limited membership of the Society and a decision was made to approach Mr Gardiner, of whom nothing more was known to us other than that he had joined the Society in 1949, was a chartered accountant and lived in London. It was left to me as Honorary General Secretary at the time to write to Mr Gardiner, which I did, receiving a reply from Mrs Gardiner saying he was abroad but she thought that he would consider our invitation favourably. That he did so was to be very much to the benefit of the Society and I have never ceased to wonder how fortuitous our choice had been. Jack Gardiner was born on 20 November 1905 at Shahjahpur in India where his father, following several generations of military service by the Gardiner family, was serving with the Royal Engineers. His father was killed on active service in 1914, during World War I, when Jack was nine years old. As a result Jack was educated at Wellington College, where he developed an interest in science, and for a while he oscillated between following an academic scientific career and entering the business world as an accountant. After qualifying as a Chartered Accountant in 1929, and gaining one of the Institute’s prizes in the process, he became a science student at a University Extension Course. There he met his future wife, Marjorie (Wendy) Massey, whom he married in 1933. It was, however, a common love of music rather than test tubes that had brought them together. The responsibility of marriage made him settle for the business world and Jack became a partner in a City firm of accountants but was seconded to the Ministry of Food in the war years. In 1959, a year after becoming Honorary Treasurer of the B.S.B.I., he was persuaded by Sir Charles Clore to accept the post of executive director of Sears Holdings Limited, an office he was to hold for the following 13 years. By a series of takeovers the concern gained control of much of the British retail trade, including that of footwear, cars and Selfridges. His involvement in the motor industry was somewhat to his own amusement as he could not drive a car and had no desire to do so. He often said that in his youth he preferred to ride horses. His period of office as Honorary Treasurer of the B.S.B.I. coincided with the closing stages of the Maps Scheme and the publication of the Atlas of the British Flora in 1962, which was to prove to be a turning point in the study of natural history. The attentions of the Society were then moving towards problems of plant conservation, causing Jack to play no small part in the fight to try to save Upper Teesdale. In all this time, although living in London, Jack had lost none of an inborn love of the English countryside and with the ties of business relaxed for one day in the week he escaped to the 240 OBITUARIES rural areas near to London every Sunday. This could be the more enjoyable with some object in view and at around the time of the end of the war in 1946 he found this in the study of the flora. In 1957 the Surrey Flora Committee was formed, its object being to ensure that the Maps Scheme records for that county would be as complete as possible. With the assistance of public transport it was possible to reach many parts of Surrey in the morning and return by another route within one day. He lost no time in assisting the Committee, to which he was co-opted in 1962. By then he had shown himself to be a more than competent field botanist, with his own neatly compiled crib and documented field notes. J. E. Lousley’s Flora of Surrey was published in 1976, but long before this Jack’s advice was being sought in a wider field and his plant studies became broader by extending them to bryophytes which could be studied more fully in the field in the winter months in which Jack felt the same need to escape to the countryside. Above all his interest in Surrey had introduced him to bryologists of the calibre of W. R. Sherrin of the South London Botanical Institute and E. C. Wallace. He joined the British Bryological Society in 1964 and was elected to sit on its Council from 1972 to 1975. His only published work was ‘A Bryophyte flora of Surrey’, in J. Bryol. 11: 747-841 (1981), which has been described as a model for other local Flora writers. In addition to spending many happy days in the countryside it had entailed much time searching herbaria. With this task completed he turned his attention to Middlesex. The B.S.B.I., the Surrey Flora Committee and the British Bryological Society were all very much in the control of amateur field workers but another society, the Linnean Society of London, which had royal patronage, rooms of its own, a paid staff, a valuable library and unique collections, was almost entirely in the hands of professional biologists — who nevertheless at times could need advice on other matters. Jack joined the Linnean Society in 1961, being elected to serve on its Council for two periods from 1966 to 1970 and again from 1973 to 1979, a Vice-President 1975 to 1979 and as Honorary Treasurer in that same period. In the year of his death he was the recipient of the Bloomer Award which is given from time to time by the Linnean Society to “‘an amateur naturalist who has made an important contribution to biological knowledge’. More worthy recipients of the award have been few. Otherwise he had given the Society much valuable advice on its publication policy. In the meantime, having been Honorary Treasurer of the B.S.B.I. for 12 years, he relinquished that office in 1970. He had played no small part in giving some much needed stability to the Society. His last important service to the Society was in helping to persuade Mr M. Walpole, another accountant, to follow him in office. The joint custody for 30 years by two such devoted officers has meant much to the Society. Jack continued to be as active as he had always been until 1983 when the illness of his wife Wendy, who had been his companion for so many years on his excursions into the London countryside, brought a change. He felt that his place was by her side assisting in any way he could her recovery, until his own final illness overtook him. He was essentially a modest man, being a ready listener with a keen sense of humour. His advice, so often sought by others, was always given with some hesitancy but invariably found to be sound. We shall not see his like again. Notwithstanding his very full life, Jack was essentially a family man and is survived by his widow, two daughters and a son. J. G. Dony CHRISTOPHER C. HAWORTH 1934—1989 Chris Haworth died on 2 December, 1989. He was born and bred in Lancashire, and although he lived long in West Cumbria, and put down many roots there, it was never difficult to recognise his Lancastrian origins. After National Service in the R.A.F., he graduated in Natural Sciences from Queens’ College, Cambridge and worked for five years as an industrial chemist with British Petroleum. It was not long, however, before his natural bent drew him into the world where he was to make his indelible mark. Although he had graduated as a biochemist, he became a teacher of biology, first, for three years, in Essex, and then, for some 20 years, at Whitehaven Grammar School in Cumberland. He was an inspiring teacher of young biologists, many of whom now grace OBITUARIES 24] the higher ranks of education, and who would happily acknowledge the debt that they owe to his influence in their formative years. What made him distinctive as a teacher was the rare combination of, on the one hand, quiet authority in his own subject, and on the other, an intellectual curiosity and incurable desire to stimulate concern about the widest range of cultural, political, and intellectual matters. The liveliness and all-embracing nature of his interests, indeed, was as much in evidence in the influence he had on his colleagues, as in that on his pupils. Whether it was in one of the various unorthodox musical ensembles in which he played the recorder, or discussing the most recent novel that he had discovered, or explaining the potential for computers in education to sceptical colleagues, it would be difficult to exaggerate the beneficient effect that he had on those who came in contact with him, and consequently, the sense of loss felt by so many at his untimely death. He was the first person that I knew to express a deeply-held philosophical belief in the frailty of the earth’s environment, and man’s obligation to protect it. There were very few others back in the sixties who shared his doubts about the propriety of leaving radioactive waste as an inheritance for future generations. It was wholly in keeping that he should have been one of the two founding members of the local branch of Friends of the Earth, which he always saw very much more as a positive force in the defence of the environment, rather than just a vehicle of protest. When, however, he felt protest was called for, as at the time of the Windscale Inquiry, this essentially quiet man was quite extraordinarily persistent and single-minded in the enormous amount of effort and skill he put into organizing the local case against the planned development. As a field botanist, he was an exemplar. He combined encyclopaedic knowledge of most branches of the British flora, a comprehensive first-hand knowledge of a very large area of the Lake District, a meticulous care in the ordering of his herbarium and records, and the skill to execute a programme of garden cultivation in pursuit of further research. Yet, he combined these skills with such modesty and quietness of demeanour, that there must have been many who were quite unaware of his distinction as a botanist, and few indeed were those who penetrated to the limits of his knowledge. It is difficult to convey the sense of excitement and fun that attended a day in the field with him. It was uncanny how often he would say “‘I rather feel that there is something interesting around here’’, to be proved right only a few minutes later. He was pleased, and would have been justified in feeling proud (though that would have been out of character), at the number of times that he had refound plants from 19th century or earlier records. He was, for many years, one of the leading members of the Flora of Cumbria project, in which he was responsible for supervising the recording over a large tract of west Cumbria, much of which, of course, he had recorded in person. It was through his work on the Flora of Cumbria project that he first became interested in the genus Taraxacum. Having, in the early days, sent some specimens for identification, he became determined to master this genus in his area. Over the next eight years, he proceeded, in the words of John Richards, to “transform the history of British Taraxacology,” with his unique mixture of ability, energy, enthusiasm, organization and scholarship. He would always, however, insist on emphasising how much, like all taraxacologists, he owed to John’s pioneering work in establishing the foundations upon which he worked. He rapidly built up a long list of correspondents, initiated a British Taraxacum newsletter, and built up a computer-generated Taraxacum database which gave rise to a long series of frequently updated and extensively annotated British check-lists, available to all correspondents. It was characteristic of Chris, as a taraxacologist, to take nothing for granted. He believed in no species until he had thoroughly checked it out for himself! This involved seeing the types of almost all the British species, which in itself led to an extended programme of lectotypification, the fruits of which are at present in press. It also led to extensive travels during which he visited the type localities of many British species, in Ireland, northern Scotland, Orkney and southern England. Like all the best taraxacologists, Chris was an enthusiastic cultivator, who maintained a large and fascinating series of comparative cultivations in pots. He remained entirely sceptical about any species which he had not checked against the type, and which he had not grown alongside its relatives. Chris’s real enthusiasm was for the special dandelions of western Britain, the Celtica and Naevosa. He discovered, or described, many splendid new taxa; his taxonomic reassessments, based on sound method as well as on inspirational judgement, are far too many to list in full, but he made it 242 OBITUARIES possible to foresee the future publication of an account of the genus in Britain. Although, sadly, he did not live to see it published, he had done enough for us now to envisage the successful culmination of his work in the shape of a B.S.B.I. Handbook of British Taraxacum, which his colleagues hope to see concluded, in what will surely be a fitting memorial to Chris’s work. Three years ago, he was appointed to a new post, as Head of Sixth Form at Cockermouth School. Unfortunately, he learnt at the same time of the illness that was to prove fatal, and he was never fit enough to take up the full burden of his new office. Even so, few realised how ill he was and he characteristically fought his illness with inspiriting courage to the very end. He carried on an international correspondence, he continued to act as referee for Taraxacum, and he continued, though less and less strenuously, to botanise in the field. He never complained, and very few were allowed to realise how near the end was. Even more important to Chris than his botany was his home and family, of whom he was intensely proud; his hospitality and that of his wife was renowned, and there was always a welcome for family, friends, neighbours or colleagues. The support of his family, and in particular of his wife, Bertha, in his last illness were incalculable. Happily, he acknowledged something of the debt that he knew he owed her by the naming of the Cumbrian Dandelion, Taraxacum berthae. One of his last full days out was last July, at Wasdale Head, for a field meeting with many friends on the Flora of Cumbria project. It was a heavenly Lakeland day, cloudless and windless; the surface of Wastwater was like glass. While the more energetic left for the high fells, Chris, by now much weakened, was perforce confined to the lowland tetrad at Wasdale Head itself. When I pointed out the list of previous distinguished recorders on the card, and suggested that, in what must in any case be one of the most visited tetrads in the county, we weren’t likely to find very many new records, Chris cheerfully expressed greater optimism, and within a hundred yards, said, ‘“‘What about that, for instance?” Needless to say, it was a new record, and, by the end of the day, we (more usually, he) had found a further 20 records for the square. It was a fine day by which to remember him. A. DUDMAN C. C. HAWORTH’S CONTRIBUTION TO THE FLORA OF CUMBRIA Sixteen years ago, Chris was the first to respond to my appeal for recorders for the Flora of Cumbria project. So began a close and productive friendship during which he accumulated, almost single- handed, the bulk of our west Cumberland database. It was a friendship nourished by frequent letters telling of new finds and rediscoveries and enlivened by characteristic flashes of his dry wit, but a friendship overshadowed in more recent years by the darkening cloud of his illness. As the flora progressed, and preliminary accounts started to circulate, I particularly appreciated his restraining hand in my more shaky generalizations, verbal infelicities and factual errors. During his last few months, family accounts were dispatched post-haste off the word-processor, and seldom was he more than a few days in replying; but his most impressive achievement was the revised account of Cumbrian Taraxacum which, with Bertha’s help, he completed and gave me only six weeks before his death. He made the most of his last summer in the field, and we were all amazed at his courage and single-mindedness in travelling to join our August bramble foray in the far east of the county. It was typical of him that, in rapidly declining health, and with the dandelion account to finish, he should nevertheless want to get to grips with Cumbrian brambles. His Taraxacum herbarium and part of his general herbarium is now with Andrew Dudman, the more important Cumbrian specimens from the latter being in the Lancaster University herbarium (LANC). G. HALLIDAY ARCHIBALD GRAHAM KENNETH (1915—1989) Archie Kenneth of Stronachullin, near Ardrishaig, Argyll who died on 27 July 1989 at the age of 74, had been a member of the B.S.B.I. since 1957. Field botany was a consuming interest for him and OBITUARIES 243 circumstances allowed him to devote a considerable amount of time to its pursuit. For many years his main energies were directed towards the botanical exploration of the districts of Knapdale and Kintyre, comprising v.c. 101, for which area he was appointed vice-county Recorder in 1961. This work led first of all to the publication in 1964 of the Flora of Danna, an account of the vegetation of a small island in Knapdale in which he had taken a particular interest, and this was followed by a supplement in 1971. In 1979 the Flora of Kintyre was published, compiled in collaboration with Miss M. H. Cunningham of Campbeltown. Continuing field work in the vice-county soon created the need for a supplement to the latter Flora and this Archie himself published in 1985 as Additions to the flora of Kintyre. Archie had a keen eye for a new or unusual plant and this facility led to the discovery of many a rarity which would have escaped the notice of less observant botanists. The discovery of Lathyrus palustris and Cirsium dissectum in Knapdale, and the finding of a fine colony of Diphasiastrum complanatum subsp. issleri in West Sutherland are just a few examples of his talent for turning up things of more than ordinary interest. He was also an enthusiastic and competent bryologist and a very complete list of the bryophytes of v.c. 101 was included in the Flora of Kintyre. Archie developed a special interest in critical plant genera, particularly Hieracium, Taraxacum and Rubus, not merely from the point of view of local Flora compilation but from a genuine and absorbing interest in such groups. This brought him into contact and correspondence with the specialists — Peter Sell and Cyril West for hawkweeds, and Eric Edees for the brambles, all of whom at one time or another enjoyed the hospitality generously extended at Stronachullin by Archie and his wife Janet. On more than one occasion Cyril West accompanied Archie on hawkweed-hunting trips to north-western Scotland, an experience much appreciated by Cyril in his later years. Of all the critical plant groups, that which most caught Archie’s attention was the section Alpina of Hieracium, especially its members inhabiting the rather species-poor hills of Wester Ross and Sutherland. In search of these attractive hawkweeds he would tramp many miles in inhospitable country, perhaps to visit a hill previously unknown botanically, and to find, if lucky, one or more species of this interesting group. This dedication resulted in a greatly enhanced knowledge of the alpine hawkweeds of the north-west and the discovery of several new and as yet undescribed species. It is fitting that one of these is to be named in his memory. Resulting from the discovery of some puzzling colonies in his home area Archie became very interested in the marsh orchids (Dactylorhiza) and after studying populations in western and north- western Scotland he became involved, as joint author with David Tennant and others, in the publication of papers on D. incarnata subsp. cruenta, D. lapponica and the enigmatic D. francis- drucei which was shown to be referable to D. traunsteineri. Archie was a rather rare attender at organized botanical meetings, although he rarely missed the annual B.S.B.I. Exhibition Meetings held in Glasgow or Edinburgh, where he obviously enjoyed the opportunity to foregather and exchange news with fellow botanists. In the field he preferred his own company or that of one or two companions. During an excursion it was not unusual for him to go off on his own in order to carry out some private investigation or visit a particular spot he had in mind and which the slow (by his standards) rate of progress of the party would otherwise have prevented. Those who have accompanied him in the hills will readily testify to his speed and agility. His stride was deceptively unhurried, but companions were often left trailing far behind. His dress was unconventional by modern standards and consisted of an old pair of trousers, tweed jacket, open-neck shirt, knitted woollen bonnet and, if it looked like rain, an old gabardine raincoat. He never wore climbing boots, preferring a strong pair of brogues or perhaps wellingtons. He travelled light on the hills with no encumbrances such as camera or binoculars, and a small satchel sufficed to carry his lunch and any specimens collected. He was never without a stout hazel crook which he found indispensable in assisting the crossing of burns or reaching awkward places on the crags. Like his mother, Katherine Graham-Campbell of Shirvan, Lochgilphead, Archie was a keen cultivator of rhododendrons, and many fine specimens grow in the moist, wooded environs of Stronachullin. His other absorbing interest was the music of the highland bagpipe of which he was a very competent exponent in his younger days. Latterly an affliction of the tendons of the hands meant that playing to a high standard was no longer possible, but he remained a well-known composer of pipe music and was much in demand as a judge at piping competitions throughout Scotland. Archie Kenneth was a fine Highland gentleman of a style becoming increasingly rare. We shall 244 OBITUARIES miss him for his kindly, unpretentious manner and cheery companionship in the field, and Scottish field botany will be the poorer for his passing. PRINCIPAL PUBLICATIONS OF A. G. KENNETH 1964 The flora of Danna. Trans. Bot. Soc. Edinb. 39: 489-501. 1970 (With A. McG. Stirling) Notes on the Hawkweeds (Hieracium sensu lato) of western Scotland. Watsonia 8: 97-120. 1971 The flora of Danna — a supplement. Trans. Bot. Soc. Edinb. 41: 155-164. 1979 (With M. H. Cunningham) The flora of Kintyre. 1983 (With D. J. Tennant) The Scottish records of Dactylorhiza traunsteineri (Sauter) So0. Watsonia 14: 415-417. 1984 (With D. J. Tennant) Dactylorhiza incarnata (L.) So6 subsp. cruenta (O. F. Mueller) P. D. Sell in Scotland. Watsonia 15: 11-14. 1985 Additions to the flora of Kintyre. Glasgow Naturalist 21: 1-12. 1985 A hybrid club-moss, Diphasiastrum X issleri in Scotland. Glasgow Naturalist 21: 101. 1986 (With M. R. Lowe & D. J. Tennant) The status of Orchis francis-drucei Wilmott. Watsonia 16: 178-180. 1987 (With D. J. Tennant) Further notes on Dactylorhiza incarnata subsp. cruenta in Scotland. Watsonia 16: 332-334. 1988 (With M. R. Lowe & D. J. Tennant) Dactylorhiza lapponica (Laest. ex Hartman) So6 in Scotland. Watsonia 17: 37-41. A. McG. STIRLING A. G. KENNETH — AN APPRECIATION Archie Kenneth has contributed more to our knowledge of the Hieracium flora of Scotland than any other botanist since before World War I. Starting in his native Kintyre and moving on to Argyll, Perth, Inverness, Ross and Sutherland, as well as some of the islands, he developed an eye for hawkweeds in the field which brought him a wealth of new records, and resulted in a joint paper with his friend of many excursions, Allan Stirling. Nothing seemed to be too much trouble for him, and if more material of a plant was requested, off he would go the following year in search of it, however long and arduous the walk. One of his important early finds was a new species, H. solum, a disjunct member of the Series Alpestria found only in two localities in Kintyre. Some of his other most important finds are in Ross, and contain several new species still to be published. My last letter to him was to inform him that one of them was to be called “‘kennethiv’’. Archie had a wonderful way with people. He even controlled Mary McCallum Webster on a B.S.B.I. excursion, giving her all manner of excuses why he had not taken the party to see the Danna dactylorchids. He explained afterwards, that actually he was taking me the next day so that I could look at them in peace without any outside distraction. I prayed that Mary would not find out, but that thought did not seem to bother him at all. He holds a special place in my heart for the way he enabled Cyril West to continue looking at living hawkweeds long after he was unable to get to them himself. His wife even cooked special dishes that Cyril was fond of to keep him happy during his stay at Stronachullin. Archie loved his coffee, and I can see him now, explaining to the manageress of a Ross hotel that a demi-tasse was no fit amount of drink for a man who had been on the hills all day. For the rest of our stay at least, everyone in the hotel dining room had a large cup of coffee (or two) if they wanted it. May I give a special thank you from those south of the border, like myself, who benefited by his quiet and happy companionship in the hills. P D. SELL OBITUARIES 245 GUY MALCOLM SPOONER (1907—1989) All who knew Malcolm Spooner will mourn the death of a distinguished marine biologist, a remarkable and versatile field naturalist, and a formidably knowledgable but unassuming and immensely likeable man. Born in Yelverton, Devon, on 26 June 1907, he was educated at Charterhouse School and Christ’s Collge, Cambridge, where he graduated with a First in Zoology in 1929. On leaving Cambridge he joined the staff of the Plymouth Laboratory of the Marine Biological Association, where his work included pioneer research on the genetics of gammarids and, after the war, investigations of the uptake of radioactive fission products by seaweeds. He was awarded the M.B.E. for his wartime code-breaking work at Bletchley Park. He and his botanist wife Molly were married in 1943. For many years he edited the Journal of the Marine Biological Association. Outside his professional field, he was a fine entomologist, with a wide field knowledge of diverse groups of British insects, and he was elected a fellow of the Royal Entomological Society in 1959. He was a leading authority on British aculeate Hymenoptera, and it was through his useful papers on various genera of these insects that I first came to know of him in my student days. Later, he was quick to correct the identification of a hoverfly I had miscaptioned. It was he who first showed me a brown hairstreak butterfly, and the rare ant Formica exsecta on field excursions in Devon. I well remember his keen pleasure in the flies and solitary wasps on the umbels of Angelica which grew in their comfortably natural garden at Yelverton — and in the Gymnocarpium dryopteris which had long become naturalized in the shade of the house. Malcolm Spooner’s botanical activities were centred on his native south-west Devon. Wistman’s Wood on Dartmoor was a particular and abiding interest. He had known the wood for many years when, in late summer 1964, we sat talking among the gnarled oaks with two Nature Conservancy staff, who expressed misgivings at the wood’s apparent state of decline. Malcolm’s very characteris- tic response was to seek out the sites of old published photographs of the wood, and to take comparable new photographs from the same spots. These showed what surprised almost everyone at the time; that, far from receding, the wood had almost doubled in area since the early years of the 20th century. Malcolm and Molly repeated the measurements in the wood made in the early 1920s by R. H. Worth, and over the years he built up a comprehensive bibliography of the many references to the wood in botanical and topographical literature. The results of this work appeared in a paper of which I was privileged to be a joint author in the Transactions of the Devonshire Association for 1980. In 1953, he and F. S. Russell re-edited the published papers of R. H. Worth as Worth’s Dartmoor, still a valuable reference work on the Moor. After his retirement from the Plymouth Laboratory, Malcolm devoted much time and many miles’ walking to recording the flora of the Dartmoor tors, and he and Molly made an important contribution to tetrad recording for the Atlas of the Devon Flora. He was for 15 years a member of the Dartmoor National Park Committee. For many years he was active in the Devonshire Association, whose 7ransactions he edited from 1967 to 1972, and of which he was President in 1979. He was a founder member of the Devon Wildlife Trust, and a valued and active member of its Council. M. C. F. Proctor ANNA YOUNGER (1901—1989) The death of Mrs Anna Younger has removed from the Scottish botanical scene a true stalwart who was deeply revered and much respected, who was untiring in her efforts to encourage field botany, particularly in Scotland, although her interests were by no means confined to that country. She was deceptively frail in stature and uncomplainingly bore the scars of several field accidents, but she remained a person of unbounded determination in searching for and finding the rarest species in the British flora. Even when in her late eighties, she was accustomed to dash off to see plants in any corner of the British Isles when a finding of a new species to her meant a further drawing in her Bentham & 246 OBITUARIES Hooker Illustrations. In her haste, she had several brushes with authority from which she managed by her charm to escape conviction! She was, however, no mean botanist and she was a particularly keen student of critical groups such as the genera Carex and Potamogeton. She was a most generous patron of the Arts and local charities, and she herself had a high reputation as an accomplished embroideress. She will be much missed within the wide circle of her friends in Scotland. G. TAYLOR WATSONIA Botanical Society of the British Isles Patron: Her Majesty Queen Elizabeth the Queen Mother Applications for membership should be addressed to the Hon. General Secretary, c/o Department of Botany, Natural History Museum, Cromwell Road, London, SW7 5BD, from whom copies of the Society’s Prospectus may be obtained. Officers for 1990-91 Elected at the Annual General Meeting, 12th May 1990 President, Professor D. A. Webb Vice-Presidents, Dr H. J. M. Bowen, Dr F. H. Perring, Dr A. J. Richards, Mr J. Ounsted Honorary General Secretary, Mrs M. Briggs Honorary Treasurer, Mr M. Walpole Back issues of Watsonia are handled by Messrs Wm Dawson & Sons Limited, Cannon House, Folkestone, Kent, to whom orders for all issues prior to Volume 18 part 1 should be sent. Recent issues (Vol. 18 part 1 onwards) are available from the Hon. Treasurer of the B.S.B.1., 68 Outwoods Road, Loughborough, Leicestershire. GRAY HERBARIUM MAR 07 199% Watsonia, 18, 247-255 (1991) 247 Presidential Address, 1990 D. A. WEBB GENERA, HOLLOW CURVES AND BOOJUMS Fellow-botanists, As most of you know I live in a remote western isle, and have not been able until today to attend a Presidential address. When I inquired what would be expected of me, some said that I should instruct, others that I should challenge or puzzle, others again that I should amuse. Unwisely, perhaps, I have tried to combine all three, and I can only hope that the resulting rather ill-mixed cocktail may contain at least one flavour to everybody’s taste. What is a genus? That is the question to which I want to devote most of my address. Hollow curves and boojums will be touched on, but they are only incidental décor, designed, in the immortal phrase, to give artistic verisimilitude to an otherwise bald and unconvincing narrative. Most of us, I imagine, take species more seriously than genera: we argue passionately for or against the recognition of species A as distinct from species B, but on disputed points such as whether we say Scirpus caespitosus or Trichophorum caespitosum we are apt simply to follow our favourite Flora, as if it had been written on stone and handed down from Mount Sinai. What, then is a genus? The standard reply is that it is the rank in the agreed taxonomic hierarchy which stands between the family and the species. Fair enough, but this tells us only the relationships of the genus, not its essence. What does it look like? How do you know when you have caught one? Until recently many people would have replied: it is the primary and basic unit of plant classification. It is the plain man’s concept of a distinct ‘kind’ of plant, whereas what we call species he usually thinks of as varieties. A genus has, after all, the form of a noun, under which species are distinguished by adjectives. ‘“‘A rose is a rose is a rose’’ said Miss Gertrude Stein, and broadly speaking she was correct. By this I mean that any plant which the plain man calls a rose without qualification (and thus excluding rock-roses, roses of Sharon and so on) belong to the genus Rosa. (I am aware that some people transfer Rosa persica to the genus Hulthemia, but personally I am not convinced.) Conversely I mean that the plain man would recognize as a rose any plant belonging to the genus Rosa. (Of course if the plain man is too plain he may simply call it a briar, and include under the same head species of Rubus or even Smilax. But we must postulate a plain man who gives at least a cursory glance at leaves, flowers and fruits, and not merely at the condition of his clothes.) This suggests that a genus is the aggregate of all those plants which the observant but uninstructed layman denotes by a vernacular name. Let us see if we can find other genera as satisfactory in this respect as Rosa. What about Viola? Most of its members are regarded as violets, but although few taxonomists want to raise the subgenus Melanium to the level of genus, gardeners and unsophisti- cated wild-flower-lovers usually distinguish violets from pansies. Well, what about Ranunculus? Many of its species are plainly buttercups, but the white-flowered aquatics are not, and what about the Lesser Celandine? If we go abroad the case is even worse. When I was in the Lebanon in spring many years ago I saw, as I imagined, sheets of scarlet anemones. It was not till I chanced to see the calyx of a flower that had been trodden down that I realized they were Ranunculus asiaticus. But they were certainly not buttercups. Finally, let us consider the gentians. In the good old days (by which I mean, of course, the days when I learnt my botany) all those blue jobs with campanulate or stellate flowers were species of Gentiana. But then some bright boys at Cambridge decided to resurrect Moench’s long-dead genus Gentianella and apply it to the annual or biennial species with a little fringe of hairs at the top of the corolla-tube. To the plain man, however, they are all still gentians, and it is not easy to persuade him that Gentiana nivalis is more closely related to Gentiana lutea than it is to Gentianella amarella. Indeed I sometimes wonder myself whether it is. This leads me on to another suggested criterion. Many botanists today, when discussing these questions, say that a good genus should have what it is now fashionable to call a characteristic ‘jizz’. This is, of course, what the plain man relies on when he sorts plants into ‘kinds’. But the examples 248 D. A. WEBB just considered show that while many genera have this characteristic ‘jizz’ there are many others that do not. I do not think that anybody questions the inclusion in the genus Fuchsia of both Fuchsia excorticata, which can grow into a tree of 15 m, and Fuchsia procumbens, a creeping herb rarely more than 1 cm off the ground. They agree in floral structure, but in little else. We just have to accept the fact that from this point of view some genera are satisfactory and others are not, and that no amount of tinkering will make them entirely satisfactory. It is, of course, easy to find plenty of smallish genera whose species all look much the same to the plain man. Myriophyllum is one such, even when surveyed on a world-scale. But the allegedly popular name ‘water-milfoil’ is redolent of the book rather than the ditch. The Latin name was formed by Clusius in the 16th century; the English equivalent has been widely used for Hottonia and Utricularia as wellsas for Myriophyllum. But the plain man either ignores these plants or calls them “‘one of them mucky water-weeds.”’ This is all rather disconcerting to the seeker of the generic concept. The gentians teach us that a genuine popular name can straddle two genera, and indeed we know that already if we call to mind pimpernel, loosestrife and celandine. Moreover, not all members of a genus share the same vernacular name: primrose and cowslip are old names firmly established, but nobody questions the inclusion of both in Primula; Bentham, indeed, wanted to unite them in a single species. I might say here, in parenthesis, that it is not always easy to say whether a name is truly vernacular or not. We can easily recognize and reject such nineteenth-century inventions as ‘few-flowered spike-rush’; nobody ever talked of a spike-rush till Bentham was driven to invent an English name for Eleocharis. But what about ‘goosefoot’? Is it a relatively late translation from the Latin, or is Chenopodium a Latinization of an old vernacular name? To answer such a question needs a lot of research in early herbals and in dictionaries — not only English, but also German and Swedish. It would appear in this particular case hahnenfuss is an old German vernacular name; it was latinized in the 16th century and then, relatively recently, translated to English. This is a warning against insularity, a point to which I will return later. It is also easy to fall into the error of assuming that because an English name is very familiar, and understood by many whose knowledge of botany is slight, that it must be vernacular. ‘Saxifrage’ provides a good example. Until the mid-eighteenth century the use of the word in English was confined to Saxifraga granulata, and in this sense it was certainly a translation by Gerard of the Saxifraga used in medieval herbals. Linnaeus was the first author to give the genus its modern circumscription, and it is a bit much to suppose that the untutored sheep-farmers of Cumbria were applying the name saxifrage to such species as Saxifraga aizoides or Saxifraga stellaris, when Ray and Tournefort were firmly including the first in Sedum and the second in Geum. Still less likely is that they were calling them rockfoils, as William Morris would have wished. There are indeed some English names beloved and cherished by Flora-writers which I find it difficult to regard as truly vernacular. I try to imagine myself hearing Farmer Hodge in the Plowright’s Arms saying: “‘It’s the wettest April I can remember; the stream by the cow-byre is nigh choked with blinks already’’. I try, but I find it difficult. And I am reassured when I discover that ‘blinks’ as a name for Montia was more or less invented by Hooker. If, bearing all these cautions in mind, we make a list of genuinely vernacular names we find that many, indeed, do correspond fairly closely to modern concepts of a genus, but there are almost as many which designate a single species, and are not applicable to other species of the same genus. On the one hand we have buttercup, clover, lime and poppy, with a generic denotation; but against these we must set weld, tutsan, sycamore and meadow-sweet, which denote only a single species within a somewhat varied genus. The fact that many British plants, common enough to have been given a vernacular name, belong to genera of which there is only one British species introduces an ambiguity, but personally I find it hard to believe that such names as sea-kale, horse-radish, traveller’s joy or kingcup would be applied to other species of Crambe, Armoracia, Clematis or Caltha if we had them. Linnaeus attached tremendous importance to the genus as the corner-stone of taxonomy, and a few modern authors are inclined to follow him blindly. But we should remember the tag Vixere fortes ante Agamemnona: Agamemnon was not the first of heroes, nor was Linnaeus the first of taxonomists. He was not working on a tabula rasa, but was to some extent in thrall to his predecessors — not only men like Ray and Bauhin, but also to the unknown Greeks who distinguished parsley from hemlock. Walters (1961) has done a useful service in emphasizing this; PRESIDENTIAL ADDRESS, 1990 249 but I cannot follow him when he affirms his adhesion to Linnaeus’ view of the genus as the corner- stone of taxonomy. For I am convinced that if there is a single fact which the so-called ‘New Taxonomy’ (now just over 50 years old) has taught us, it is that the species has (in most cases) an objective reality which other ranks do not possess. Talk as much as you like about Euphrasia and Festuca, it remains true that the great majority of plants fall into groups which are interfertile within the group but reproductively isolated from other groups, and which are morphologically distinctive and only moderately morphologically variable. These are what we call species, and they represent our primary data. A saying is attributed to the German mathematician Kroneker, that God made the integers, but all else is the work of man. Substitute ‘species’ for ‘integers’ and, if you like, ‘evolution’ for ‘God’, and I believe the statement to be true. So far I have been explaining, with some diversions, what a genus is not. It is time to return to my original question. I can only answer it by saying that a genus is a convenient bundle of species, with enough features in common for us to be able to apprehend the bundle as a unit and to make some generalizations about it. This description could, of course, be applied to a tribe or a section as well as to a genus, but nobody bothers about tribes or sections until genera have been established. The difficulty lies, of course, in the word ‘convenient’. A bundle of species convenient to one man may be inconvenient to another. I need only remind you that the gardener finds it convenient to keep his Cactaceae and Crassulaceae in the same greenhouse, while the taxonomist finds it convenient to put them in different volumes of his treatise. A generic bundle which is convenient to the man obsessed with chromosome numbers is likely to be inconvenient to the pollen electroscan man or the phytochemist. We find ourselves, whether we like it or not, in the shifting sands of tradition, common sense, consistency and the fight between stability and improvement. Walters (1961, 1986) has considered the opposition between these last two, and has come down fairly heavily on the side of stability. It is an attitude with which I have a lot of sympathy, but I think that he skates lightly over some difficulties. To begin with, is it reasonable to accept improvements made fairly recently, which in their day were looked on as just as disturbing as those which it is now proposed to reject? I am reminded of Tansley’s famous answer when he was asked what exactly he understood by conservation. “It usually means’, he replied, “keeping things much as they were when the speaker was young.”’ Some 50 years ago Gilbert-Carter described Hooker’s Student’s Flora of the British Isles as ‘“‘the only Flora fit to be put into the hands of a student’’, but it is rather disconcerting to see how many genera which most of us take for granted today are not recognized by Hooker: they include Sinapis, Filipendula, Aphanes, Sorbus, Berula, Knautia, Cirsium and Vulpia. And I am talking, of course, not merely of nomenclatural but of taxonomic distinctions. The general acceptance of these genera suggests that the improvements of this century have been useful and welcome. For this reason I have some hesitation in deciding that a line must be drawn just now. But I hold that new generic fusion or splitting should be done only for a very good reason. In Flora Europaea we had a working rule that if a species couldn’t be honestly keyed out without a profusion of ‘usually’, ‘more or less’, or overlapping measurements, then it wasn’t a good species. I think the same should apply to genera. I am myself responsible for a fusion which is probably not very popular, but when I sank Peplis in Lythrum it was simply because among the species of south- eastern Europe I was unable to find a clear distinction. I shall return to these genera later. There is, however, a further and more compelling reason for considering that a ‘freeze’ of genera in their present form is impracticable. This is the lack of agreement among recent and authoritative Floras and monographs, coupled with the fact that for some groups in some parts of the world there has been no authoritative treatment at generic level for well over a century. It is all very well to say “let us follow Cronquist for families and put an end to disputes”, but there is no modern equivalent to the Genera Plantarum of Bentham and Hooker. Anybody who tries to sort out into genera a world-wide collection of, say, the Andropogoneae (a tribe of grasses), or — God save us all — certain families of ferns, soon finds that he is forced to take arbitrary decisions because of lack of agreement among even recent revisions. For few of these are world-wide in their scope. It is all too easy to accept as definitively authoritative a scheme based primarily on the Flora of our own region, and perhaps by a taxonomist we know and respect; but this respect may not be shared by taxonomists in other parts of the world. We have with some difficulty digested the relegation by Melderis of our familiar species of Agropyron to Elymus, but Agropyron still exists, and there are many species in Central Asia for which we have no guidance as to their correct generic niche. Again, Californian Floras refuse to recognize a distinction between Convolvulus and Calystegia, because the 250 D. A. WEBB 70 60 50 ne c ro) 40 fo) we) 7 re} = = 30 20 10 Fe DP TES I Gr 1h HF EBAY 7 AY SerG JOM OSHS IZ Number of species per genus Ficure 1. ‘Hollow curve’ for the Acanthaceae, omitting genera with more than twelve species. distinction, so clear in Europe, is blurred in California. The most widely used, and in many respects the best, Flora of Japan includes Vulpia in Festuca, and Reynoutria in Polygonum. A regime of stability if applied everywhere would mean permanent provincialism. As the detailed knowledge of species advances generic revision must occasionally follow. I have no objection to this so long as the changes are made only when all the evidence has been weighed, and not as a consequence of some idiosyncratic theory, or as a means of getting our name into the next supplement of Index Kewensis. I turn now, very briefly and inevitably superficially, to the topic of hollow curves, and I do so partly because they have lately been in the news, but also because my last point — that we have not reached finality in the delimitation of genera or families — is relevant to our assessment of them. The phrase, ‘hollow curve’, introduced by Willis (1922) nearly a century ago, indicates a phenomenon to which he was the first to draw attention. In nearly every large family of angiosperms it seems that there are a very few really large genera (that is containing some hundreds of species), a moderate number of middle-sized genera, and a large number of very small genera, many of them with only one species. It follows, therefore, that if you plot number of species per genus against number of genera containing that number of species, you get a curve which, at least at a superficial glance, looks like a hyperbola (Fig. 1). There are various other ways of plotting the same data, but the resulting curve is always much the same (Fig. 2). PRESIDENTIAL ADDRESS, 1990 251 300 Ranunculus 250 Clematis a 200 = ® D © Q wn & ary «D0 Q yn ao) Anemone fe) za 100 Thalictrum 50 Viorna group of small monotypic genera genera 1 2 3 4 5 6 7 8 Dogs) | 1 HZ tse + 14 No. of genera Ficure 2. ‘Hollow curve’ for the Ranuculaceae, tribe Anemoneae. The explanation of this curve was originally proposed by Willis in evolutionary terms which nobody accepts now, but a dispute is current as to whether — as Walters (1986) maintains — the curve is a man-made phenomenon, dependent on the way that our concept of the family and the genus developed, or, as is argued in a recent paper by Cronk (1989), the shape of the curve is at least in part determined by the pattern of variation which exists in nature. I confess that I find my sympathies divided in this argument, not least because I cannot entirely understand either side of it. My real motive, however, for bringing hollow curves into this address at all is to suggest that both sides are taking too seriously the figures on which their curves and their arguments are based. It is no use imagining that numbers of species per genus and family limits are fixed for ever. Opinion is very evenly divided as to whether we should treat the Papilionaceae as a family, or lump it in with others to make the Leguminosae. Whichever we do, of course, the curves will remain hollow, but some will be hollower that others. Again, pressure is steadily mounting for fragmenting the genus Polygonum into five or six, and, as it is one of the two really big genera in the family, if it is dismembered the curve for Polygonaceae will develop something of a middle-aged spread. Table 1, on which Fig. 2 is based, illustrates the same point. It is clear that a number of the monotypic genera are recent splits, and I should be very surprised to learn that they are universally 252 D. A. WEBB accepted. Again, Haworth’s attempt to split Saxifraga into about a dozen genera, made almost 200 years ago, has been taken up by some quite respectable American Floras; if it were followed the hollow curve for Saxifragaceae (a family which, incidentally, everyone delimits differently) would be in a sorry state. None of these considerations will abolish the phenomenon of the hollow curve, but the curves will vary to an extent sufficient to suggest that the somewhat sophisticated mathematical treatment proposed by Cronk rests on a very unstable basis. I would go so far as to suggest that his analysis is of more interest to the applied mathematician than to the biologist. This is not to deny that some of the suggestions and speculations made elsewhere in his paper are interesting and stimulating, as also are those of Walters, but they lose rather than gain by being linked to a geometry which is liable to be upset by the next taxonomic revision. May I be allowéd to end on a rather more frivolous note? I re-read recently that somewhat neglected masterpiece of Lewis Carroll’s, The hunting of the snark. As many of you probably know it tells of the voyage of a ship to a land where the crew hope to catch a rather mysterious monster — the snark. The crew are an oddly assorted list; all they have in common is that their occupations begin with the letter B. I shall pass over without comment the billiard-marker, the broker, the boot- and the bonnet-maker, but I shall consider briefly the others, who have some relevance to my general theme; these are the beaver, the butcher, the barrister, the bellman and the banker. I take these as symbolizing various types of what I conceive to be the wrong approach to the problems of generic delimitation. The beaver is the man like the late-lamented H. P. Fuchs of Basel, or Dr Josef Holub (whom God preserve) of Prague; such men spend their lives beavering away in ancient and TABLE 1. SIZE OF GENERA IN RANUNCULACEAE, TRIBE ANEMONEAE, FROM WILLIS (1931) Genus No. of species Ranunculus 300 Clematis 220 Anemone 130 Thalictrum 75 Viorna (Clematis p.p.) 18 Adonis 10 Oxygraphis 10 Clematopsis (Clematis p.p.) 10 Myosurus 7 Naravelia (Clematis p.p.) Rhopalopodium (Ranunculus p.p.) Knowltonia (Anemone p.p.) Trautvetteria Barneoudia (Anemone p.p.) Hamadryas Halerpestes (Ranunculus p.p.) Leucocoma (Thalictrum p.p.) Anemonanthea (Anemone p.p.) Capethia Aiolon (Anemone p.p.) Arcteranthis (Oxygraphis p.p.) Aspidophyllum Beckwithia (Ranunculus p.p.) Gampsoceras (Ranunculus p.p.) Kingdonia Kumlienia (Oxygraphis p.p.) Laccopetalum (Anemone p.p.) Paroxygraphis Piuttia (Thalictrum p.p.) Stipularia (Thalictrum p.p.) Sumnera (Thalictrum p.p.) Syndesmon (Anemone p.p.) Viticella (Clematis p.p.) eee OLN HD WD PDP PON OO ~) ~) PRESIDENTIAL ADDRESS, 1990 253 obscure literature and coming up with totally unfamiliar names. These are mostly specific epithets, but occasionally they resurrect distressing genera such as Logfia or Ifloga. The butcher is the man who, when he sees a large genus, has an irresistable impulse to cut it up into joints of household size. This is a very natural impulse, for very large genera are a nuisance; unfortunately, however, some of them seem very resistant to butchery. I am afraid I must typify the butcher by Dostal, a taxonomist for whom I have a considerable respect; but he caused a crisis in Flora Europaea by proposing to split Centaurea into eight or nine pieces. We allowed him to detach a few small fragments, and he might have got away with the major butchery had he not disclosed that the genus containing several very familiar species was to bear the name of Sagmen. At that the committee struck. The barrister is the taxonomist who believes that generic limits can be settled by laws or rules: the rule most frequently put forward is that if two plants can hybridize they must belong to the same genus. If orchids are mentioned the barrister is apt to change the subject. The bellman, who is, incidentally, the captain of the ship, reveals his weakness in his own words: ‘Just the place for a snark’, the Bellman cried As he landed his crew with care. ‘Just the place for a snark’. I have said it twice That alone should encourage the crew. ‘Just the place for a snark’; I have said it thrice. What I tell you three times is true. We have all come across bellmen, commoner, I am glad to say, across the Atlantic than here, who publish an idiosyncratic generic revision, and then republish it twice in different journals and assume that it is generally accepted. The banker is, like all bankers, a natural conservative. He values stability of exchange and interest rates above all other considerations, and assumes that any change in them will be for the worse. He shuts his eyes to the fact that on many problems of generic delimitation there is no general agreement: the yen and the dollar, as we have seen, do not always follow the pound. He also ignores, as I have suggested earlier, the fact that some of the things he regards as lynch-pins of his stable world were regarded by the previous generation of bankers as dangerous innovations. In a different way from the others, but just as confidently, he is sure that his concept of the genus is one with which all right-thinking men must agree. I have omitted one important member of the crew, the baker. I am afraid that here I must cheat a little, for, although he has some relevance to my main problem, he reminds me most, not of an errant taxonomist, but of a particularly troublesome species. The baker, as some of you may remember, arrived for the voyage with 42 boxes, with his name painted clearly on each, but owing to some misunderstanding they were left behind on the beach. He was wearing plenty of clothes, but without the boxes he couldn’t remember his name. He would answer to ‘Hi!’ or to any loud cry Such as ‘Fry me!’ or ‘Fritter-my-wig!’ To ‘What-you-may-call-um!’ or ‘What-was-his-name!’ But especially “‘Thing-um-a-jig!’ His intimate friends called him ‘Candle-ends’ And his enemies “Toasted-cheese’. I think we all know species to which this applies. I found one when I was writing the Lythraceae for Flora Europaea, and I was in doubt as to whether it belonged to Lythrum or to Peplis. My doubt had been shared by others, as can be seen from its full synonymy: Lythrum borysthenicum (Bieb. ex Schrank) Litwinov in Maevski, Fl. Sred. Ross., 3rd ed, 5: 1209 (1917). Lythrum nummulariifolium Loisel. in Desv. J. Bot. 2: 330 (1809), non Persoon, Syn. 2: 8 (1806). Peplis borysthenica Bieb. ex Besser, Enum. Pl. Volhyn. 81 (1822), nomen nudum. P. borysthenica Bieb. ex Schrank in Flora (Regensb.) 5: 643 (1822). 254 D. A. WEBB P. erecta Req. ex Bentham, Cat. Fl. Pyr. Bas-Languedoc 111 (1826), nomen nudum. Ammania borysthenica (Schrank) DC. in Mém. Soc. Phys. Genéve 3 (2): 79 (1826). Peplis australis Gay ex Arnott in Edinb. New Phil. Jour. 2: 250 (1827), nomen nudum. P. biflora Salzm. ex DC., Prodr. 3: 77 (1828). Lythrum bifiorum (Salzm. ex DC.) Gay in Ann. Sci. Nat. 26: 227 (1832). Peplis erecta Req. ex Moris, Fl. Sard. 2: 67 (1840). P. tithymaloides Bertol., Fl. Ital. 4: 233 (1841). Ammania boraei Guépin, Suppl. Fl. M.-et-L. 39 (1842). Middendorfia borysthenica (Schrank) Trautv. in Flora (Regensb.) 22: 496 (1842). M. hamulosa Trautv. in Mém. Acad. Sci. Petersb. (Sci. Phys. Math.), ser. 6, 4: 491 (1842). Peplis nummulariifolia Jordan, Obs. pl. crit. 3: 85 (1846). P. boraei (Guépin) Jordan, Obs. pl. crit. 3: 81 (1846). P. timeroyi Jordan, Obs. pl. crit. 3: 83 (1846). P. hispidula Durieu in Duchartre, Rev. Bot. 2: 431 (1847). Lythrum hispidulum (Durieu) Koehne in Sitzb. Bot. Ver. Brandenb. 22:29 (1880). Lythropsis peploides Welw. ex Koehne in Bot. Jahrb. 1: 305 (1881). Lythrum loiseleurii Rouy & Camus, Fl. Fr. 7: 164 (1901). Peplis australis Gay ex Sampaio, Fl. Portug. 305 (1911). Lythrum boraei (Guépin) Klokov, Fl. R.S.S. Ucr. 7: 401 (1955), in obs. With such a rich synonymy one can be excused for thinking that the addition of “‘Toasted-cheese’ to the list would scarcely be noticed. It will be seen that the species has been assigned to four different genera. Middendorfia and Lythropsis (which can be regarded as synonymous) represent desperate attempts to find a home for those plants which seem intermediate between Lythrum and Peplis as traditionally interpreted, but their recognition would merely mean that we had on our hands three genera hard to define instead of two. The fact is that Linnaeus and his predecessors, impressed by the numerous and conspicuous differences that separate Lythrum salicaria from Peplis portula, put them without hesitation in different genera; it was only the gradual discovery later of various dingier and dingier Lythrum-like plants from southern Europe that eroded the distinction and convinced me that only one genus could be maintained. So, in spite of disapproving noises from some conservative bankers I sank Peplis in Lythrum, and I seem, rather to my surprise, to have got away with it. I have strayed rather far from the snark and its hunters, whom, I may remind you, I portray as seekers after the concept of the objective, scientific, universally accepted genus. They set about their hunt in a business-like way: They sought it with thimbles, they sought it with care; They pursued it with forks and with hope; They threatened its life with a railway-share; They charmed it with smiles and with soap. But they receive a nasty shock when the nameless baker reveals to them a warning given him by his uncle on his deathbed. If your snark be a snark that is right; Fetch it home by all means — you can serve it with greens — And it’s handy for striking a light. But oh, beamish nephew, beware of the day, If your snark be a boojum! For then You will softly and suddenly vanish away, And never be met with again! The crew were shaken by this, but they resolve all the same to proceed with the hunt. After many adventures they saw the baker gesticulating from a distant crag before he plunges into a chasm. PRESIDENTIAL ADDRESS, 1990 2D) ‘It’s a snark!’ was the sound that first came to their ears, And seemed almost too good to be true; Then followed a torrent of laughter and cheers; Then the ominous words ‘It’s a boo~’ In the midst of the word he was trying to say, In the midst of his laughter and glee, He had softly and suddenly vanished away, For the snark was a boojum, you see. Well, it is my contention that all snarks are boojums, and that snark-hunting is best abandoned; but by this I do not mean generic scrutiny and possible revision, for I remind you once again that I use the snark as the symbol of the would-be absolute and eternal definition of a genus, determined on a priori principles. Let us recall the nursery rhyme: One, two, buckle my shoe; Three, four, shut the door; Five, six, pick up sticks; Seven, eight, lay them straight. Picking them up is the job of the species-maker. Laying them straight is what I have mainly been talking about, if we interpret ‘straight’ to mean ‘arranged in bundles of convenient size and composition’. ‘“‘Let us’, in the sonorous words of Ecclesiastes, ‘“‘hear the conclusion of the whole matter’’. Approach the business of generic revision in the spirit in which the 1662 Prayer Book exhorts us to approach matrimony: not unadvisedly, lightly or wantonly, but discreetly, advisedly and soberly. Don’t disturb wantonly such bundles as you find, but don’t be afraid to make discreet adjustments if you have considered the matter from all angles and are confident that it will improve the convenience of the bundles concerned. Above all, don’t go snark-hunting on the basis of a single general principle, for if you do you will deserve, even if you do not actually meet with, the fate of the baker. REFERENCES Cronk, Q. C. B. (1989). Measurement of biological and historical influences on plant classification. Taxon 38: 357-370. WALTERS, S. M. (1961). The shaping of angiosperm taxonomy. New Phytol. 60: 74-84. Wa ters, S. M. (1986). The name of the rose: a review of ideas on the European bias in angiosperm classification. New Phytol. 104: 527-546. WILLIs, J. C. (1922). Age and area. Cambridge. WILLIs, J. C. (1931). A dictionary of the flowering plants and ferns, 6th ed. Cambridge. i ] we es Sane nai er witial enlist (Sch ove apt An win + o—, ns eat . ot Ma rye - ee dw (RA REQ Oy os bEracoen malt yer | es | . ) ae ileal ? ape! sir to tebimn ot nl err a . ? Ly iiehth chs wade ban y tytcz ha oH he a oan Y ecete | ‘iid + a5 edsehz) 2d oy pl . &. ere Ph Adath. |. wer. 7s (R42 snath ale eye fiawe-ds he ends bate. tall fateheig caine fy sad ae ea ah i ix Pitas tps 27 wit aeeel nities Dus yativts< oi hense teat soe Ob: a gin 53 etme < Have ailia lodemya cork 20 Le _siqioniny ; t ; 7 Pel 1 i a werent. eet AP |Lna: T ay oaks famine. « : a5 4 ry 4 as ; * 2 : by od ‘+s 4 ay: * + ’ poten Ochi ts ~~ | t - 71 a = 1 7 -yt 24 + oo r r J j “ ‘ a ? 4 soe t J F ” ’ ) 5 Oi { ’ me fee Watsonia, 18, 257-295 (1991) 257 An account of Orobanche L. in Britain and Ireland F. J. RUMSEY and S. L. JURY Department of Botany, University of Reading, P.O. Box 221, Reading, Berkshire, RG6 2AS ABSTRACT Morphological descriptions are given of the 14 species of Orobanche (Orobanchaceae) recorded in the British Isles, together with separate keys for identifying fresh material and herbarium specimens. Accounts of the history of the species are presented together with illustrations and distribution maps. The variation in Orobanche minor is accounted for with the recognition of four varieties. INTRODUCTION The genus Orobanche is renowned as a taxonomically very difficult one. In most cases this is a result of many of the useful characters becoming lost on drying, and the lack of adequate field notes. Plants which are very distinct in the field become reduced to a hideous brown uniformity when pressed. Therefore, herbarium specimens are often incorrectly determined (an average of S—10% in fact). The loss of characters on drying, considerable intra-specific variation, confusing synonymies, incorrectly cited names and badly described species with poor types (often with different species on the same sheet) have done little to generate interest in the genus. Too many botanists have shown a reluctance to deal with this genus in herbaria, perpetuating the myth that the species are impossible to identify once dried. Certainly, Orobanche minor Sm. and its close relatives often cannot be positively determined without descriptive notes made at the time of gathering, but all other species from the British Isles are distinct enough not to need any additional information. It is hoped that this account will stimulate other botanists to study, identify and record members of this fascinating parasitic genus in Britain and Ireland, as well as clear up some errors and confusions made in the past. If more than one English name is in common usage, then they are all given, but the first is the name recommended by Dony, Jury & Perring (1986). Chromosome numbers are given where known. Counts from British material (Hambler 1958) are marked by an asterisk. All other counts are from European material (Moore 1982). Distribution maps are based on herbarium material from BM, CGE, E, K, LIV, MANCH, NMVW and RNG, together with selected records from the Biological Records Centre. CHARACTERS OF TAXONOMIC IMPORTANCE Populations of Orobanche species are often enormously variable with regard to such characters as size, colouration and pubescence, yet these details are seldom recorded on specimen labels. Because of this variation and their use in the taxonomy of the genus, care should be taken not to rely on a single character in isolation. Most of the British species have a restricted host range and it is therefore important to identify and record this. Unfortunately, this seemingly simple task is often almost impossible: it is not easy (nor often desirable!) to dig up the plants to observe connections between host and parasite. If the host species has to be guessed, this should be stated, and other surrounding species noted. Too often an incorrect plant has been given as the host and this has wrongly influenced later determination. Other characters to pay particular attention to include: the overall stature of the plant; the degree of swelling at the base (though where a species has several host species, such as Orobanche minor, these host taxa will obviously influence the Orobanche morphology); colour and degree of stem 258 F. J. RUMSEY AND S. L. JURY pubescence; length and pubescence of bracts; presence or absence of bracteoles; length, shape and venation of the calyx (though this may vary considerably on a single plant); position, shape and size of the lobes of the upper and lower corolla lips, the height of insertion of the stamens, the position and degree of the stamen pubescence, and the colour and degree of fusion of the stigma lobes. Pubescence of the anthers has been used as an important diagnostic character for taxa within section Trionychon Wallr. but not for those within other sections of the genus (Webb & Chater 1972). All British taxa of section Orobanche have hairless anthers. Preliminary investigations of seed morphology (Rumsey unpubl.) reveal great variability in size and shape from single capsules and preclude reliable discrimination of the taxa considered here with a single exception. O. ramosa L. subsp. ramosa is distinct in having a secondary layer of reticulation to the surface of the cells and not just the small circular to ovoid pits seen in all other British taxa. KEYS TO THE SPECIES Two keys are provided. The first is for fresh material and includes characters not available from herbarium specimens; for these the second key should be used. However, as more reliable characters are included in the first key, this should be used whenever possible. KEY TO FRESH MATERIAL 1. . Flowers with bracteoles; capsule valves free aDOVE oo. - 222.225 cend-aucasacs ce neee tee eee 2 1. Flowers without bracteoles; capsule valves coherent above .. 202... ..scccne-cnce.aeoeeee eee eee 4 2. Stem usually simple, c. 8 mm diameter; corolla narrowly campanulate; on Achillea TIRE CL OVID ceo wre cigs toe Sepa las A oan Se le ee Se eee 3. purpurea 2. Stem branched, slender, c. 5 mm diameter; corolla infundibuliform or tubular-infundibuli- FOTN? Of) VATIONS, NOSUS: so- cneso seis ce oe ctlaero oso Doss» esi eee Sete eee Oe ea oan a 3 3. Flowers 18-35 mm, infundibuliform; pedicels to 20 mm; anthers hairy ......... 2. aegyptiaca 3. Flowers 10-18 mm, tubular-infundibuliform; pedicels to 8 mm; anthers glabrous 1. ramosa 4. Plant with ComSpICUOUS SWEEE SCOT ae. 205 cage c5 cee sae os cons Sede ec daar ee =) 4. Plant tnscented, or if witha SCene, then TOCIE «on. xx 9 le 3 \ @ §=Seees FiGure 16. Distribution of O. minor var. minor; @ = post-1950 record, © = pre-1950 record. The map includes all records of the species other than those attributed to the rarer varieties. is Fee Coren the ce ee ee ee ee ee ee eee OROBANCHE IN BRITAIN AND IRELAND Bie) FicureE 17. Distribution of O. minor var. maritima (Pugsley) Rumsey & Jury; © = post-1950 record, © = pre- 1950 record. 292 F. J. RUMSEY AND S. L. JURY ° xm £00 =, MILES eee Ff ¢ ee ee a : | tN | — — Ficure 18. Distribution of O. hederae Duby; @ = post-1950 record, © = pre-1950 record, + = introduction or casual. Py ees pon tlda en dev a We peated ee Yt aes ee ee ae ee ee ee eS oe ee ee OROBANCHE IN BRITAIN AND IRELAND 293 FicurE 19. Distribution of O. crenata Forskal; + = introduction or casual. 294 F. J. RUMSEY AND S. L. JURY Flowering stem simple, 15-50 cm, glandular-pubescent, scarcely swollen at the base, yellowish. Inflorescence rather dense. Bracts 13-25 mm, oblong to lanceolate, acute to acuminate. Calyx 8-15 mm, unequally bidentate or entire. Corolla 15-22 mm, yellow tinged with red, erecto-patent to patent; upper lip + emarginate, porrect; lower lip with conspicuously ciliate sub equal lobes. Stamens inserted 2-3 mm above the corolla base; filaments hairy. Stigma lobes yellow becoming reddish with age. Style scarcely exserted, slightly convolute. Flowering period June to August. Introduced deliberately, parasitic on Berberis. Sometimes difficult to distinguish from the closely related O. flava Mart. Material from Oxford could also be confused with O. rapum-genistae from which it differs in its hairy filaments and its flowers not dark red inside. Erroneously reported from Surrey, one specimen is of somewhat atypical O. elatiox (CGE), another is surprisingly O. caryophyilacea (herb. H. C. Watson, K). O. lucorum is currently known only from the Botanic gardens at Oxford (Bowen, RNG) and St Andrews (Ratter, E). At Oxford it parasitizes a range of Berberis cultivars. Given the scarcity of Berberis in native stations, if it is to become established in this country it will be found in parks and gardens. ACKNOWLEDGMENTS We would like to thank the following people for their assistance: Chris Preston, Lynne Farrell, Mike Jones, June Underwood, all the B.S.B.I. recorders and members who have provided information and the herbarium curators who have allowed access to material. REFERENCES ApaMS, K. J. (1984). Orobanche crenata Forskal — native in the British Isles? Watsonia 15: 161-162. BECK VON MannacetTrta, G. (1930). Orobanche in A. ENGLER, ed. Das Pflanzenreich 4 (261): 1-348. BENTHAM, G. & Hooker, J. D. (1930). Handbook of the British flora, 7th ed. Ashford. Bevis, J., KETTELL, R. & SHEPARD, B. (1978). Flora of the Isle of Wight. London. BicHarD, J. D. & McCuintock, D. (1975). Wild flowers of the Channel Islands. London. Bowen, H. J. M. (1968). Flora of Berkshire. Oxford. BuTCHER, R. W. (1961). A new illustrated British Flora, 2. London. CLAPHAM, A. R. (1952). Orobanche L., in CLAPHAM, A. R., TuTIN, T. G. & WARBURG, E. F. Flora of the British Isles. Cambridge. Dony, J. G., Jury, S. L. & PERRING, F. H. (1986). English names of wild flowers, 2nd ed. Reading. GARRARD, I. & STREETER, D. (1983). Wild flowers of the British Isles. London. Gi_mour, J. & WALTERS, S. M. (1954). Wild flowers — botanizing in Britain. London. Goon, R. (1984). A concise Flora of Dorset. Dorchester. HamB_er, D. J. (1958). Chromosome numbers in some members of the family Orobanchaceae. Bot. J. Linn. Soc. 55: 772-777. Hux ey, A. & TayLor, W. (1977). Flowers of Greece and the Aegean. London. JERMYN, S. T. (1974). Flora of Essex. Fingringhoe. Jones, M. (1987). Orobanche hederae Duby in the British Isles, in WEBER, H. Cur. & FORTSTREUTER, W.., eds. Parasitic flowering plants. Marburg. LeSueEur, F. (1984). Flora of Jersey. St Helier. LINNAEUS, C. (1763). Species plantarum, 2nd ed. Stockholm. Lous.ey, J. E. (1950). Wild flowers of chalk and limestone. London. Makrcetrs, L. J. & Davip, R. W. (1981). A review of the Cornish flora 1980. Redruth. MARQUAND, E. D. (1901). Flora of Guernsey and the Lesser Channel Islands. London. Martin, W. K. (1965). The concise British flora in colour. London. MILNE, M. (1986). Plant records. Watsonia 16: 183-198. MILNE-REDHEAD, E. (1985). More hosts of Orobanche. B.S.B.I. News 41: 29. Moore, D. M. (1982). Flora Europaea checklist and chromosome index. Cambridge. MUSSELMANN, L. J. & PARKER, C. (1982). Preliminary host ranges of some strains of economically important broomrapes (Orobanche). Economic Botany 3%: 270-273. PERRING, F. H. & FARRELL, L. (1983). British red data books: 1. Vascular plants, 2nd ed. Lincoln. Petcu, C. P. & Swann, E. L. (1968). Flora of Norfolk. Norwich. ey a ea Ee OROBANCHE IN BRITAIN AND IRELAND 295 PHILLIPS, R. (1977). Wild flowers of Britain. London. Puitp, E. G. (1982). Atlas of the Kent flora. Maidstone. PoLunin, O. (1969). Flowers of Greece and the Balkans. Oxford. PoLunin, O. & Hux ey, A. (1965). Flowers of the Mediterranean. London. Pus ey, H. W. (1940). Notes on Orobanche L. J. Bot., Lond. 78: 105-116. Ray, J. (1690). Synopsis methodica stirpium Britannicarum. London. Ray, J. (1724). Synopsis methodica stirpium Britannicarum, 3rd ed. London. RIDDELSDELL, H. J., HEDLEY, G. W. & Price, W. R. (1948). Flora of Gloucestershire. Bristol. Rog, R. G. B. (1981). Flora of Somerset. Bridgewater. Ross-Craic, S. (1966). Drawings of British plants, 23. London. RosvaLL, A. (1979a). Snyltrotslaktet i Sverige. Svensk Bot. Tidskr. 73: 27-37. RosvaL_, A. (1979b). Orobanche alba, Timjansnyltrot pa Gotland. Svensk Bot. Tidskr. 73: 1-6. Rumsey, F. J. (1985). Orobanche caryophyllacea Sm. in North Wales. Watsonia 15: 277. Rumsey, F. J. (1986). Orobanche crenata Forskal. B.S.B.I. News 43: 18-19. Simpson, F. W. (1982). Simpson’s Flora of Suffolk. Ipswich. SITWELL, N. (1984). Shell guides to Britain’s threatened wildlife. London. Sowerby, J. (1794). English Botany, 3. London. Sowersy, J. (1797). English Botany, 6. London. Sowersy, J. (1799). English Botany, 8. London. Sowerby, J. (1807). English Botany, 25. London. Sowerby, J. (1866). English Botany, 3rd ed. (Ed. J.T.B. Syme). London. Strip, A. (1980). Wild flowers of Mount Olympus. Kifissia. Sutton, G. (1797). A description of five British species of Orobanche. Trans. Linn. Soc. Lond. 4: 173-188. Wess, D. A. (1985). What are the criteria for presuming native status? Watsonia 15: 231-236. Wess, D. A. & CHATER, A. O. (1972). Orobanche L., in Tutin, T. G. et al., eds. Flora Europaea 3: 286-293. Cambridge. (Accepted July 1990) ‘oran 7 a vi Teed he sgl, 25-50 cla _ - Pewecig athee dente. Mekete ; num, uaeoU ‘lly bidentate or entire, G arent, tf. «ap | Staregige ici cited 33 nin alee oe) ) eee. SrvGe “ESAT sa LS et) ET a Maries, deionkth & thatog is Vom aes ye init aie | Ur aD. | ; = idea: a phe meree irri! ear le “ay be 2. CTE 2 Lakota tt a a | ab seenplin Agee a ee 8 eenreT>/ Say nye tens onterrm .d vest lex’ | .oobao! .& peach ddignd Ray t oie! 2S erated AANA 4 . = nite? ofeene? 1) Jibs GS) bs iné ERIE, Adena 40085) .0 iti A «worw\) ino jo-eewah, Sb ce Bint » Se sir 1 entiti. wbopetesl> 5) ps3 fini eit? neaiiqixye df aanrat Oagiete avila ornare 3 ap} PDA! yah oa rel Ate: .* 40el wer eset {31 Crt 0) A #2TAH 2, ‘ai Wi vu, StiqsanA,) = s (as a t ¢) ni J at ms) cl s a . my ~ a 13 5 . F » Rae oi ' commucalty Wnptige x mA 7 g a F Lino, Watsonia, 18, 297-301 (1991) 297 A re-assessment of Pteridium aquilinum (L.) Kuhn in Britain F. J. RUMSEY, E. SHEFFIELD Department of Cell & Structural Biology, University of Manchester, Oxford Road, Manchester, M13 9PL and C. H. HAUFLER Department of Botany, University of Kansas, Lawrence, Kansas 66045, U.S.A. ABSTRACT Starch gel electrophoresis has been used to assess the recent recognition of Pteridium aquilinum (L.) Kuhn subsp. atlanticum C. N. Page and subsp. /atiusculum (Desv.) C. N. Page from Scotland. The former has been described as a new subspecies and the latter raises a variety to subspecific rank. Page’s subsp. atlanticum is isozymically indistinguishable from typical subsp. aquilinum. Scottish material referred to subsp. latiusculum, on the other hand, is additive for isozymic markers characteristic of North American /atiusculum and European aquilinum and may represent a hybrid between those taxa. Morphological intermediates between the hybrids and pure aquilinum, indicate that these plants may have crossed extensively with their neighbours. Preliminary isozymic studies of northern European plants which have been morphologically referred to North American latiusculum indicate that they may also be the product of past hybridisation. Our isozymic analyses and the DNA studies of others indicate that northern hemisphere Pteridium variants are closely related; the elevation of Tryon’s var. /atiusculum to subspecific rank therefore seems untenable. INTRODUCTION Tryon (1941) assigned all British bracken to a single variety, Pteridium aquilinum subsp. aquilinum var. aquilinum. Recent studies, however, have demonstrated considerable phenotypic plasticity (Page 1976, 1986) and genetic polymorphism (Wolf, Haufler & Sheffield 1987, 1988a,b; Sheffield, Wolf & Haufler 1989; Wolf, Sheffield & Haufler 1990). In 1989, Page used morphological features to diagnose two further subspecies of British bracken (subsp. atlanticum C. N. Page and subsp. latiusculum (Desv.) C. N. Page) and he hypothesised that hybridisation between these could have initiated the vigorous subsp. aquilinum. Given the sympatry of these three taxa, he suggested that introgression among them was commonplace, and was responsible for the spectrum of intergrading variability seen in British bracken today. The presence of more than one bracken taxon in this country and the suggestion of active evolution within the species are of considerable importance to our understanding of this economically important weed and may have a profound influence on the formulation of control strategies. The aim of this study was therefore to elucidate the status and identity of these taxa using starch gel electrophoresis of their protein constituents (isozymes). This is a well established technique for the study of genetic variability (e.g. Gottlieb 1981) and systematic relationships (Crawford 1983, 1985). Work with pteridophyte species has shown that isozymes can be particularly useful in distinguishing cryptic species (Paris, Wagner & Wagner 1989) and in reconstructing episodes of hybridisation and allopolyploidy during reticulate evolution (Werth 1989). Pteridium isozymes have been the subject of intensive study both in Britain and the USA, and isozyme markers which reliably distinguish taxa recognised on morphological grounds have been found for all those investigated to date (Wolf 1986; Wolf, Haufier & Sheffield 1988a). Two taxa have been studied particularly closely; subsp. aquilinum sensu stricto and subsp. Jatiusculum. Both vary at 298 F. J. RUMSEY, E. SHEFFIELD AND C. H. HAUFLER many of the 28 loci currently investigated, but two enzymes have alleles which are unique to one or the other variety: isocitrate dehydrogenase (IDH) and triosephosphate isomerase (TPI). MATERIAL AND METHODS Material was collected from the first recorded Scottish locality (Page 1988) of subsp. latiusculum in the company of C. N. Page and Y. C. Golding. Eight samples were taken from fronds at regular intervals across a patch c. 65m X 3m with scattered outliers. Isozyme analysis (see Sheffield, Wolf & Haufler 1989) demonstrated this to be a single genetic individual (clone). The woodland immediately behind the clone is of planted Larix and Betula and cannot be considered ‘native pinewood’ (cf. Page 1989). Immediately surrounding this clone are more extensive patches of a plant morphologically intermediate between it and subsp. aquilinum (which exists c. 30m away). Plants of this intermediate morphology have also been noted elsewhere in Scotland during intensive searches for more subsp. Jatiusculum material by C. N. Page (in litt.), FJR and others; many such samples were collected and isozymically analysed during the present study. Nine fronds of material collected at the locus classicus of subsp. atlanticum, and fertile material referred to subsp. /atiusculum, were collected by CNP and YCG and kindly made available for analysis. Gametophytes were raised from the spores of the latter; some were electrophoretically investigated, others were crossed with aquilinum gametophytes, and the resultant sporophytes analysed. Two Pteridium fronds were also collected from each of a woodland in Grantham, New Hampshire, U.S.A., and a conifer plantation on Men Island, Denmark, and electrophoretically analysed. Electrophoretic analysis of 28 loci followed the methods established for Pteridium by Wolf (1986). (See also Wolf, Haufler & Sheffield 1987, 1988 a & b; Sheffield et al. 1989; Sheffield, Wolf & Haufler 1989; Wolf, Sheffield & Haufler 1990). RESULTS AND DISCUSSION SHOULD PTERIDIUM AQUILINUM SUBSP. ATLANTICUM BE RECOGNISED AT SUBSPECIFIC RANK? Comprehensive isozymic analysis of material collected at the locus classicus of subsp. atlanticum revealed banding patterns consistent with the range of variation expressed by subsp. aquilinum in all samples. Although these enzymes represent only a tiny part of the genome, all Pteridium taxa previously investigated, which include those recognised at varietal level, have had some discrete and distinct banding pattern for at least one of the isozymes studied (Wolf 1986; Wolf, Haufler & Sheffield 1988a). We therefore conclude that recognition of subsp. atlanticum at subspecific level is not consistent with the other well defined Pteridium taxa. The basicolous nature of subsp. atlanticum has been strongly stressed, although without supporting evidence from soil analysis. Many authors (see Sheffield et al. (1989) for review) have reported P. aquilinum sensu stricto from base rich habitats. The morphology of subsp. atlanticum intergrades with that of subsp. aquilinum s.s. It has been suggested that such plants represent hybrids and introgressants between these taxa (Ruther- ford 1989). Given the acknowledged variability within the genus, we believe that final judgement on this question should be deferred until cultivation reveals whether the phenological and morphologi- cal characters described by Page (1989) are environmentally or genetically determined. Until such time, we consider that the electrophoretic evidence necessitates the reduction of subsp. atlanticum to synonymy with subsp. aquilinum var. aquilinum. IS PTERIDIUM AQUILINUM SUBSP. LATIUSCULUM A BRITISH PLANT? All of the material referred by Page (1989) to subsp. /atiusculum possessed banding patterns for the enzymes TPI and IDH that would be expected to result from an amalgamation of the isozymes previously found in subspp. /atiusculum and aquilinum (Fig.1). This strongly suggests that Scottish plants are hybrids of that parentage. Morphologically these plants more closely approach subsp. latiusculum but often lack marginal hairs to the pinnules and possess more or less ciliate indusia, characters considered diagnostic of subsp. aquilinum s.s. by Tryon (1941). I a a a PTERIDIUM AQUILINUM IN BRITAIN 299 Ficure 1. Starch gel stained for isocitrate dehydrogenase, a dimeric enzyme. Two left hand lanes, P. aquilinum sensu stricto from Manchester showing the most common single banded fast allele. Two central lanes, P. aquilinum (L.) Kuhn subsp. latiusculum (Desv.) C. N. Page from Scotland, showing the three banded heterozygote pattern that would result from a cross involving parents with the fast allele (lanes 1 & 2) and the slow allele (lanes 5 & 6). Two right hand lanes, P. aquilinum subsp. aquilinum var. latiusculum (Desv.) Underw. from New Hampshire, U.S.A. showing the most common single banded allele. Electrophoretic analysis of the gametophytes raised from these plants showed segregation of the marker alleles (Fig. 2), as would be predicted for a plant with a diploid genetic system with disomic inheritance patterns (Wolf, Haufler & Sheffield 1987). Half the gametophytes were, as expected, electrophoretically indistinguishable from subsp. aquilinum, the other half indistinguishable from subsp. latiusculum. Gametophytes of the putative hybrid are fertile, and proved fully inter-fertile with subsp. aquilinum. Crosses between gametophytes from the putative hybrid with the ‘latiusculum’ pattern and aquilinum gametophytes gave sporophytes with banding patterns indistinguishable from those of the Scottish plants (e.g. Fig. 2). Cross fertilisation of gametophytes bearing the ‘latiusculum’ markers would reconstitute a sporophyte electrophoretically indistinguish- able from subsp. latiusculum, as would self-fertilisation of these gametophytes. Interestingly, no such plants have been found in Scotland. As Pteridium is an outcrossing species (Wolf, Haufler & Sheffield 1988b), sporophytes are routinely derived by cross-fertilization. This explains the absence of sporophytes of selfed origin but the absence of Jatiusculum-type sporophytes can only be attributed to the far higher numbers of aquilinum than ‘latiusculum’ spores (which would make crosses between aquilinum and ‘latiuscu- lum’ far more likely than ‘latiusculum’ xX ‘latiusculum’). On morphological and phenological grounds we assume that some plants with the subsp. latiusculum markers represent introgressants to the subsp. aquilinum parent. Putative introgres- sants from other Scottish sites have not been analysed. In the absence of plants referred by us to subspp. /atiusculum X aquilinum they must be derived either from wind-borne hybrid spores, or be relics of previous subspp. latiusculum X aquilinum or latiusculum populations now extinct. ee ® FiGure 2. Starch gel stained for isocitrate dehydrogenase. Eight left hand lanes, single gametophytes grown from spores of P. aquilinum (L.) Kuhn subsp. latiusculum (Desv.) C. N. Page showing segregation of fast and slow alleles. Two right hand lanes, sporophytes grown from gametophytes taken from the same culture, crossed with gametophytes of P. aquilinum sensu stricto, showing a three banded heterozygote pattern. 300 F. J. RUMSEY, E. SHEFFIELD AND C. H. HAUFLER IS PTERIDIUM AQUILINUM S.S. A HYBRID OF SUBSPP. ATLANTICUM AND LATIUSCULUM? Although P. aquilinum sensu stricto shows features indicative of hybridity as stated by Page (1989) (and tentative support for this has been obtained from DNA analysis for some specimens (Thomson, in litt.)), the isozymic evidence mitigates against claims that subspp. latiusculum and atlanticum can be considered potential parents. CONCLUSIONS The discovery of an alien genome within British material is of considerable interest. Initial investigation of Danish material attributed to var. /atiusculum has revealed an isozyme pattern similar to that of Scottish material. European authors have shown greater reluctance than their American counterparts to distinguish var. /atiusculum from other taxa within subsp. aquilinum and we agree with Page (in litt.) that non-American material may not be conspecific with var. latiusculum (Desv.) Underw. From the limited electrophoretic evidence currently available we hypothesise that the boreal Eurasian taxon has been derived at some stage from the hybridisation of the now European var. aquilinum with the now American var. Jatiusculum and that Scottish material referred by Page (1989) to the latter is best viewed as hybrid stock. This may be a native component of the British flora, or could have resulted from more recent spore dispersal from elsewhere within Europe. Subsp. atlanticum C. N. Page must, on current evidence, be regarded as synonymous with subsp. aquilinum. The elevation of var. latiusculum (Desv.) Underw. to, and the creation of subsp. atlanticum at, subspecific rank is untenable within the currently accepted global view of the genus Pteridium (Tryon & Tryon 1982). Brownsey (1989) reported that hybrids between the subspecies recognised by Tryon (1941) give rise to sterile progeny (although fertile material has been obtained from one of the stands cited by Brownsey; see Tan & Thomson (1990)), whereas vars. latiusculum and aquilinum have proved to be fully interfertile. Recent chloroplast DNA analysis (Thomson, in litt.) emphasises the distinction between the essentially Southern hemisphere subsp. caudatum and the essentially Northern hemisphere subsp. aquilinum; these taxa are more distinct from one another than vars. /atiusculum and aquilinum. To reflect this, and preserve a sensible infraspecific hierarchy we prefer to consider subsp. latiusculum (Desv.) C. N. Page at varietal level (as in Tryon 1941). Evidence is accruing that Pteridium is probably best subdivided into several species, and resurrection of subsp. latiusculum (Desv.) C. N. Page may therefore be deemed appropriate at some time in the future, but until a global monograph is completed, the uneven taxonomic treatment which currently plagues the genus should, in our opinion, be avoided. ACKNOWLEDGMENTS We are grateful to C. N. Page and Y. C. Golding for their help, to S. Challinor for technical assistance, to K. Wolf, M. Wolf, P. Haufler, H. Haufler, H. Balsler, S. Churchill, C. N. Page and Y. C. Golding for field collections, and to the Nuffield Foundation and Royal Society for financial support. REFERENCES Brownsky, P. J. (1989). The taxonomy of bracken (Pteridium: Dennstaedtiaceae) in Australia. Aust. Syst. Bot. 2: 113-128. CrawrorD, D. J. (1983). Phylogenetic and systematic inferences from electrophoretic studies, in TANKSLEY, S. O. & Orton, T. J., eds. Isozymes in plant genetics and breeding, part A, pp. 257-287. Amsterdam. CrawrorD, D. J. (1985). Electrophoretic data and plant speciation. Syst. Bot. 10: 405-416. Gorr.ies, L. D. (1981). Electrophoretic evidence and plant populations. Prog. Phytochem. 7: 1-46. Pace, C. N. (1976). The taxonomy and phytogeography of bracken — a review. Bot. J. Linn. Soc. 73: 1-34. Pace, C. N. (1986). The strategies of bracken as a permanent ecological opportunist, in SMitH, R. T. & TAYLOR, J. A., eds. Bracken — ecology, land use and control technology, pp. 173-181. Lancaster. PTERIDIUM AQUILINUM IN BRITAIN 301 Pace, C. N. (1989). Three subspecies of bracken, Pteridium aquilinum (L.) Kuhn, in Britain. Watsonia 17: 429- 434. Paris, C., WAGNER, F. S. & WAGNER, W. H. (1989). Cryptic species, species delimitation and taxonomic practice in the homosporous ferns. Am. Fern J. 79: 46-54. RUTHERFORD, A. (1989). Atlantic bracken. B.S.B.I. News 52: 17-18. SHEFFIELD, E., WoLF, P. G. & HAUFLER, C. H. (1989). How big is a bracken plant? Weed Res. 29: 455—460. SHEFFIELD, E., WoLF, P. G., HAUFLER, C. H., RANKER, T. & JERMy, A. C. (1989). A re-evaluation of plants referred to as Pteridium herediae (Colmeiro) Love & Kjellqvist. Bot. J. Linn. Soc. 99: 377-386. TAN, M. K. & THomson, J. A. (1990). Variation of genome size in Pteridium, in THomson, J. A. & Situ, R. T., eds. Bracken: biology and management, pp. 95-103. Sydney. Tryon, R. M. (1941). A revision of the genus Pteridium. Rhodora 43: 1-31, 37-67. Tryon, R. M. & Tryon, A. F. (1982). Ferns and allied plants. New York. WERTH, C. R. (1989). The use of isozyme data for inferring ancestry of polyploid pteridophytes. Biochem. Syst. Ecol. 17: 117-130. Wo LF, P. G. (1986). Electrophoretic evidence for diploidy and outcrossing in the genus Pteridium (bracken fern). M.Sc. thesis, University of Kansas. Wo Fr, P. G., HAuFLER, C. H. & SHEFFIELD, E. (1987). Electrophoretic evidence for genetic diploidy in the bracken fern (Pteridium aquilinum). Science 236: 947-949. Wo tr, P. G., HAUFLER, C. H. & SHEFFIELD, E. (1988a). Maintenance of genetic variation in the clonal weed Pteridium aquilinum (bracken). SAAS Bull. Biochem. Biotech. 1: 46-50. Wo LF, P. G., HAuFLER, C. H. & SHEFFIELD, E. (1988b). Electrophoretic variation and mating system of the clonal weed Pteridium aquilinum (L.) Kuhn (bracken). Evolution 42: 1350-1355. Wo tr, P. G., SHEFFIELD, E. & HAuFLER, C. H. (1990). Estimates of gene flow, genetic substructure and population heterogeneity in bracken (Pteridium aquilinum). Biol. J. Linn. Soc. (In press). (Accepted July 1990) Nh hyo ntietintl oi a to it ‘ra afl Orr ranagt (nod etazive isa for rm f % 7 Deve > to ait oi a vivelt ee eae ae BRE-STE RE cud nhl Pater te viele Re GSO T Agim? 4 AL noamon'l a, suds a | seb sect 80 rave | teat EB aches. be ‘Seal HNO juclspaliienioeecean iy si Selb ihe i eee cies Aver te: 7 AT ee We ae ot oe , Ant Bodsis Lio) Whodtiive atelier at Th hier | othees Cato, Mote St SRR ome 2 eae Bigangetyets mrt qin satan tity ie! rere ered) OP Aches soe ty rie % se ead 1 Ape? Si Serato i), Musab ansh, ee Sie Oh PRY LOREM ONG quae Tye" ty otis tors bARAAT i eyequeranter®: ak Bho heh — Teh Aas “ae Sn wwitainy S om bye eo powery Aa pay. ay er a ee ee Heb el - (oz g ite WAj | f aye} ‘oi aay PE isviga } ear wa *' sess hie Ciosagorsied be is he ny AU Ft, onli i tiie: t oktakion en athe, atten a eS Bae ; a cot pte statal view of the nein, § AY taceseied Cut brytortals net wi eo thes ~bspecies 466 is gett lan | a et yf ru ae tah eet i Tea -obtened fmt oy BY fel + O00) ), Wwoeneis vieS. lone . wn dor iteieeth... Aco chioroniast reece ath : Hes nell Cou becnunliie =e cada py ne ore tore distin? Frome aia Ne.’ wy conten A suet es tt iraspreitie Ba .. $2 a av) La Tyee ta ot WERK ve ht) Several: SPRRiOs, 9 mew ther: eo ti eswnal aero DAY 4 iW 7 it i Conmeted, et Uae FREOW rr Wi oe On, OG a | nae te vit ah hs fi. I : ‘4 bie) ©. e nu : le | y *, { hal] at ray o™ " : nrchlil OB ae ad Love Soutety: orn py faateiale ~\ oy Aiteteiva je yi | ea, doernoetorredse: atubdien jit 1a “# rt ely y Ne at 1 yay ve Aowbeseaga) iM oes ; ue et, $A: 405416, : pho vou. Phytases Fhe, a shy ann : revs Hat: 3 Late Sod ‘be ue ép Moa tutal ovisbaniat, of Mauphe, FED, a pile pe. De TR, of ae. ree) Watsonia, 18, 303-305 (1991) 303 The status of Populus nigra L. in the Republic of Ireland D. D. HOBSON St John’s College, Oxford, OX1 3JP ABSTRACT The presence of the Black Poplar Populus nigra L. in Ireland is confirmed and it is shown to be locally common in rural habitats in the Midlands and along the Shannon river-system. It is suggested that P. nigra is an Irish native and evidence supporting this is put forward. INTRODUCTION The results of the B.S.B.I. Survey of the distribution of Populus nigra L., the native Black Poplar, in England and Wales have recently been published by Milne-Redhead (1990). I have been interested in the Black Poplar since E.M.-R. showed me some when I was staying in Suffolk in 1977, and I started to find them near my home in Dorset. I have given the Survey records from many other places in England and rediscovered the tree on Putney Heath (v.c. 17), recorded by J. Fraser in 1924 but missed by local botanists! I have also planted out near my home in Dorset 13 seedling Black Poplars raised by E.M.-R. in 1977 from seed gathered in Cheshire, and observed their not inconsiderable genetic variation. I am, therefore, thoroughly familiar with the tree. So, when E.M.-R. asked me to undertake a reconaissance of P. nigra in Ireland, I willingly agreed. Very little was known about the Irish status of this tree. A few records had come in from the Survey, including two from Co. Kildare and another from Co. Cork, but not enough to suggest a natural distribution, and none from Northern Ireland, so the map (Milne-Redhead 1990) was left blank. Populus nigra was not considered a native tree in Ireland by Colgan & Scully (1898). Praeger (1934), in his account of Irish vegetation, mentions that ‘poplar’ occurs in some areas but does not say which species. Black Poplar seemed to be most likely. But the view held by Irish botanists has been that it is a very rare, introduced tree, planted from English stock by Anglo-Irish settlers. OCCURRENCE IN IRELAND A circuit was planned round the Republic of Ireland south and west of Dublin. It was felt that, because of the limited time available, it would not be worthwhile to explore farther north, as the tree has a southern distribution in Britain. It seemed that the tree was most likely to be found in the low- lying fertile river-basins of the Irish Midlands, but I also had to investigate elsewhere and check several possible records. My route took me from Co. Wexford, north up the Barrow and Slaney rivers and up to Dublin. I also looked at specimens named P. nigra in the herbaria at DBN and TCD. Some of the specimens were obviously hybrids, but others were very clearly correctly named. From Dublin, I travelled west to the Bog of Allen, an area of wet loamy farmland and vast lowland peat-bogs which covers much of Co. Kildare and Co. Offaly around the headwaters of the River Liffey. P. nigra proved to be locally common across a wide area between Edenderry and Clane, and I collected 32 records. I then moved to the Shannon river-system and based myself at Portumna on the northern tip of Lough Derg. At Ballyshrule Bridge, just to the west of Portumna, I found P. nigra to be the commonest tree, but of an unusual, unbossed clone. E.M.-R. tells me that he knows of only one unbossed clone in England. However, in all other respects these trees conformed to the typical P. nigra. In the close vicinity of the bridge, 80 trees were recorded. Around the lake, a further 77 trees were found, with a particularly impressive group of 17 at Ballinderry on the eastern side. 304 D. D. HOBSON Ficure 1. Distribution of Populus nigra L. in Ireland. South of Limerick, the banks of the River Maigue yielded a further eight trees, but in the Blackwater and Suir river-valleys farther south, only another four trees were found. A map of the known distribution in Ireland is given in Fig. 1. POPULUS NIGRA IN IRELAND 305 Contrary to existing opinion that all Irish P. nigra have been planted from introduced stock, only a few of these trees are associated with farmyards and farmhouses. Most of them are growing in hedges in wet farmland, often with deep, water-filled ditches draining the fields. The trees were never found in peat-bogs but always in the surrounding loamy soils. Moreover, the trees were found almost exclusively in the river-valleys and floodplains, which E.M.-R. considers to be their natural habitat in England and Wales, and rarely on hillsides. As P. nigra has been considered an introduction to Ireland, it seemed probable that one method of entry of the tree into the Irish countryside might have been during the building of the extensive canal system across Ireland in the late 18th century, during the era of British domination. In Co. Kildare, I searched along a stretch of the Grand Canal south-west of Clane and discovered that the only poplars were old P. X euramericana hybrids. It appears that, as early as 1794 when this stretch was built, P. nigra had been replaced with cultivated hybrid trees. This seems to indicate that any introduction of P. nigra must have occurred before the 18th century. Furthermore, I found Black Poplars in the hedges of the pre-existing field-system, through which the canal had been built. Older planned tree-planting would have occurred on the extensive parkland that grew up in Ireland. However, only one of the trees found could be the result of such a planting — the vast old tree near Cahir. It seems reasonable to conclude that Populus nigra is a native tree of the Irish Midlands. It is not a woodland tree, so has had to survive amidst agriculture. It is almost exclusively propagated from cuttings so, as in England and Wales, relies on man’s assistance. Even now it is still being planted by Irish farmers; several trees found were no more than ten years old. In Co. Galway the timber is used for joists and floorboards. I am inclined to consider the tree native because it does not appear necessarily to be associated with settlements and its distribution is very tightly linked with river- valleys and flood-plains — too closely to be haphazard introduction. The local density and distribution of the tree in Kildare and around Lough Derg is such that, if indeed the tree has been introduced, it must be an ancient introduction. But why then is this ancient introduction not concentrated along the eastern coast — the easiest point of access for English settlers and the area of densest English settlement? It seems necessary for further research to be conducted into whether the tree is indeed native; possibly pollen analysis could provide evidence. [ think it is also necessary to build up a complete picture of the distribution of the tree across Ireland; for instance, north up the Shannon river-system from Lough Derg to Lough Allen. I hope local B.S.B.I. members will carry on where I left off and not rely on visiting botanists from England or Wales. Perhaps someone will discover it in Northern Ireland. In all, I discovered 210 trees and it appears that Ireland is a very important station for this rare and unusual species. I am confident that the status there of this tree of ‘rugged grandeur” (Rackham 1986), is, at the moment, secure. ACKNOWLEDGMENTS The work was funded by a grant from the Welch Bequest Fund, administered by the B.S.B.I., to whom I am most grateful. I am deeply indebted to E. Milne-Redhead for his constant help and encouragement over many years and to his daughter, Annette Harley, for typing the manuscript. For help with the map I am grateful to C. D. Preston of the Biological Records Centre, Monks Wood. Thanks are also due to S. C. Holland for generously passing on her Irish records. Finally, I wish to thank the staff of the Botany Department and Herbarium at Trinity College, Dublin, and the many other people in Ireland who volunteered information. REFERENCES Coiaan, N. & ScuLty, R. W. (1898). Cybele Hibernica, 2nd ed., p. 509. Dublin. MILNE-REDHEAD, E. (1990). The B.S.B.I. Black Poplar survey. Watsonia 18: 1-5. PRAEGER, R. L. (1934). The botanist in Ireland, para. 239. Dublin. RackHaM, O. (1986). The history of the countryside, p. 207. London. (Accepted May 1990) tm bsosbosin: aot betnslg abed s¥as al K.. 4 ehh Ne 4 vii nokmige rreitee aed ‘eo oe | a en my ie es uh i” ' CMA ri WHO fib, 0408 ray di & A a brs ; abrusigarr ise win ‘| A bot aac L, A 7 . a ; iy ey " My » AUBIN DD > VSR ee ssf diced Rome. pln ‘¢ Hi etissted esse toed) levnwil a “aos i ee “sy ’ ; 40 POT LOS ) BV ANT 14 tev Tee i 1s phil 4 i i it q r i bi ¥ iu af i ; 7 eh ’ Mm) ite “7 7 . 4 o) t ; tt ) i % : ' t t i j i a a 4 * IE i 710" ‘4 “ad i ’ ' eT * Fe ~ Watsonia, 18, 307-309 (1991) 307 Two new varieties of British Dactylorhiza D. M. T. ETTLINGER Royden Cottage, Cliftonville, Dorking, Surrey, RH4 2]F ABSTRACT Dactylorhiza fuchsii var. rhodochila D. M. T. Ettlinger, var. nov., and D. majalis subsp. purpurella var. atrata A. J. Richards, var. nov., are distinguished from typical specimens chiefly by their blotch-marked labella. INTRODUCTION The labellum of many species of European Orchidaceae has small purple dot or dash markings on a paler background, these being very occasionally replaced by a solid central or overall reddish-purple blotch. In most species this variation is rare and occurs only in isolated individuals, e.g. in Corallorhiza trifida Chatel., Orchis anatolica Boiss., O. morio L. and O. coriophora L. In the genus Dactylorhiza it seems to be more frequent, the best-known example being D. insularis (Sommier) Landwehr var. bartonii Huxley & Hunt. This blotch marking on the labellum may be accompanied by heavier-than-average leaf spotting (e.g. in D. fuchsii), heavy leaf spotting where little or none is usually present (e.g. in D. majalis subsp. purpurella), and sometimes even overall suffusion of the leaves with anthocyanins (both). However, no change in leaf marking is seen in D. insularis var. bartonii or in the blotch-marked variants in Orchis, so blotch marking of the labellum cannot be simply an effect of heavy anthocyanin presence in the plant as a whole: some mutation in the genes that code for labellum marking seems likely to be the cause. That this sort of variation may have evolutionary potential is indicated by the pollination advantage gained by having two colour morphs in D. sambucina (L.) So6 (Nilsson 1980). This species offers no reward to its bumblebee pollinators (Bombus spp.) and attracts them by deceit; the presence of the red var. rubra Winterl., in addition to the commoner yellow morph, presents an additional search pattern and extends the period before individual pollinators learn to avoid the species as a whole. For D. fuchsii a similar picture of deceit is clouded by the finding (Dafni & Woodell 1986) that unrewarded visits by naive bumblebees were supplemented by visits from honeybees (Apis mellifera), which were to some extent rewarded by a stigmatic exudate. Nevertheless, I believe that Dafni & Woodell’s study area (with which I am familiar) carried an atypically high honeybee population and there must be many D. fuchsii sites, e.g. in dune systems and on open downland, where honeybee presence is negligible. No similar work has been published on the pollination ecology of D. majalis subsp. purpurella but it is a reasonable assumption, in view of the high incidence of hybrids between the two where they coexist, that it is similar to that of D. fuchsii. It therefore seems worthwhile to distinguish by name those examples of the blotch variation which at least occur regularly. Populations with these variants also contain typical specimens and intermediates do occur. No form of isolation seems to be involved and the rank of variety therefore seems to me to be appropriate. Bateman & Denholm (1989) referred to three, possibly four, British dactylorchids in which a blotched variant occurs: Dactylorhiza maculata (L.) So6 subsp. ericetorum (E. F. Linton) Hunt & Summerhayes, D. fuchsii (Druce) So6, D. majalis (Reichenb.) Hunt & Summerhayes subsp. purpurella (T. & T. A. Steph.) D. M. Moore & So6, and possibly a single specimen of D. incarnata (L.) Sod. They mentioned without comment the already-named blotch-marked D. maculata var. concolor Vermeulen and they refrained from giving names to the others. The analogous variety in D. fuchsii appears to be commoner in the British Isles than D. maculata 308 D. M. T. ETTLINGER var. concolor, and Bateman & Denholm (1989) listed six British sites, including one of their own study areas where it formed c. 20% of the D. fuchsii population. To these should now be added a drain bank near Belton, N. Lincs. (Weston 1979), alkaline grassland at Blackhall Rocks, Co. Durham (M. Bradshaw 1967 per M. R. Lowe, pers. comm. 1985), a second limestone meadow in Derbys. (P. M. Torry, pers. comm. 1989), a wood on Wealden Clay near Warnham, W. Sussex (D. C. Lang, pers. comm. 1990) and the edge of a chalk beechwood near Duncton, W. Sussex (J. M. Scott per D. C. Lang, pers. comm. 1990). There can be little doubt that more sites will come to light once the taxon is formally recognised. The blotch-marked variety of D. majalis subsp. purpurella is only known at present from one site near Hartlepool (Bateman & Denholm 1989), where a population of over 100 specimens was found in 1978 by A. J. Richards and has persisted since: plants approaching its description have been seen near Kilmore, Co\Wexford, and near Hornhead, W. Donegal, though these need confirmation (R. Piper, pers. comm. 1989). Richards proposed the epithet atrata for this taxon but did not actually publish it. However, it has already become widely known on a hearsay basis and has appeared in print as a nom. illegit. (Graham 1988). To rectify this situation, A. J. Richards has kindly supplied the diagnosis of var. atrata below. In both these varieties the chief distinction lies in labellum colour. Since colour cannot at present be preserved in herbaria, the obligatory holotype specimens have been supplemented by additional material in the form of photographic prints on Cibachrome made from colour transparencies; these should have a useful life in herbarium conditions of 50—100 years. THE VARIETIES Dactylorhiza fuchsii (Druce) So6 var. rhodochila D. M. T. Ettlinger, var. nov. Ho orypus: Beacon Hill, GR 41/608.226, N. Hants, v.c. 12, chalk grassland, 11 June 1990, D. M. T. Ettlinger (K). Additional photographic material from the same site and from near Buxton, GR 43/ 09.75, Derbys, v.c. 57, limestone grassland (photographs by D. M. T. Ettlinger, June 1989 and June 1984 respectively) is with the holotype in K, copies in BM. A D. fuchsii differt sequentibus: folia subdensiore punctata, labellum signatum macula rhodopur- purea ad centrum cum margine pallidiore, vice punctibus linulibusque. Speciminibus extremibus pagines superiores foliarum omnino a purpureo tectae, et totum labellum atropurpureum sine margine pallida. It differs from typical D. fuchsii as follows: leaves rather more heavily spotted, the labellum marked with a broad central reddish-purple area with paler edges in place of dots and small lines. In extreme examples the leaves are suffused on the upper surfaces with purple and the whole labellum is a rich dark purple without a pale margin. Dactylorhiza majalis (Reichenb.) P. F. Hunt & Summerhayes subsp. purpurella (T. & T.A. Steph.) D. Moresby Moore & So6 var. atrata A. J. Richards, var. nov. Hotortypus: Locality (details withheld) in Co. Durham, v.c. 66, dune grassland, 13 July 1979, A. J. Richards (BM). Additional photographic material from the same site (photographs by D. M. T. Ettlinger, leg. A. J. Richards, 11 July 1985) is with the holotype in BM, copies in K. A D. majalis subsp. purpurella differt sequentibus: folia grosse atromaculata, in pagine superiore maculis rotundis solidis ultra 5 mm in diametro, in pagine inferiore regulariter punctata ad margine, punctae circa 1 mm in diametro. Labellum saepe toto suffusum atropurpureum sine maculis vel linebus, aliquando cum margine pallidiore. It differs from typical subsp. purpurella as follows: leaves heavily and darkly spotted, on the upper surface with solid round spots of over 5 mm diameter, on the lower surface regularly marked at the TWO NEW VARIETIES OF DACTYLORHIZA 309 edges with spots of c. 1 mm diameter. Labellum usually entirely suffused with dark purple without spots or lines, sometimes with a paler edge. ACKNOWLEDGMENTS I am grateful to I. Denholm, P. F. Hunt, M. R. Lowe and F. Rose for advice and comments on parts or all of the manuscript; to the Nature Conservancy Council for permission to take the holotype of var. rhodochila from a National Nature Reserve; and particularly to A. J. Richards for advice and permission to include his var. atrata in this paper. REFERENCES BaTEMAN, R. M. & DENHOLM, I. (1989). A reappraisal of the British and Irish dactylorchids, 3. The Spotted- orchids. Watsonia 17: 319-349. Darni, A. & WOooDELL, S. R. J. (1986). Stigmatic exudate and the pollination of Dactylorhiza fuchsii. Flora (Jena) 178: 343-350. GraHaM, G. G. (1988). The flora and vegetation of County Durham. Durham. Nitsson, L. A. (1980). The pollination ecology of Dactylorhiza sambucina. Bot. Notiser 133: 367-385. Weston, I. (1979). A variant of Dactylorhiza fuchsii (Druce) So6 in N. Lincs. Watsonia 12: 399-400. (Accepted August 1990) --, 7? © Laas 7 fetucdy antes vitae Some’ c, 2% drain bank vcur Belton, N. reg Derhecté Rradshaw 1967 per Mi. pte mai Derorval®. Tocty, pork Conmpha, TSRR Sac t Laoag, pcs. Comat Ley poet s¢ — sme Ds inincabqresiebarat Rereransoscat greets eee ulz nt ; : hrmadiits ictaiedsts vet pe fer ae) Pei * teri 4! /Hatewion & Deshaky “ry) ncrentt benwee Coes ees shure fr - | Sucturds oot hes povelienl wiaer- Mien eporcudnng its desctiption haven ne ’ Weakness ond nter Meee, 0 Oeemen. hough these need coniicns hy cod pe aiwrw “oe le mi et Gd pot actually pablish it. Ho av aricig?): Fisgal a einshett ve cn aha nace eds (RE preligmecnaie eric: < : kas rigdlpVietledths “18 ath Ben we hasbersa he cenpilices sabes ious Saeargie Ave) 1.2 -s18000W a — AD = THE ciel pee Lo ro | Real ares. nT bo aohmangy | pee era Bis ar Eerie tae nti at sues wi? titans 'y, Tt it ¥20lties BAe Mi oa AGH ; Cl alee yo cee oT eit eset} 3 tala G i - CUT q . L mocdin odopiay f ir hes exrreniiram Hon SLOp iran atten aaiNes "Sy ! ; ne Dei ‘TO pee f dots ated seal) ines. Ing Lute Torr es . slant, 4 Jule 1 7 7 fomotographs by Ed ' Bel Yuen on BA, 4 Bled ay ‘t) pAgine et} punctate ae ae MPurcm MAO @ oe ; ter Liv epottedd, cai tng c regularly markexd ier s WY Watsonia, 18, 311-313 (1991) 311 Two new species of Cotoneaster K. E. FLINCK Villa Magnolia, Quai Alfred Chatelanat 10, 1820 Veytaux, Vaud, Switzerland and B. HYLMO Bygatan 30, S-267 00, Bjuv, Sweden ABSTRACT Two new species of Cotoneaster are described: C. atropurpureus Flinck & Hylm6, sp. nov. and C. hjelmqvistii Flinck & Hylm6, sp. nov. Both are common in cultivation and are sometimes found naturalised in the British Isles. C. ATROPURPUREUS FLINCK & HYLMO C. atropurpureus is a native of western China, but it is frequently planted elsewhere. In private gardens in the U.S.A. shrubs cultivated as C. horizontalis are mainly C. atropurpureus. In Europe C. atropurpureus is common in cultivation as C. horizontalis var. prostratus Hort. or as Cotoneaster cv. Prostratus Vilmorin (non Baker) (Grootendorst 1946). It occasionally naturalises in the British Isles (e.g. at Cambuslang, Lanarks., v.c. 77). The holotype of C. atropurpureus was identified by Rehder & Wilson (1912) as C. horizontalis var. perpusillus Schneider (= C. perpusillus (Schneider) Flinck & Hylm6). In the Botanic Gardens of Edinburgh and Glasnevin, Dublin, there are still shrubs of C. atropurpureus labelled ‘‘C. horizontalis var. perpusillus, Wilson 496” — the name and number having been carefully kept for 80 years. In AAH there are several sheets of C. atropurpureus cultivated in the Arnold Arboretum as Wilson 496 (the earliest from 1911). Cotoneaster atropurpureus Flinck & Hylm6, sp. nov. Ho.otypus: China, western Hupeh (=Hubei), Ichang, altitude 1300-2000 m, bare rocky ground, prostrate, fruits red, October 1907, Wilson 496 (A). PARATypPuS: Cultus Bjuv, Scania, Sweden, 5 June 1967, Hylm6 9703 (LD). Ex affinitate C. horizontalis Decne., a quo differt altitudine majore, habitu erectiore, foliis obovatis, tenuibus, undulatis, petalis basi atropurpureis. Frutex 0-5-1 m altus, ascendens ad decumbens. Ramis primariis arcuatis; ramuli juniores distichi, initio dense luteo/flavo-villosi; ramuli annotini cinereo-fulvi, persistenter flavo-villosi. Folia praecociter decidua, 9-14 x 8-12 mm, tenuia, undulata, obovata-orbicularia, obtusa vel truncata, mucronulata, basi obtusa ad acuta; supra, clare ad obscure viridia, in gemma sparse pilosa, explicata glabra, levia; subtus pallide viridia, pilis sparsis luteis nitidis, nervis 2-3 utrinque; petioli 1-2 mm, luteo/flavo-villosi. Cymae (1-)3 florae. Pedicelli 0-2(-1) mm, pilosi. Bracteae 2 mm, subulatae, glabrae, margine villoso, rubro-violaceae. Flores erecti, 6 x 3-4 mm. Receptaculum sparse flavopilosum. Lobi calycis erecti, 1-5-2 x 1-1-5 mm, deltoidei, acuminati, sparsi pilosi, virides vel rubropurpurei, margine luteo/ flavo-villoso. Petala erecta, 3-5 x 2-5 mm, inflexa, pandurata, ungui extracto, apice sub-erosa, basi atropurpurea, margine obscure rubro. Stamina 10, erecta vel incurvata, 2-3 mm; antherae albae, margine rosaceis; filamentis late subulatis, obscure rubris. Styli 2-3. Fructus 8 x 5-6 mm, obovatus, 312 K. E. FLINCK AND B. HYLMO ruber-vermicularis hollandicus, apicem versus sparse pilosus; lobis calycis oblique erectis; pyrenae 2-3. C. atropurpureus belongs to section Cotoneaster, subsection Adpressi Hurusawa, series Adpressi Flinck & Hylm6. It is related to C. horizontalis Decne, but the leaves are larger, thinner, obovate, undulate, and fall earlier; and the petals are darker and have a purplish-black base. C. atropurpureus is tetraploid, with 68 chromosomes (as determined by Zeilinga (1964) as C. horizontalis var. perpusilla 13077 (LD), and by Hensen (1966) as C. atropurpureus). Its characteris- tics are as follows. Shrub; decumbent to ascending, 0-5-1 m (or, if supported, to 3 m). Branches arched; young branches irregularly distichous, initially densely yellow-villous; 1-year-old branches greyish-brown, persistently yellow-villous, Leaves deciduous; lamina 9-14 x 8-12 mm, papery, slightly undulate, obovate-orbicular, apex obtuse to truncate, mucronulate, base obtuse to acute; upper surface mid/ dark-green, sparsely pilose initially but becoming glabrous, veins 2-3 pairs & not impressed; lower surface pale-green, subglabrous (the hairs shining golden-yellow); petiole 1-2 mm, yellow-villous. Inflorescence a (1—)3—-flowered cyme. Pedicels 0-2(-1) mm, pilose. Flowers erect, 6 X 34 mm. Receptacle sparsely yellow-pilose. Calyx-lobes erect, 1-5-2 x 1-1-5 mm, deltoid, acuminate, sparsely pilose, green or reddish-purple, margin yellow-villose. Petals erect & incurved (leaving no, or only a small, opening), 3-5 x 2-5 mm, fiddle-shaped, small drawn-out claw at base, apex irregularly denticulate, dark-red with purplish-black base and very narrow white marginal band. Stamens 10, erect or incurved, 2-3 mm; anthers white with margins tinged pale pink; filaments broadly subulate, dark purplish-red but white at apex (below anther). Styles 2-3. Fruits 8 x 5—6 mm, obovoid, orange-red (RHS 40A), glabrous except for sparsely pilose apex; calyx-lobes obliquely erect; nutlets 2-3. C. HJELMQVISTII FLINCK & HYLMO The taxon described in this section has been common in cultivation for more than 50 years, but various names have been used for it. An early record is a sheet in AAH of material cultivated in the Arnold Arboretum in 1940 as C. racemiflorus (Desf.) K. Koch cv. Fontanesii (Vilmorin 353-1936). It was initially distributed by nurseries under that appellation, and then under the wrong name C. horizontalis Decne. var. wilsonii Havemeyer ex Wilson (= C. ascendens Flinck & Hylm6). Subsequently it has also been sold as C. horizontalis cv. Coralle and cv. Robusta. This taxon has frequently been planted in Europe, and is also cultivated in, for example, Argentina, Chile, China, and the U.S.A. It is not uncommon in cultivation in the British Isles, where it may naturalise. Its wild origin is unknown, but it is likely to be a plant of western China (where related species are native). It appears to be a wholly apomictic species, and breeds true from seed. Cotoneaster hjelmqvistii Flinck & Hylm6, sp. nov. Ho tortypus: Cultus Hortus Botanicus Alnarp, Sweden, 4 June 1959, Hylm6 9300 (LD). PARATYPUS: Cultus Hortus Botanicus Alnarp, Sweden, 14 October 1958, Hylmo6 9300 (LD). Ex affinitate C. horizontalis Decne, a quo distat habitu erectiore, altitudine majore, omnibus partibus majoribus, foliis majoribus orbicularibus tenuioribus praecociter deciduis, calyce glabro. Frutex 1-1-5 m altus, ascendens ad decumbens. Rami primarii arcuati; ramuli juniores stricte distichi, initio dense luteo- ad flavo-strigillosi; ramuli annotini badii, persistenter flavo-villosi. Folia decidua, 13-20(—25) « 10-18(—25) mm, tenuia, plana, orbicularia ad rotunde obovata (in ramulis floriferis obovata), obtusa ad apiculata mucrone pusillo, basi obtusa; supra nitentia, clare viridia (in autumno intense rubropurpurea), glabra, superficie levi, nervis 3-4(—5S); subtus pallide viridia, sparse persistenter flavo-pilosa, ad nervos densius pilosa. Petioli 2-3 mm, pilosi. Cymae (1-)3(-4) florae. Pedicelli 0-5—2(—3) mm, glabri vel sparse pilosi. Bracteae 2 mm, subulatae, glabrae, margine villoso, rubro-violaceae. Flores erecti, 5 x 3-4 mm. Receptaculo glabro. Lobi calycis erecti, 2 x 1-5 mm, deltoidei, acuminati, glabri, virides vel rubropurpurei, margine luteo-villoso. Petala erecta, 4 = 2 bs te 6 ode, - Wiebe eon ee TWO NEW SPECIES OF COTONEASTER 313 x 4mm, leviter incurvata, orbicularia ad obovata, ungui parum conspicuo, apice sub-erosa, rubra ad obscure rosacea, margine rosaceo. Stamina 11—14(—16), erecta, 2-3 mm; antherae albae; filamentis late subulatis, rosaceis. Styli (1-)2(-3). Fructus 8 x 8 mm, orbicularis ad obovatus, luteus, glaber; lobis calycis oblique erectis; pyrenae 2-3. C. hjelmqvistii belongs to section Cotoneaster, subsection Adpressi Hurusawa, series Adpressi Flinck & Hylm6. It is related to C. horizontalis, but has a taller, more erect habit, is larger in all parts (the leaves considerably so), has leaves that are also thinner and fall earlier, and has a glabrous calyx. Its characteristics are as follows. Shrub; decumbent or ascending, to 0-5-1 m (or, if supported, to 7 m). Branching distichous; branches curved at tips when young but becoming straight, persistently villous (initially dense, yellow to yellowish-white); 1-year-old branches brownish-red. Leaves deciduous; lamina 13—20(- 25) X 10-18(—25) mm, thin, flat (or becoming concave), orbicular to broadly ovate (always ovate on fertile shoots), apex obtuse to mucronate, base obtuse; upper surface shining pure-green (becoming intense red-purple in autumn), glabrous, veins 3-4(—5) and not impressed; lower surface pale-green, persistently yellowish-white pilose; petiole 2-3 mm, pilose. Inflorescence a (1—)3(-4)-flowered cyme. Pedicels 0-5—2(—3) mm, glabrous or sparsely pilose. Flowers erect, 5 xX 3-4 mm. Receptacle glabrous. Calyx-lobes erect, 2 x 1-5 mm, deltoid, acuminate, glabrous, green or reddish-purple, margin villose. Petals erect but slightly incurved (leaving centre open), c. 4 X 4 mm, orbicular to obovate with small claw at base, apex irregularly toothed, red to dark-rose with rose margin. Stamens 11—14(—16), erect, 2-3 mm; anthers white; filaments broadly subulate, rose. Styles (1—)2(- 3). Fruits 8 X 8 mm, globose to obovoid, orange-red (RHS 40A), glabrous; calyx obliquely erect; nutlets 2-3. ACKNOWLEDGMENTS The authors are grateful to J. Fryer, B. A. Gale and C. A. Stace for assistance in preparing this paper for publication. REFERENCES GrooTenporst, H. J. (1946). Het sortiment Cotoneaster. Jaarboek de proeftuin te Boskoop, p. 56. HENSEN, K. J. W. (1966). Het geslacht Cotoneaster. Dendroflora 3: 17-19. REHDER, A. & WiLson, E. H. (1912). Rosaceae. Plantae Wilsonianae, p. 155. Cambridge, Mass. ZEILINGA, A. E. (1964). Polyploidy in Cotoneaster. Bot. Notiser 117: 262-278. (Accepted June 1990) ¢ 7 7 on low Sileas wv ne ei aoe ee me fete KARA areca stil nepiergoe OLE sain t se AT18 > f ‘ 7 = fn meto .ar x % ontaur pin 3 » arctins 4 he i geen | pay ¢ ' 7 c os J ' ? it vr ~ Sif gy ; ' ’ es BST ae aw ati pat 31 Cay 12 apn. 2 Sty sO ap , marlin oben Ma ities) eed ss ek bsealey ait aie th du: we egy : = AL oF a PEEA, Oy ABET thi nner act Miia } 2G aha oe “id 5 sian, vi roll? 2 28 Ps ran nth zaew - ¥ (¢ Reaves MeaeleyA ri meer ‘a — nt Watsonia, 18, 315-321 (1991) 315 Short Notes A NEW SUBSPECIES OF FESTUCA RUBRA L. Work carried out in this laboratory over the past 15 years has resulted in a new classification of the Festuca rubra L. group. The purpose of this note is to describe the only new taxon recognised. Festuca rubra L. subsp. scotica S. Cunn. ex Al-Bermani, subspecies nova Ho.otypus: Ardlamont Point, near Tighnabruaich, Argyllshire, v.c. 98, Scotland. H. McAllister s.n. Grown in University Botanic Garden, Leicester, 1983. C. A. Stace F157 (LTR). Planta rhizomatosa; culmi sparsim fasciculosi usque ad 70 cm alti; panicula densa ramis strictis; spiculae (ad apicem quarti flosculi) (8) 9-4—-11-8 (12-5) mm longae; lemmata basalia (5-5) 6-1—7-9 (9) mm longa cum arista (0-3)1-2—2-3(3-7) mm longa; folia innovationum involuta, fasciculis scleren- chymae parvis discretis; chromosomatum numerus 2n=56. This subspecies is distinguished from most taxa within the F. rubra group by its long spikelets, rather long lemmas, and long awns. F. rubra var. commutata Gaudin (F. nigrescens Lam.) has equally long awns, but the spikelets and lemmas are shorter and the plant lacks rhizomes. F. rubra subsp. multiflora Piper (F. diffusa Dumort., F. heteromalla Pourret) has equally long spikelets and lemmas but the panicle is very diffuse, the culms usually much taller (often 1 m tall), and the leaf-blades are flat. F. juncifolia St Amans also has equally long spikelets and lemmas but the culms are not tufted, the rhizomes are much longer, and the innovation leaf-blades have continuous or nearly continuous abaxial sclerenchyma. F. rubra subsp. scotica is a boreal taxon; outside Scotland we have seen material from Scandinavia (including Iceland), and it is likely to occur more widely. In Scotland it occurs in a range of grassy, usually wet, habitats, from near sea-level to at least 825 m in the mountains. It is found from the type locality in Argyllshire northwards to North Mainland, Shetland Isles, and Lewis, Outer Hebrides. The only other taxon within the F. rubra group that has a similar distribution is F. rubra subsp. arctica (Hackel) Govoruchin (F. richardsonii Hook.), but the latter has shorter spikelets and lemmas which are usually glaucous and densely covered with white hairs and have usually very short (or no) awns, and is hexaploid (2n=42). F. rubra subsp. scotica is evidently either octoploid (2n=56) (type specimen; plant from Iceland) or decaploid (2n=70) (plant from Torridon, W. Ross). This taxon was first recognised by C. E. Hubbard as characteristic of high altitudes in the Cairngorms (see Webster (1978, p. 489) and Hubbard (1984, p. 374), where it is called “‘subsp. (ex Cairngorms)’’), but he did not name it. It was also recognised as a distinct taxon by S. M. Cunningham, working in Leicester in 1979-82, and he provisionally named it subsp. scotica and determined specimens from outside the Cairngorms (see Scott & Palmer (1987, p. 352)); he did not formally describe it. We have now grown it at Leicester for many years and confirmed its distinctness. Further details of morphology, anatomy, cytology and distribution will follow in a later paper, as will the results of hybridisation experiments. ACKNOWLEDGMENTS We are grateful to Hugh McAllister for providing us with living material, and to Stewart Cunningham for making his unpublished notes available to us. 316 SHORT NOTES REFERENCES HusBArD, C. E. (1984). Grasses, 3rd ed. revised by J. C. E. Hubbard. Harmondsworth. Scott, W. & PALMER, R. (1987). The flowering plants and ferns of the Shetland Islands. Lerwick. WEBSTER, M.McC. (1978). Flora of Moray, Nairn and East Inverness. Aberdeen. A.-K. K. A. AL-BERMANI & C. A. STACE Department of Botany, University of Leicester, Leicester, LE1 7RH ABNORMAL FLOWERS IN PLATANTHERA CHLORANTHA (CUSTER) REICHENB. AND GYMNADENIA CONOPSEA (L.) R.BR. SUBSP. CONOPSEA On 7 July 1989, a plant of Platanthera chlorantha (Custer) Reichenb. was found in flower near Dent in N.W. Yorks., v.c. 65. It had 19 flowers but only the uppermost one was abnormal. The height of the plant was 30 cm. The column, median outer perianth segment and lateral inner perianth segments of this abnormal flower were normal, but the lateral outer perianth segments were absent. The labellum was sepaloid, and without a spur. The dimensions of the labellum were 4-6 < 9-3 mm; it was shallowly deflexed, curled up at the distal end and trilobed. It was white with some light green at the tip and very light green veins. The sepaloid nature of the labellum of this abnormal flower is of particular interest because it bears a striking resemblance to those featured in Plate 2 and Figs. 1A & B of McKean’s (1982) description of the supposed intergeneric hybrid, x Pseudanthera breadalbanensis McKean (Platan- thera chlorantha X Pseudorchis albida (L.) A. & D. Léve). In X P. breadalbanensis all the flowers are similar (McKean 1982), each having a labellum similar to that of the abnormal P. chlorantha flower found at Dent. In the author’s opinion, x P. breadalbanensis is not a hybrid but an abnormal variant of P. chlorantha, for the following reasons. Firstly, the labella are similar to the labellum in the abnormal Dent flower. Secondly, the only morphological features of Pseudorchis albida evident in X P. breadalbanensis are the trilobed labellum and the flower length; the latter, however, merely reflects the former, and thus there are no intermediate features requiring a hybrid explanation. Thirdly, there is abundant evidence of the frequency of floral abnormalities in P. chlorantha. Hemsley (1907) wrote that in “‘P. bifolia in the broad sense, that is including P. chlorantha, many deviations from the typical floral structure are on record, . . .’’. This is borne out by Camus & Camus (1929). Only a few of the abnormalities observed in P. chlorantha need be mentioned. Flowers with no spur, two spurs and three spurs were found to be sufficiently frequent to be given names: ecalcarata (Druce 1921; Godfrey 1933), bicalcarata (Camus & Camus 1929) and tricalcarata (Hemsley 1907) respectively. These abnormalities were variously accorded the rank of variety, form and lusus. Hemsley (1907) reported that “True peloria of two kinds has been observed in Platanthera, namely lip-peloria and petal-peloria, in which the transformed organs, being of the petal series, are either like the lip or the lip takes the shape and colouring of the other two petals’. Camus & Camus (1929) specifically referred to the former kind of peloria in P. chlorantha. Thus, types (a) and (b) peloria sensu Bateman (1985) have been observed in Platanthera. Interestingly, Henslow (1858) reported a plant of P. chlorantha in which only the lowest flower was abnormal, although only approximately half the inflorescence was expanded. The flower was abnormal in that the median outer perianth segment was united with one of the lateral outer perianth segments, and instead of the usual two anther-lobes, there were four. The fourth reason for preferring a peloric interpretation of x P. breadalbanensis derives from the five observations made by Bateman (1985). Fifthly, of the habitat of the alleged hybrid McKean (1982) remarked “‘A site with so many plants of Platanthera chlorantha as well as several specimens of Pseudorchis albida must be very rare in Britain.” On 7 July 1985 at the Dent site over 200 flowering spikes of P. chlorantha were counted in the corner of the hilly pasture where the abnormal flower was found. On 28 June 1986, at the same site, six flowering spikes of P. albida were found just above this concentration. This is the only such site known to the author in Watsonian Yorkshire (v.cc. 61-65 inclusive). That similar labella should appear at similar sites, one labellum undoubtedly not the product of hybridisation, would seem to be significant. The mounted labellum of the abnormal flower of P. chlorantha found at Dent has been retained by the author. SHORT NOTES 317 The next example of an abnormal orchid flower illustrates the more common situation where all the flowers in an inflorescence are similarly and clearly abnormal. A plant of Gymnadenia conopsea (L.) R.Br. subsp. conopsea with abnormal flowers was found on 14 July 1989, near Rievaulx in v.c. 62, N.E. Yorks. All the flowers in the inflorescence were abnormal in exactly the same way. The labellum, the inner lateral perianth segments and the outer median perianth segments were normal. The outer lateral perianth segments were abnormal in that they were folded forward, not spread horizontally or downwardly curved. Further, their margins were not revolute. Clapham (1987) said of the genus Pseudorchis “‘Close to Gymnadenia but with the outer lateral perianth segments connivent with the outer median and the inner perianth segments into a hood so that the flower becomes almost campanulate’’. The plant found near Rievaulx thus had flowers with features common to both genera. To my knowledge, no other members of sub-tribe, Gymnadenii- nae (which includes Platanthera, Pseudorchis and Gymnadenia) have been recorded from this site. However G. conopsea subsp. densiflora (Wahlenb.) G. Camus, Bergon & A. Camus is not uncommon here. Camus & Camus (1929) commented that abnormalities in G. conopsea are frequent. REFERENCES BATEMAN, R. M. (1985). Peloria and pseudopeloria in British orchids. Watsonia 15: 357-359. Camus, E. G. & Camus, A. (1929). Iconographie des Orchidées d’ Europe et du Bassin Méditerraneen 2: 382-3, 405-6. Paris. CiapuHaM, A. R. (1987). Pseudorchis albida (L.) A. & D. Léve, in CLapHaM, A. R., TuTin, T. G. & Moore, D. M. Flora of the British Isles, 3rd ed., p. 570. Cambridge. Druce, G. C. (1921). New county and other records. Rep. botl Soc. Exch. Club Br. Isl. 6: 399. Goprery, M. J. (1933). Monograph and iconograph of native British Orchidaceae. Cambridge. HEMSLEY, W. B. (1907). Platanthera chlorantha Custer var. tricalcarata Hemsl. J. Linn. Soc. Lond. (Bot.) 38: 3— Spe Eig Os HENSLow, J. S. (1858). On a monstrous development in Habenaria chlorantha. Proc. Linn. Soc. Lond. (Bot.) 2: 104-105, t. IB. McKean, D. R. (1982). X Pseudanthera breadalbanensis McKean: a new intergeneric hybrid from Scotland. Watsonia 14: 129-131. F. HoRSMAN 7 Fox Wood Walk, Leeds, LS8 3BP A NONRESUPINATE ABNORMALITY IN ORCHIS PURPUREA HUDSON Rose (1948) described peloria in Orchis purpurea Hudson in Kent and various floral abnormalities have since been described. Ettlinger (1987) has drawn attention to a group of O. purpurea on a nature reserve in E. Kent (v.c. 15) displaying various degrees of peloria and nonresupination. We report a different nonresupinate abnormality without peloria, from the same nature reserve, of which we have been unable to find a previous description. A single plant was observed in flower in 1986, 1987 and 1988 and was consistently abnormal in each year. It failed to appear in 1989 and 1990. Each flower was completely inverted; the peduncular ridges were not spiralled, showing that resupination had not occurred. The usual floral parts could be identified, including pollinia, but the flowers, the inflorescence and the rosette leaves were all narrower than in associated normal plants (Fig. 1 and Table 1). Seed capsules did not mature in any year. However, seed-set was patchy among normal plants on the reserve. The complete failure to set seed might be due to the difficulty facing pollinating insects when attempting to settle on the inverted flowers and, in any event, they would probably fail to place the pollina from normal flowers on the stigmatic surfaces of the nonresupinate forms. The abnormality might be explained by the loss of a fragment of DNA carrying the gene for resupination and some accessory gene(s) controlling robust growth. 318 SHORT NOTES Ficure 1. a) The flowering spike of the nonresupinate plant (traced from a photograph). Note the rectilinear peduncular ridges. b) Individual fiorets from a normal (above) and the abnormal plant. The abnormal flower was drawn from a specimen preserved in spirit. TABLE 1. COMPARISON OF NORMAL AND NONRESUPINATE PLANTS OF ORCHIS PURPUREA HUDSON Normal plants on the same reserve Character Nonresupinate (dimensions in cm) plant n mean range S.D. Height of flowering stem 35 8 47-3 39-56 5-87 Inflorescence height 12-5 x 14-1 10-22 3-73 diameter 2-5 8 5-3 4-57-0 0-76 no. of flowers 30 8 35-9 2849 7-94 Width of flower” across ‘arms’ 0-9 12 1-38 1-0-2-0 0-33 across “skirt” 0-5 12 1-28 0-9-1-7 0-26 Rosette leaves“ length 13, 14, 13 16 15-3 11-19 2-23 width 3-5, 3-5, 3-5 16 5-0 4-3-6-3 0-52 *Plants were measured when all but the top 2-3 flowers were fully open. ” A single flower was measured on 12 different plants; dimensions appeared relatively constant for a given plant. ~ Three leaves were measured on the nonresupinate plant, but only two on each normal plant. | | | SHORT NOTES 319 ACKNOWLEDGMENTS We are grateful to D. C. Lang for drawing the florets and for with help in presenting this note; to A. C. Jermy and J. J. Wood for bibliographic help; and to Mrs Peter Morgan for translation of a Dutch article. We thank the Kent Trust for Nature Conservation for permission to take material for study and to publish. REFERENCES ErrinGeR, D. M. T. (1987). Peloric and duplex examples of Orchis purpurea Hudson in Kent. Watsonia 16: 432. Rose, F. (1948). Orchis purpurea Hudson. J. Ecol. 36: 366-377. G. I. C. INGRAM* & C. W. DUNSTER *Staple Lees, Hastingleigh, Ashford, Kent, TN25 5HG HERBARIUM SHEETS OF THE RARE FERN x ASPLENOPHYLLITIS MICRODON (T. MOORE) ALSTON IN EDINBURGH Two previously unknown Victorian herbarium sheets of the rare Guernsey Fern (x Asplenophyllitis microdon (T. Moore) Alston) have recently come to light at the herbarium of the Royal Botanic Garden, Edinburgh. In excellent condition, the two sheets (both originally ex herb. Neill Fraser, from whom they were purchased in 1905) include at least five separate gatherings made between the years 1860 & 1865, containing a total of ten beautifully preserved fronds, varying from 15 to 26cm in length. This unusual intergeneric hybrid fern between Lanceolate Spleenwort (Asplenium billotii F. W. Schultz) and Hart’s Tongue Fern (Phyllitis scolopendrium (L.) Newm.) has always been of limited wild occurrence. First found in Guernsey in 1855, it was subsequently known to occur in the mid- 19th century sporadically in Cornwall, Devon and possibly Wales (Alston 1940), while more positive records of it came from Guernsey through the latter half of the 19th century, where it still survives (McClintock 1968, 1975). It is illustrated as silhouettes in Page (1982: 129), and as field photographs in Page (1988: 131 & 132). The two newly-found Edinburgh sheets appear to include specimens from both Guernsey and Cornwall. As might be expected of a hybrid between two such morphologically differing parents, the frond form of the hybrid is variable between specimens. Previously-known original Victorian herbarium material (at BM and K) of this fern is sparse, and amounts to less in total than these new finds. The discovery of these well-preserved Edinburgh sheets thus not only doubles the amount of known herbarium material, but also helps add substantially to our knowledge of the variation pattern. REFERENCES AtstTon, A. H. G. (1940). Notes on the supposed hybridisation in the genus Asplenium found in Britain. Proc. Linn. Soc. Lond. 152: 132-144. McCuintock, D. (1968). Plant notes: Asplenium billotii x Phyllitis scolopendrium = X Asplenophyllitis microdon (T. Moore) Alston. Proc. Bot. Soc. Br. Isl. 7: 387-389. McCuintock, D. (1975). The wild flowers of Guernsey. London. Pace, C. N. (1982). The ferns of Britain and Ireland. Cambridge. PacE, C. N. (1988). Ferns. Their habitats in the landscape of Britain and Ireland. London. C. N. PAGE Royal Botanic Garden, Edinburgh, EH3 5LR A NEW SUBSPECIES OF GYMNADENIA CONOPSEA (L.) R.BR. Gymnadenia conopsea (L.) R. Br. subsp. borealis (Druce) F. Rose, comb. et stat. nov. Habenaria gymadenia Druce var. borealis Druce in Rep. botl Soc. Exch. Club Br. Isl. 5: 172 (1918). 320 SHORT NOTES Gymnadenia conopsea var. borealis (Druce) Godfery, Mon. Icon. Br. Orchidaceae 143 (1933). G. conopsea vat. insulicola Heslop-Harrison in Proc. Univ. Durham Phil. Soc. 10: 260 (1941). G. odoratissima auct. angl., non (L.) Rich. (1817). A compact plant with small, dark-pink to purple flowers; labellum about twice as long as broad, much less divided than in subsp. conopsea, though middle lobe broader than the lateral; sepals oval, acute. Flower scent strong, reminiscent of cloves. Flowering July and August. Hill pastures up to 700 m, which may be acid, and apparently the predominant subspecies from Scotland southwards to Derbyshire, but also locally in Devon and Cornwall, and in the New Forest, Hampshire, in base-rich flushes, and one site in Ashdown Forest, Sussex. A full account of the taxonomy of G. conopsea is in preparation. ~ F. ROSE Rotherhurst, 36 St Mary’s Road, Liss, Hants., GU33 7AH NEW NAMES AND COMBINATION FOR THREE HYBRIDS Opinions differ over the use of names as opposed to formulae for interspecific hybrids, but where the hybrid has a distribution pattern independent of that of its parents, due to natural or man- assisted spread, it surely demands a name in its own right. The following fall into this category. Atriplex < taschereaui Stace, hybrida nova Ho .otypus: Beckfoot, Cumberland, v.c. 70, England. Occasional on exposed coastal beach, 23 September 1978, P. M. Taschereau s.n. (LTR). Hybrida fertilis inter A. glabriuscula Edmondston et A. longipes Drejer. Ab A. glabriuscula bracteolis nonnullis ad 20 mm longis, pedicellatis ad 10 mm, herbaceis apicibus differt. Ab A. longipes bracteolis connatis in dimidio inferiore, pedicellatis ad 10 mm differt. Ab A. x gustafssoniana Taschereau (A. longipes X A. prostrata) bracteolis connatis in dimidio inferiore differt. Most plants are similar to A. glabriuscula in vegetative characters, but differ in at least some bracteoles being up to 20 mm long, herbaceous at the apex and with pedicels up to 10 mm long. The hybrid differs from A. longipes and other hybrids involving that species in having its bracteoles united at the base for up to half their length. A. longipes itself has even larger bracteoles (up to 25 mm) with longer pedicels (up to 25 (30) mm). A. X taschereaui is named in honour of Dr Pierre Taschereau, who not only collected the holotype but has contributed much to our knowledge of this genus in Britain. Taschereau (1989) has provided a more detailed description and a drawing of this hybrid, which occurs on exposed beaches in Scotland, northern England and the Isle of Man, usually (but not always) with A. glabriuscula but usually without (and commoner than) A. longipes. Mahonia < decumbens Stace, hybrida nova Ho.otypus: South of Newmarket to Cambridge road, West Suffolk, v.c. 26, England. In shady beech plantation, 2 July 1973, D. M. McClintock s.n. (BM). Hybrida (? fertilis) inter M. aquifolium (Pursh) Nutt. et M. repens (Lindley) Don. A M. aquifolium foliolis (3-)7, hebetatis, minus quam duplo longioribus quam latis differt. A M. repens foliolis 4-8 cm longis, plus quam 1-5-plo longioribus quam latis differt. This hybrid is closer to M. repens in habit, having procumbent, stoloniferous stems rarely more than 30 cm high and leaflets with a matt upper surface, but the leaflets are 4-8 x 2-4—4-5 cm (1-5 to 2 times as long as wide) with 8-11 teeth on each side. This hybrid is a garden plant (often mis-named M. repens) to which several cultivar names refer. SHORT NOTES 6 Ft | Some untypified binomials might be applicable to this hybrid, but N. P. Taylor (Kew) has kindly advised me (in litt. 1989) “*. . . if you really need a binomial I recommend you to publish a new name. The typification of the earlier names is too problematic and uncertain’. It is planted as ground cover, which is the origin of the Newmarket population, where it has been known for many years. Doronicum X excelsum (N.E.Br.) Stace, comb. et stat. nov. D. plantagineum var. excelsum N.E.Br. in Gard. Chron., ser. 2, 20: 230 (1883). This taxon is of garden origin, its most likely parentage being D. columnae Ten. x D. pardalianches L. x D. plantagineum L., as suggested by Leslie (1981). REFERENCES Lesuiz, A. C. (1981). A note on naturalized Doronicum in Britain. B.S.B.I. News 27: 22-23. TASCHEREAU, P. M. (1989). Taxonomy, morphology and distribution of Atriplex hybrids in the British Isles. Watsonia 17: 247-264. C. A. STACE Department of Botany, University of Leicester, Leicester, LEI 7RH i at ton ae omit : a - 1 gesrcome, bas 3 peers 2.008 i eeNEA IsiTES taleog fs nunca offs to igito od me | nasd 200 li SISW .2OE = i _— . Lork-gank 4 purple Boqwegg: Jat # ? D, (fweoprec. 4 rok lot é ¢ 7 ? 5 eee VIE i VY Pin = | ’ r * > r> a a e +2 ¢ : r - se ‘ q . * : X 7@ ty ch. - 5 43% 12 SATS AR SEO Re tx SiG, 2. 4 ‘ ,fugrIO ast a >= : 4 = rs a . & (Te22) sags oi baesygue 26 , lwarig ; ¥ x a m% . 1 - s AAMT ES ; steed : eta! o Sic r TOM .Yrmonwcs Zi et} MA 7 th S ' ag ii? ; rare “> haves tpt Watsonia, 18, 323-329 (1991) 323 Book Reviews Flora of the East Riding of Yorkshire. E. Crackles. Edited by R. Arnett. Pp. xii+271, with 5 text figures, 41 colour plates and 465 distribution maps. Hull University Press and Humberside County Council. 1990. Price £30 (ISBN 0-85958-487-9). The publication of Eva Crackles’ Flora of the East Riding is the most important event in Yorkshire botanical literature since the appearance of the last Flora of the East Riding of Yorkshire by J. F. Robinson in 1902. Since then there have been accounts of the plants of smaller parts of the county but never a full vice-county flora. The present work is the culmination of a lifetime’s interest and 40 years of systematic recording by Miss Crackles and fellow field botanists. It has been edited by Roger Arnett, a lecturer in the School of Geography and Natural Resources at Hull University. He also produced camera-ready copy for the printers. In his foreword Professor C. A. Stace states: ‘“‘Clearly the Flora will immediately become the authoritative and indispensable source of floristic data concerning the East Riding’’. Part I of the Flora comprises: 1) a definition of the area covered, which is divided into four geographical regions, namely Upper Derwentland, The Vale of York, The Wolds and Holderness; 2) a comprehensive chapter on geology and soils written by Dr Arnett; 3) a historical section on botanists and earlier publications; 4) an interesting chapter on habitats with examples of sites and the plants therein; 5) an explanation of the distribution patterns shown on the maps; 6) a short chapter emphasising the need for conservation. Part II is composed of the systematic part of the Flora and the distribution maps. Only vascular plants are dealt with. For each species we are given the date of the first record, its status, habitat and frequency in v.c. 61, and a code to indicate its European distribution where it was felt that this would be instructive. For records of uncommon species the name of a nearby village is given along with the date and the initials of the recorder. Changes in frequency are indicated. Details of critical species are given where they have been determined by specialists. Publications and herbaria cited in the flora are listed, as are recorders. The excellent photographs of habitats and plants are nearly all Miss Crackles’ own. An innovation in this Flora is that some of the maps are arranged in sets according to the habitat requirements of the plants. Maps of very common and very rare plants are omitted, but there are 200 additional maps which show interesting distribution patterns. Overlays are provided to show altitude; chalk and limestone; sand or gravel; clay; rivers, canals and lakes; springs; built-up areas; disused railways. However, their value is somewhat diminished by the fact that the dots on them obscure the dots on the maps. Criticisms are few. There is the occasional typographic error. Parapholis incurva is not at its “absolute northern limit’ at Spurn. Perhaps it is not well known that it also occurs in Northumberland. One might also quibble with the statement that Carex paniculata “requires highly alkaline conditions’. Outside the East Riding, it thrives in sites over millstone grit and, even allowing for a thin cover of clay till which may possibly include a few, small limestone erratics at one site, the conditions are by no means highly alkaline. Nevertheless, this splendidly produced, informative book will rank amongst the best of the recent, large format vice-county Floras and may act as both a spur and a daunting challenge to the other Yorkshire vice-county recorders. Miss Crackles has expressed a hope that this will be a flora that will be used. Its size will prevent its being taken into the field and some may be deterred by the price from buying it for themselves but, 324 BOOK REVIEWS as the standard work of reference, it will be removed frequently from library shelves and enjoyed by all those who have an interest in the flora of south-eastern Yorkshire. P. P. ABBOTT The herbarium handbook. Edited by L. Forman & D. Bridson. Pp. iv+214, with numerous illustrations. Royal Botanic Gardens, Kew. 1989. Price £12 (ISBN 0-947643-20-6). This new handboak certainly fills an important gap on the bookshelf. It provides information which until now was only available by asking experienced herbarium staff. All important herbarium techniques are dealt with in a clear and easy way, from the mounting and labelling of specimens to organizing and cataloguing a collection, and discussing modern techniques like the use of a computer data base. There are special chapters on the curation of groups that need special treatment like pteridophytes, bryophytes, fungi and lichens, and the management of a spirit collection has also been described. As the book is mainly based on the techniques used at Kew it does not always give the whole range of options available. For example, the possibility of freeze- drying fungi is not mentioned, although this is a fairly easy way of preservation which for some groups can give very good results. However, this small book is more than just a herbarium handbook. It also covers many aspects related to the work in a herbarium, like collecting specimens or illustrating and photographing plants. Furthermore it is a very friendly reference book, providing most useful information one ever so often tends to forget, such as the meaning of Latin abbreviations or dates written in Roman numerals, or the difference between a holotype and a syntype. To make it even more useful the index could have been a little more detailed, and maybe important page numbers printed in bold would have helped to save searching. Nevertheless it is an excellent book and these very minor points of criticism should not obscure its merits. It is undoubtedly essential for all herbarium staff but also very valuable for anybody who is interested in collecting and preserving plants. To me it is a most helpful guide which I would not like to have to do without. H. HOFMANN Spartina anglica — a research review. Edited by A. J. Gray and P. E. M. Benham. Institute of Terrestrial Ecology & H.M.S.O., London. 1990. Price £8 (ISBN 0-11-701477-X). The story of Spartina anglica is now well-known — a vigorous and fertile plant with immense abilities to invade large areas of estuarine mudflat. Since evolution from a sterile hybrid in the late 19th century it has spread to many intertidal areas in the temperate zone. This review combines some new material with the proceedings of a conference held in 1985. The story of S. anglica unfolds through a series of complementary papers dealing with its biogeography, ecology and genetics. The volume covers the sensitivity of the species to oil pollution and heavy metals and considers its productivity and possible uses as a bio-fuel. The dilemma for mankind (S. anglica — friend or foe) is covered in the final inconclusive paper dwelling on the advantages of this highly successful plant for land reclamation and prevention of erosion against the disadvantages of the swift and fundamental changes that the plant can bring to estuaries of national and international value for birds. Although tending towards the academic, the botanist will find much in this volume of interest, but a more detailed examination of the impact of Spartina on substrate, and on mud-dwelling invertebrates, would have been a welcome addition. A more holistic view of the overall impact of S. BOOK REVIEWS 325 anglica and a glimpse of the likely future spread both of the species and its associated ‘die back’ would have been informative. P. ROTHWELL Plant names of medieval England. A. Hunt. Pp. 400. Boydell & Brewer Ltd., Woodbridge. 1989. Price £35 (ISBN 085991-—266-3). Popular medicine in thirteenth century England. A. Hunt. Pp. xiit+476. Boydell & Brewer Ltd., Woodbridge. 1990. Price £39.50 (ISBN 085991-290-4). With these two books Tony Hunt establishes a new reference point for vernacular plant taxonomy in 13th—15th Century England. Those interested in medieval herbals or vernacular names will want to have Plant names of medieval England on their shelves. Making progress in this field is like grasping smoke, and this book should be used with caution. Hunt comes to this field as a scholar of Anglo- Norman French and botanists will question many of his identifications, even though the standard is high. Hunt’s achievement is to distil the Latin nomenclature of 64 texts dated between 1280 and 1500, with their associated vernacular glosses, into the easily digestible form of Plant names. He has started more lexical hares than he has caught, but for that we should be grateful. The introduction to Plant names sketches the problems of such a study and lists the sources, witha thorough bibliography. The main body of text is a dictionary of the Latin or Greek names in alphabetical order: Abaligustra, Abies, Abitumon, etc., as they appear in medieval herbals. Under this is the botanical identification, and the English names that gloss the Latin: Abies = fir-tree, etc. Herein lies the main problem of the work, an over-reliance on medieval Latin names over English vernacular. Some English names are found as glosses for many medieval Latin names, for instance ‘mugwort’ (for 22). Hunt concludes that they are polysemous, which they are not. Mugwort has, since Anglo-Saxon times, meant only one thing to an Englishman, Artemisia vulgaris. It is the Latin names which are polysemous. It seems to me that this polysemy has a consequence of tremendous importance: out of it comes the concept of the genus, alien to the almost totally uninomial folk taxonomies of Europe. For instance, in trying to interpret the Latin uninomial ‘Consolida’, confused herbalists applied it to several different plants, and to resolve this confusion, these were later distinguished as Consolida major (Symphytum officinale), Consolida media (Chrysanthemum leucanthemum) and Consolida minor (Bellis perennis). The idea of genus and epithet so established led directly to the Linnaean binomial system. Hunt repeats the blunder (pace Grattan & Singer, Anglo-Saxon magic & medicine (1952, p. 87) and Earl, English plant names from the tenth to the fifteenth century (1880, p. 5)) of associating ‘bothen’ with Rosmarinus officinalis. The Anglo-Saxon ‘bothen’ (later corrupted to bothel) means only one plant, Chrysanthemum segetum, although it has been applied to other weedy Compositae, as in white bothen for Chrysanthemum leucanthemum (in the appendix of Gerard’s Herbal) and blue bothen for Centaurea cyanus (hence bluebottle). There are some details that might make a botanist blanch, such as Pulegium montanum identified as Thymus serpyllum, not Thymus pulegium or Thymus spp., and for that matter the persistent use of ‘ssp.’ when ‘spp.’ is intended. Likewise the Index of Botanical Names is irritating with the initial of every epithet capitalised. The author touches upon, but does not solve, the problem of polarity in translation names. For instance, is Oculus Bovis just a translation of ox eye (Chrysanthemum leucanthemum), or is it the other way round? The existence of a good Anglo-Saxon name, meadwort, for this plant leads me to suspect that ox eye has never been more than a translated book-name. The 19th century flora writers who were so indefatigable in coining pseudo-vernacular book-names, may just have been at the head of a long line. In Popular medicine Hunt gives a picture of 13th century botany (indistinguishable from medicine at this date). In the preface he draws attention to the fact that 80 per cent of the world’s population relies still on popular medicine, and thus the study of vernacular herbalism is far from being an irrelevance. The introduction is the most interesting part of this book for the general reader, covering the history, form and content of the medical receipt from Babylonian times to the 13th 326 BOOK REVIEWS century, the preparation of compound medicines, collection of herbs, weights and measures and the archaeological evidence. After the introduction there follow 272 pages of well edited medieval texts. For the first time these works can be studied with the problems of palaeography and lexis largely removed. The reader is saved from laziness however, as no translations from the Latin, Anglo-Norman French or Middle English are provided — to do so would have unreasonably increased the bulk and price of the book. There follow 127 pages of notes and glossaries. Not all the botanical information is correct. Bresil/ Brasile (pp. 202 & 309) is absurdly glossed as Sequoia sempervirens, when it is actually Caesalpinia sappan, imported from the East Indies. For a big book the index is slight. Q. C. B. CRONK & ~ Atlas Florae Europaeae, Volume VIII: Nymphaeaceae to Ranunculaceae. Edited by J. Jalas & J. Suominen. Pp. 261. Committee for Mapping the Flora of Europe and Societas Biologica Fennica Vanamo, Helsinki, Finland. 1989. Price 460 Finnish Marks (ISBN 951-9108—07-6). Chorology, the study of the geographical distribution of species on the earth’s surface, is nowadays a more exact science than our Victorian counterparts knew. In particular, the spread of dot grid mapping continues to provide objectively justifiable maps for botanists and zoologists in many fields. None of these mapping projects is more impressive than the Atlas Florae Europaeae, begun as an international cooperative offshoot of the Flora Europaea project in 1965, and administered from Helsinki. The previous seven volumes, covering Pteridophyta, Gymnospermae and the beginning of the Angiospermae, appeared between 1972 and 1986, and were reprinted as a compendium in three volumes by Cambridge University Press in 1988. The arrangement of the work follows that of Flora Europaea itself. It is now 26 years since the first volume of Flora Europaea was published, and in many respects the text of this first volume is known to be out of date and needing modification in the light of newly published work. One of the strengths of the Aflas project is that the editors have consistently not only indicated where their treatment differs from that of Flora Europaea but have also given chapter and verse for that difference. Since the Adlas is still, in this eighth volume, only dealing with families in the first volume of Flora Europaea, it is not surprising to find quite large changes, both taxonomic and nomenclatural, between the two works. The volume is very largely devoted to the family Ranunculaceae, and many familiar British plants are naturally featured. As an example of change, the Atlas gives ten subspecies of Aconitum napellus: these correspond roughly with the ‘A. napellus group’ of five species in Flora Europaea, and the British plant, mapped in the Af/as under the name of A. napellus subsp. napellus, is indicated as almost, but not quite, endemic to Britain, with a single dot in south-western France. The widespread plant of northern France is separately mapped as subsp. lusitanicum Rouy. In avowedly critical groups, the Atlas is often commendably frank. Thus, the maps of Batrachian Ranunculus taxa are prefaced by a carefully-phrased warning that concludes as follows: ‘“Neverthe- less, we consider that the ... maps below, with all their faults and unevenness, are more informative than collective maps and more useful for future research. . .”., asentiment with which I feel sure most users will agree. The remarkably high standard of production achieved so far in the Atlas has been fully maintained in this volume. In particular, the excellent paper means that the maps are reproduced with perfect clarity. As for minor proof-reading errors, I can only report that I have yet to find a single one, though I have referred to many pages. It is perhaps a pity that no information accompanies this volume as to the future intentions, in collaboration with Cambridge University Press, of continuing the publication in compendium form. A quick calculation based on Flora Europaea pagination suggests that the now published volume 8 of the Atlas, together with the four projected volumes, would conveniently fit into another three volumes of the compendium. Before the end of the century, European botanists can confidently expect a completed companion Atlas to Flora Europaea volume 1. But by then, it is rumoured, we shall have a brand new second edition of that F. E. volume, so presumably the process can start again! BOOK REVIEWS 327 In the Preface, the editors express confidence that the project will keep to its schedule to finish mapping all the taxa in Volume 1 of Flora Europaea by 1995, and indicate that this will require four more volumes. All European botanists will wish them success! S. M. WALTERS Plants of Upper Teesdale. C. Lowe, with contributions from M. Lowes & G. Ward. Illustrations by J. Fielding. Pp. ix+75, with 50 line drawings and 1 map. Privately published. 1990. Price £4.75 by post from Chris Lowe, 4 Brick Kiln Cottages, Boxford, Colchester, Essex, CO6 SNT. Responsibly excluding the National Nature Reserve (apart from the Cow Green Nature Trail) and all plants not accessible by path, this book understandably had difficulty in meeting an apparent target of 50 ‘Teesdale specialities’. Indeed, only 35 such species are included. The objective, in a book aimed at the intermediate level botanist, is to give a flavour of the flora of Upper Teesdale. Myrrhis odorata (L.) Scop., Trifolium medium L., Peucedanum ostruthium (L.) Koch, and Mimulus spp. are tenuously included in the 35. By popular visitor demand, eight species of orchids are described, with an informed reference to conservation, and the 50 are made up with seven late- flowering meadow species “‘not originally destined for inclusion’’. Eight routes or sites, chosen to see these plants, are carefully explained and clearly benefit from local knowledge. These are accompanied by a map and a chart. The arrangement is phenological, apart from the orchids. The illustrations with each plant vary in quality: Gentiana verna L. and Minuartia verna (L.) Hiern are well caught, but that of Dactylorhiza purpurella (T. & T. A. Stephenson) So6 is unhelpful. Whether the stem is solid or hollow (with two safe tests given) is usefully emphasised in a key to the orchids. In many of the 50 plants described the differences from similar species are helpfully explained. The form of the footnotes rather creates the impression of afterthoughts, and the statement that “Several other “Teesdale specialities’ are not in fact dealt with in this book”’ is patently incorrect. Some local names are given; I particularly like ‘Prying Annie’ for Succisa pratensis Moench. Given that several references are made to the various geographical elements represented in the flora of Upper Teesdale, and to “. . . the relict late-Glacial arctic/alpine flora’, a short section explaining why the flora is thus unique would have been helpful. One reference is made to a discovery, namely: “John Binks discovered [Equisetum pratense Ehrh.]| in early Victorian times’! Binks died in 1817 (The botanical exploration of Upper Teesdale, F. Horsman in prep.). A party including James Backhouse Snr. and Jnr. discovered it in Upper Teesdale in 1844 (Phytol. 1: 1065-69, 1844). Given the constraints placed upon this book, dealing with such a sensitive area, it is a good compromise. However, for those few members who have not yet made their pilgrimage to this mecca, I suspect they will find the book rather disappointing. F. HORSMAN Plants of Dhofar. A. G. Miller & M. Morris; illustrated by S. Stuart-Smith. Pp. xxvii+361, with numerous colour illustrations. Published by the Government of Oman, Muscat. 1988. Price £30 (ISBN 0—7157-0808-2). This beautiful and scholarly book highlights the remarkable flora of Dhofar, a little known botanical and cultural enclave on the southern coast of the Arabian Peninsula. I accompanied Tony Miller (of the Royal Botanic Garden at Edinburgh) on his first plant collecting trip to this Southern Region of Oman in September 1979. Despite being familiar with the plants of northern Oman, most of the species and many of the genera here were unknown to me and this book will become a valued and interesting reference work for other ecologists, botanists and travellers who are similarly greeted by the grand vistas of the moist jebels of Dhofar. About 190 of the approximately 750 species of higher plant found in Dhofar are illustrated, all in 328 BOOK REVIEWS colour. The illustrations are excellent in terms of their lay-out, artistry and their attention to botanical detail; dissections of flowers and fruit and details of the bark and thorns and succulent stems are included where appropriate. Most of the plants were painted from living specimens and many have never been illustrated before; Susanna Stuart-Smith deservedly won a gold medal when some of these paintings were exhibited at the Royal Horticultural Society. Facing each plate a single paragraph of 7-10 lines gives a clear and detailed botanical description (in English) of the species concerned. This is followed by text describing the distribution and habitat of the species (and genus) in Dhofar and elsewhere, along with other information on the plant’s biology, the origins of the Latin name, etc. This makes interesting reading and presents a challenge to the plant geographer as there are two genera and 50 species endemic to Dhofar and a number of taxa with markedly discontinuous distributions; the most remarkable of these is Thamnosma (Rutaceae) which.has one species in Somalia and South Arabia, another species on Socotra (with which the flora of Dhofar has close affinities), two in the deserts of S. W. Africa and a further two in California and Mexico. This botanical section leads into the description and discussion of the ethnobotany of the plants. Just as the botanical and artistic parts of this book are a tribute to the efforts of their creators, the clear way in which the wealth of information on the varied uses of these plants has been presented is a credit to Miranda Morris, who has conducted painstaking research into the non-Arabic languages and ethnography of the people of Dhofar for more than 10 years. To a botanist, and I hope a wider audience, these sections are eminently readable and not just a list, thanks to Dr. Morris’s style and intimate knowledge of the area and its people. One should not be deterred by the use of specialist terms as there is an informative glossary which will add some words to your vocabulary — did you know that ‘‘sternutatory” refers to an agent that causes sneezing? The Jibbalis (the hill people of Dhofar) evidently know their plants and how to benefit from them in diverse ways, and to have a botanist and linguist (plus artist) working together ensures that the information on medicinal use is accurate and relates to a particular, named species. In years to come, the information may not be available from its original source as the old lifestyle of the Jibbalis and their traditional dependence on plants disappears with the rapid development that has taken place in Oman since 1970. I have noticed a few very minor discrepancies in the book, e.g. on p. 6 Blepharis linariaefolia is said to be the only species of Blepharis in Oman, other than B. dhofarensis, yet on p. 8 a third species B. ciliaris is said to occur in northern Oman; there is no reference in the text to the lovely illustration of Eulophia guineensis as the frontispiece; the introductory paragraph on the genus Acacia is presented on p. 180 following descriptions of two other Acacia species on the previous pages. Information on the local distribution of a particular plant in the botanical part of the text is occasionally repeated in the ethnobotanical part (for instance Ipomoea pes-caprae) — and it is the only book I can recall that has two separate introductions! But these quirks do not detract from the positive aspects of the book. It is not a field guide (it weighs 2 kg!) and does not pretend to be one, but I think a good selection of plants has been chosen including nearly all of the dominant woody species, most of the endemics and those of especial scientific interest — thus species such as the frankincense (Boswellia sacra), the wild myrrhs (Commiphora spp.), the aloes (Aloe spp.) and other succulents and tubers (particularly in the Asclepiadaceae) are covered. The families are presented in alphabetical order which I find satisfactory, but I would have preferred the Dicotyledones and Monocotyledones to have been treated separately. Grasses are not included. Brief sections discuss the climate (the monsoon rain and the dense mist and cloud that make Dhofar so unusual), the vegetation zones, phytogeography, local plant names, traditional medicine, etc., and there are checklists of Latin and local names. Now that Oman is cautiously opening its doors to visitors, I can recommend Dhofar to those considering an adventurous holiday in an unusual and beautiful part of the World, and I certainly recommend this book as one of their companions and to take home and treasure. R. WHITCOMBE Flora of New Zealand, Volume IV. Naturalised Pteridophytes, Gymnosperms, Dicotyledons. C. J. Webb, W. R. Sykes & P. J. Garnock-Jones. Pp. Ixviii+ 1365, with 16 colour plates and 123 figures in the text. D.S.I.R., Christchurch, New Zealand. 1988. Price N.Z.$88 (ISBN 0-477-02529-3). BOOK REVIEWS 329 The study of the arrival, establishment and subsequent spread of introduced plant species has long been a subject of some interest to botanists throughout the world. Many papers and a number of books have been published on the topic, but apart from S. T. Dunn’s fragmentary Alien Flora of Britain (1905) a regional flora largely devoted to naturalized adventives is virtually a novelty. The New Zealand flora comprises over 2900 taxa, of which approximately half are established introductions originating from other parts of the globe. The latest volume of the Flora treats these except for the monocotyledons which will be dealt with in a later part. Keys and detailed descriptions of families, genera and species are provided and data are given on their introduction, dispersal mechanisms, habitat preferences and ecology. Keys are also provided to families containing aquatic and parasitic species. For genera possessing both indigenous and introduced species the former are also keyed out and described. Among the genera with the largest number of naturalized species are Trifolium, Rubus, Eucalyptus, Solanum, Salix, Senecio and Veronica. Hundreds of taxa presumed native in the British Isles are found as adventives in New Zealand, including such rarities as Crepis foetida, Erica vagans, Galium debile and Veronica verna. Many other introductions are also adventives familiar to British botanists, e.g. Galinsoga parviflora, Calystegia silvatica, Epilobium ciliatum, Rubus procerus, Heracleum mantegazzianum, Buddleja davidii and Veronica filiformis, the last mentioned a comparatively recent invader of lawns and grass verges. Notable absentees in the islands are Aethusa cynapium, Chenopodium rubrum and Lepidium ruderale. Betula pendula is naturalized but B. pubescens, though in cultivation, does not appear to have escaped to the wild. Conyza canadensis is part of a complex, the true species being rare, and the most widespread being the S. American C. bilbaoana E. J. Remy, which differs in its inconspicuous ligules, and narrow-triangular, inner involucral bracts; C. parva Cronq. from N. America, generally a smaller, more slender plant than C. canadensis with glabrous, linear bracts is also present. Do we perhaps need to reexamine our British populations of C. canadensis? An interesting factor is that, as in Europe, there has been a great decline in cornfield weeds. This began earlier for Agrostemma githago and Scandix pecten-veneris; both were widespread at the turn of the century but have not been reported from New Zealand since 1930. Agricultural pests are all too frequent and include an unlikely trio in Ulex europaeus, Berberis darwinii and Erica lusitanica. The taxonomy and nomenclature used are up-to-date and of a very high standard, but Amsinckia calycina (Moris) Chater is probably referable to A. micrantha Suksd., Anchusa barrelieri (All.) Vitman is now usually assigned to Cynoglottis barrelieri (All.) Vural & Kit Tan, while Galeobdolon luteum cv. Variegatum is probably synonymous with G. argentatum Smejkal, recently reduced to Lamiastrum galeobdolon subsp. argentatum (Smejkal) Stace in Britain. Conyza albida Willd. ex Sprengel is a synonym of C. sumatrensis (Retz.) E. Walker, naturalized for some years in the Channel Islands, and now spreading in S.E. England, while the earliest name for Silene maritima as a subspecies of S. vulgaris appears to be S. vulgaris subsp. alpina (Lam.) Nyman and not S. vulgaris subsp. maritima (With.) A. & D. Léve. These, however, are minor points in an identification manual of great excellence which should prove an invaluable addition to the bookshelves of all plant taxonomists. D. H. KENT so ORE REPLI RU! - 6 deaeth ce ne rFeh: fact en Smartt esac sab Vee combed vorielage gat “AGliey a2 ti Me si hedgaeinees oes x | m4 ; Ai ge Drill WEES? SRLS AR oy ent bee preine arene Cotas elites hw ayserlia sare ‘Sash gidecoy (sndpeettineiniconec UL Sy, 1H me Lee, Sree a sitepiteg gery Seaion aya Cane uate fhm, t a err oan tralia tosis Aux: epireul: trigual Soda ret “a eee wien be cmsarey aay UO 2 hetiawht wire bag cise, wine. a om neti 443 SAR jee eemeesit wy eee siy dbutant’, binmebanh, areiPishirentibiee ify PERG > ieee 7 tani tertks (Ot anise Shite test pray ronment RECORD Coon ea acelin, heheheh ier yan Song mecea re ae eer en snmiciy vehi boy SM vigeise abled Lote tid Seelam hime: oda pcan ale tae eal nN ts eee petertsl 7h aukneetedeieund ll Nal teatlase on idatanny watt rsh cipegayes we eles aeerediony such dase alain aaeial oe Oh Se haw ery dhewatet iene ie vanitavilers Diteoudia SSM Mate) A one aero ea ‘gah iy (otere wee toe ener ie ys Ke coletnop a Bild | eural ysl: cter Spain Wileye oth) eect siti wah Pon beb| matali pa aay aries 2a ; pls See es hieie ell eee Wate ivi ehh open ehambelbacheat ent cr Dee oak tek Heit ar ogee ever ot atin orwhpasrerrenet gia , i : a A RS i cacy ee aioe aera | fyi i : ty ot Preaphatety watbag ity rane Reap Sud ttt suede beta hee! ‘ooh eo wena eit este { , at >) 8 S y N fu i ) . gai ‘ euch wetittih Chaw.n & Dewcaauehl ante iaaetiaee t ptitev fi sive A - ES Seer) eer ietrones: peal tod iayacinnl ‘ pe i wets, WAAR weet scadayt fs Peed. Mag. tn tt ' A? Ra eA : aie dy ee Dy MR eS WR SAls Maia dei sainiehd rin AB . ua,’ aay pe cca falwiist- ie AS teh elias bdetrn bela 34 oe Folios ae. ¥ Hrtwr cia’ T7 27282, Lea Glan Hsin tits lyno sion» teomghaaang elaine weer “ Marois i i $ et 4 \ “i w ; » _ , KU ets Ee TEENA La Ree ee) OT ead Raat pe myih eb Tike meats a Fi q - : oy } , } bs haf ; ry :b ee lt ers, hy tae 9 Tet pat ashen bt i a ao a we nts Pik Siar S| ? ¥ aS th ote! hw AB LU RR, VE poy AP} en ay ¢ \ wy 16 ; ie | Oe ees b ta kh VonWemeSaeS Ee TU RINDI sai v4 | i oat ; rate % (iS) Ue eG rai oe PhS Ki Ee Chiat ir W9gi38 i 4 Me wa , ae n ria fl SU Uy Ue Cras? Gl ele tol Tis) solivelaiens nid prawn swim lihette dae aati - 1} v r | 4, ys \ be eat « \ y % § ‘{ A I it mu { - j ; " y Uy, ee nw Watsonia, 18, 331-332 (1991) 331 | Report ANNUAL GENERAL MEETING, 12 MAY 1990 The Annual General Meeting of the Society was held in the Bourne Laboratories Lecture Theatre, Royal Holloway and Bedford New College, Egham Hill, Egham, Surrey, by kind invitation of Professor W. C. Chaloner. Professor J. D. Dodge welcomed the Society to the College and 105 members were present to hear the Presidential Address, by Professor D. A. Webb, Genera, hollow curves and boojums (see page 247). This was followed by the Annual General Meeting at 11.45. Apologies for absence were read, including those of Professor Chaloner who was in France for his election as a Foreign Member of the French Academy of Sciences, on which this Society offered congratulations. Minutes of the 1989 Annual General Meeting as published in Watsonia 18: 115-117 (1990), were approved and signed by the President, Professor D. A. Webb, Sc.D., F.M.L.S., who was in the Chair. REPORT OF COUNCIL The report had been circulated to members and there were no queries from the floor. The Hon. Treasurer commented that since the Report had been written, changes in staff and work priorities in the Department of Botany at The Natural History Museum might affect the joint project for a computerised database for nomenclature and bibliography of the flora of Britain and Ireland, as approved by Council. The funding for this project from the Welch Bequest would therefore be delayed until the arrangements with the Museum had been clarified. The adoption of the Report was then proposed by Dr T. C. G. Rich, seconded by Mr. A. O. Chater, and unanimously approved. TREASURER’S REPORT AND ACCOUNTS The Hon. Treasurer, proposing the adoption of his Report and Accounts, offered to explain any points, but there were no queries. The adoption was seconded by Mr D. J. McCosh and passed unanimously. RE-ELECTION OF HON. GENERAL SECRETARY AND HON. TREASURER The re-election of these officers was proposed by Dr T. C. G. Rich, seconded by Dr F. H. Perring and unanimously approved with applause. ELECTION OF COUNCIL MEMBERS In accordance with Rule 10, nominations had been received for Miss L. Farrell, Mr D. A. Pearman and Mr C. D. Preston. Their election was proposed by Dr R. J. Pankhurst, seconded by Mr J. Bevan, and passed unanimously. RE-ELECTION OF HONORARY AUDITORS The Hon. Treasurer in proposing the re-election of Grant Thornton, West Walk, Leicester emphasised that it was an honour for the Society to present their Accounts over the name of these 332 REPORT distinguished Auditors. Their election was passed unanimously, and the President agreed to write expressing thanks from the Society. ANY OTHER BUSINESS Mr R. G. Ellis proposed with appreciation and affection a vote of thanks to the re-elected officers. This was seconded by Mrs A. Lee and carried with applause. Mr B. Gale and his assistants were thanked for their local organisation, and the meeting ended at 12,55, & ~ M. BriGGs FIELD EXCURSIONS HELD IN CONJUNCTION WITH THE A.G.M. LONDON UNIVERSITY BOTANIC GARDENS, SURREY. 12 MAY 1990 After the A.G.M., members visited the University’s Botanic Gardens — where guided tours were led by Dr B. W. Ferry, Mr B. A. Gale, Dr A. J. Hemsley and Dr M. Ingrouille. LANGHAM PONDS, SURREY. 12 MAY 1990 30 members enjoyed an evening visit to this famous S.S.S.I. — led by Mr R. Davidge and Mr J. M. Mullin. SILWOOD PARK, BERKSHIRE. 13 MAy 1990 About 40 members spent a splendid day at the country estate of the Imperial College of Science, Technology & Medicine (University of London). Dr V. K. Brown, Dr M. J. Crawley and Dr A. Gange explained the research work on plant-invertebrate interrelationships that is being underta- ken there and led tours of the extensive grounds. B. A. GALE ee ee ee ee ae Se Te See ee toe, A el & as ee at Dig eer 2 ote Eee n po ee = = re) O i W. Cornwall 1b. Scilly E. Cornwall S. Devon N. Devon S. Somerset N. Somerset N. Wilts. S. Wilts. . Dorset . Wight ‘3 . S. Hants. . N. Hants. . W. Sussex . E. Sussex . EKent . W. Kent . Surrey . S. Essex . N. Essex . Heris: . Middlesex . Berks. . Oxon . Bucks. . E. Suffolk . W. Suffolk . E. Norfolk . W. Norfolk . Cambs. . Beds. . Hunts. . Northants. . E. Gloucs. . W. Gloucs. . Mons. . Herefs. . Worcs. . Warks. S. Kerry N. Kerry W. Cork Mid Cork E. Cork Co. Waterford S. Tipperary Co. Limerick Co. Clare H10. N. Tipperary . Co. Kilkenny H12. Co. Wexford . Co. Carlow H14. Laois NAMES OF VICE-COUNTIES IN WATSONIA ENGLAND, WALES AND SCOTLAND 35 . Staffs. . Salop . Glam. . Brecs. . Rads. . Carms. . Pembs. . Cards. . Monts. . Merioneth . Caerns. . Denbs. . Flints. . Anglesey , 3S eis: . N. Lincs. Leics. 55b. Rutland 69 . Notts. . Derbys. . Cheshire iS. "Bancs: . W. Lancs. ‘S.E- Yorks. . N.E. Yorks. . S.W. Yorks. . Mid-W. Yorks. . N.W. Yorks. . Co. Durham . S. Northumb. . Cheviot Westmorland 69b. Furness . Cumberland . Man . Dumfriess. . Kirkcudbrights. . Wigtowns. . Ayrs. IRELAND . S.E. Galway H16. W. Galway H17. N.E. Galway . Offaly H19. Co. Kildare H20. Co. Wicklow . Co. Dublin . Meath . Westmeath H24. Co. Longford . Co. Roscommon H26. E. Mayo . W. Mayo . Co. Sligo . Renfrews. . Lanarks. . Peebless. . Selkirks. . Roxburghs. . Berwicks. . E. Lothian . Midlothian . W. Lothian . Fife . Stirlings. . W. Perth . Mid Perth . E. Perth . Angus . Kincardines. . S. Aberdeen . N. Aberdeen . Banffs. . Moray . Easterness 96b. Nairns. . Westerness . Main Argyil . Dunbarton . Clyde Is. . Kintyre . S. Ebudes . Mid Ebudes . N. Ebudes . W. Ross . E. Ross . E. Sutherland . W. Sutherland . Caithness . Outer Hebrides . Orkney . Shetland . Co. Leitrim . Co. Cavan . Co. Louth . Co. Monaghan . Fermanagh . E. Donegal . W. Donegal . Tyrone . Co. Armagh . Co. Down . Co. Antrim . Co. Londonderry WATSONIA Botanical Society of the British Isles Patron: Her Majesty Queen Elizabeth the Queen Mother Applications for membership should be addressed to the Hon. General Secretary, c/o Department of Botany, The Natural History Museum, Cromwell Road, London, SW7 5BD, from whom copies of the Society’s Prospectus may be obtained. Officers for 1991-92 Elected at the Annual General Meeting, 4th May 1991 President, Dr P. Macpherson Vice-Presidents, Mr J. Ounsted, Mr P. S. Green, Dr G. Halliday, Mr A. C. Jermy Honorary General Secretary, Mrs M. Briggs Honorary Treasurer, Mr M. Walpole Editors of Watsonia Papers and Short Notes, J. R. Edmondson, R. J. Gornall, E. C. Nelson, B. S. Rushton* Plant Records, C. D. Preston Book Reviews, J. R. Edmondson Obituaries, J. R. Akeroyd “Receiving editor, to whom all MSS should be sent (see inside back cover). era Orcp 2? 3 1995 GRAY HERBARIUM Watsonia, 18, 333-342 (1991) 333 The pollination of Arum maculatum L. — a historical review and new observations A. J. LACK and A. DIAZ School of Biological and Molecular Sciences, Oxford Polytechnic, Headington, Oxford, OX3 OBP ABSTRACT Previous accounts of the complex trap pollination mechanism in Arum maculatum L. (Araceae) are briefly reviewed; they conflict in some details and are incomplete. Our own study showed that the species is self- incompatible; it traps female owl-midges Psychoda phalaenoides as the main pollinator, and papillae on the spathe and column surfaces stop them escaping. A fluid, consisting of a dilute sugar solution, surrounds the stigmas in most plants. This is not taken by the flies but acts as a site for pollen deposition and germination. One pollen-bearing fly can carry over 150 pollen grains and can effect full pollination of the female flowers but, in our study sites, the numbers of flies were limited and some inflorescences failed to set fruit owing to lack of pollination. Experimental removal of parts of the inflorescence led to a reduction in the numbers of flies caught and in the number of inflorescences setting fruit, contrasting with previous work. It is concluded that all parts of the trap mechanism are essential to ensure full fruit set. In addition, the more flies that are caught, the more likely a plant is to act as a pollen donor to other plants. INTRODUCTION Lords-and-Ladies, Arum maculatum L., is one of Britain’s most distinctive native plants. It grows in woodlands and shady hedgerows where the first signs of its presence are the broad glossy emerald leaves as they emerge through the sparse February ground flora. In late April or early May a pointed shoot growing from the centre of the leaf rosette unfurls to become the bizarre inflorescence for which the plant is famed. The species belongs to the huge, mainly tropical family, Araceae and is the only common British member of the genus. The centre of radiation of the genus Arum is in the Eastern Mediterranean (Prime 1960, 1980) and Britain and Ireland form the northern and western limit of its range. Bown (1988) gives a general, and beautifully illustrated, account of the whole family. Arum maculatum spreads vegetatively to form clonal colonies, but these are restricted to a small local patch, up to about 1 m in diameter (Prime 1960). Reproduction and dispersal in the species is mainly by seed. The inflorescence of A. maculatum consists of a group of 20-40 female flowers at the base, then a whorl of hairs derived from sterile flowers, followed by 60-100 male flowers, another set of hairs and the whole is topped by a spadix (Fig. 1). A spathe surrounds this and remains fully furled around all the flowers up to the base of the spadix, acting as a trap for insects. This most unusual inflorescence, along with other peculiar features of the plant, has attracted considerable interest which culminated in the publication of the classic monograph, Lords-and-Ladies, by Prime (1960, reprinted 1981). Prime readily acknowledged that much remains to be discovered, but since 1960 very little work appears to have been done. FUNCTION OF THE INFLORESCENCE PARTS IN POLLINATION PREVIOUS ACCOUNTS In past accounts there was some confusion over the working of the trap mechanism and the importance of the various parts of the inflorescence in pollination. Prime (1960) described in detail the events surrounding pollination, his account consisting of an amalgamation of past work and 334 A. J. LACK AND A. DIAZ spadix sc ~~ Tae aes a a Taos upper whorl of sterile hairs \ x % s ll neem male flowers ; Pig tes) Tes lower whorl of sterile hairs 5 EM j Loy ee female flowers fads / ] a awe —_—_—— stigmas WA Cs ed Ficure 1. Inflorescence of Arum maculatum with spathe chamber cut open. some of his own observations. The following is a summary of his account: as the spathe unfurls the female flowers are receptive and the spadix produces a ‘“‘foul and urinous”’ smell from compounds volatilised by respiratory activity in the spadix. The smell attracts flies which breed in dung and, in particular, females of a single species of owl midge, Psychoda phalaenoides L. It is not clear where these flies alight, from Prime’s account, but, by implication, they alight at the base of the spadix. The flies crawl downwards into the spathe chamber and, once inside, the upper whorl of trap hairs makes it impossible for them to fly out (though he suggests that they could crawl out up the spathe since the hairs do not reach the chamber wall). Through walking around the chamber, they may deposit any pollen they are carrying on to the receptive stigmas. Usually by the following morning POLLINATION OF ARUM MACULATUM fe the smell of the spadix is gone, the stigmas are withered and a small drop of “‘nectar”’ (see later) is secreted by the female flowers, which the flies take. Pollen is shed by the male flowers, some of which is picked up by the flies, and the trap hairs begin to wither. Prime suggested that the flies then escape by crawling up the spathe wall. Unknown to Prime until after he had written his account, Knoll (1926) had studied Arum nigrum in detail in Dalmatia, and this was brought to the attention of British botanists by Dormer (1960). Differences from Prime’s account include the fact that the insects land on the spathe (though this is not entirely clear) and fall into the chamber (not crawl) as a result of its slippery surface. This surface has downward pointing papillae covered with oil droplets, to half way down the spathe chamber. The insects cannot climb the spathe or up the central column, owing to similar papillae on the column just above the female flowers. The upper whorl of hairs is therefore not relevant in preventing escape of the trapped flies, but stops larger insects, which may damage the inflorescence, from entering. ‘“Nectar” is secreted throughout the female phase as well as the male phase and shedding of the pollen is accompanied by a crumpling of the epidermal cells of the column, allowing the insects to crawl up the spadix, rather than the spathe, to escape. Although Knoll’s work applied specifically to Arum nigrum and not A. maculatum, Meeuse (1961), Proctor & Yeo (1973) and Faegri & van der Pijl (1979) all quoted his account with the proviso that some of the details may not apply to A. maculatum, and ignored Prime’s. Popular accounts in newspapers, newsletters and floras have variously combined these accounts and invented new variations of their own! A. nigrum is larger than A. maculatum and the lower whorl of trap hairs is similar to the upper whorl, whereas in A. maculatum the lower whorl has fewer hairs (often only two or three, Fig. 1) (Knoll 1926; Proctor & Yeo 1973 and personal observations). A. nigrum appears to attract many dung-feeding flies (all those small enough were found in the chamber (Knoll 1926)) whereas A. maculatum specifically attracts female Psychoda phalaenoides and, in places, P. grisescens Tonnoir (Proctor & Yeo 1973). OBSERVATIONS FROM THE PRESENT STUDY This investigation was set up to clarify the working of the trap mechanism. It was done mainly in a mature hedgerow on the edge of a playing field in Oxford (v.c. 23), where Arum maculatum was abundant (site A). Three further sites, one in an exposed hedgerow by fields (site B), another in a damp hollow near some young beeches (site C) and a third by paths in scrub woodland (site D), were used for comparison. Sites B, C and D were in the Chiltern region near Reading (v.c. 22). A fifth site, in a wooded garden in Cumbria (v.c. 69) was used in 1990 (site E). The self-incompatibility system was tested using nine inflorescences in site A in 1990. These were covered in semi-opaque paper bags, before opening, on 5 May 1990, and were examined each day after that. When open, two were cross-pollinated artificially using a small paint-brush with pollen from two inflorescences from other plants several metres away, on 10 May. The pollen was dusted on to as many of the female flowers as possible through a hole cut in the spathe chamber wall. Damage to the chamber was minimised and it was difficult to reach the flowers on the opposite side of the column to the hole. Two other inflorescences were artificially self-pollinated in the same way on 14 and 15 May, but using pollen from the same inflorescence. The other five were left unpollinated. Checks on the inflorescences were made regularly thereafter until a final recording of fruit and seed set on 21 June 1990. 15 further inflorescences were bagged and left undisturbed in site E between 1 and 7 May 1990 and checked regularly until 22 July 1990. Observations of the trap mechanism and the visiting insects were made using porous plastic windows inserted into the chamber wall of 30 inflorescences of Arum maculatum in late April and early May 1988 in site A and 35 inflorescences in early May 1990 in site E. Approximately 120 individuals of Psychoda phalaenoides were seen through these, over a period of several days in varying weather conditions. To examine the surface of the spathe and column a series of transverse sections were taken through the spathe chamber wall and the column and examined under a light microscope. The fluid around the stigmas (the “‘nectar’’ of Prime and others) was extracted from approxi- mately 150 flowers on 13 inflorescences using microcapillaries. Measurements of the concentration of sugar, measured as a standard sucrose equivalent, were made on extracts from 5—10 flowers combined using Bellingham and Stanley pocket refractometers adjusted to read small volumes (Corbet 1978; Lack 1982). Extracts from five further inflorescences, one of which had reached the 336 A. J. LACK AND A. DIAZ male phase and the fluid had many pollen grains in it, were analysed for their amino acid content using the method of Yarwood (1989) which can detect trace quantities. Pollen germination was tested by placing fresh pollen on 1) fresh stigmas, 2) in the stigmatic fluid, 3) in fluid from cut stems and 4) in tap water. Germination was recorded after twelve hours. RESULTS The two inflorescences which were artificially cross-pollinated set 20 out of 35 fruits (57%) and 27 out of 39 fruits (69%). These fruits had 1—5 seeds (most 2 or 3) with a mean of 1-8 seeds and 2-6 seeds respectively (excluding fruits that did not set). It was clearly noticeable that most of the fruits which failed to set on the crossed plants were those behind the column from the hole through which pollinations were made. The artificially selfed inflorescences produced 0 out of 34 fruits (0%) and 6 out of 30 fruits (20%) (mean seed number 2-5). None of the bagged and undisturbed inflorescences set any fruit. In many of the selfed and undisturbed inflorescences the fruit started to swell and only stopped about two weeks after inflorescence opening, when they withered. These results show that, normally, A. maculatum is strongly self-incompatible (confirming Prime (1960)). The few fruits set on one selfed inflorescence must have arisen either from a weak self-compatibility in the one plant or from contamination of the self pollen load on the paint brush with a few grains from another plant. Most spathes opened in mid- to late morning, but there was some variation in the timing of flowering events and some opened at almost all times of the day. The smell was normally apparent immediately. Most of the Psychoda phalaenoides entered the chambers around dusk or in the early part of the night (before midnight) during the warm dry weather in which observations were made. When the weather is wetter or cooler they may be more active in daylight hours. They almost invariably landed on the spathe and fell into the trap, usually after slipping several times; only two, out of 55 seen entering, crawled in. Once inside, the flies can and did climb the central column, past the lower whorl of hairs and the male flowers to the upper whorl of hairs. These hairs are very close together and appeared to have a slippery surface, and this prevented the flies from manoeuvring between them and escaping. On the second day no smell was apparent and copious pollen was shed from the anthers which fell into the spathe chamber and on to any insects inside, and the hairs withered. The spathe remained impossible for the flies to climb, but they climbed the spadix past withered hairs to escape. The sections through the spathe showed that papillae cover the open spathe and spadix and extend down on the spathe to half way up the female flowers (as described by Knoll (1926) for A. nigrum), with a transition zone, of approximately 5 mm, of progressively smaller papillae. The papillae pointed at 90° to the wall of the spathe and spadix. Papillae on the spadix extended down to the top of the upper whorl of hairs. The stigmatic fluid was detected in only about two-thirds of inflorescences and varied in quantity up to about 0-3 ul per flower. It was not apparent when the spathe first unfurled, but appeared after about three hours and before most of the Psychoda were caught. It surrounds the feathery stigma, when present, and appears to be resorbed soon after the trap hairs wither. The concentration of sucrose equivalent was between 9% and 12-5% in all samples. The amino acids leucine, isoleucine and/or methionine were detected in the fluid which had pollen in it, but the other four fluid samples had no detectable amino acids. The sugar is only slightly more concentrated than that in the fluid which exuded from cut stems, mainly arising from the phloem —8% sucrose equivalent was recorded from each of three cut stems. Despite several periods of observing Psychoda inside the spathe chamber we never observed them taking the stigmatic fluid (contrary to Prime’s (1960) claim) and many inflorescences had fluid and Psychoda present. Psychoda species are not known to feed during their adult lives (P. Withers, pers. comm. 1990). We must conclude either that the fluid has no function in A. maculatum, or that its function is nothing to do with a food reward for the insect visitors. It was often noticeable that, in an inflorescence’s male phase, much pollen was trapped in the fluid and many grains had germinated. Observations on the germination of pollen grains showed that they germinate readily in the stigmatic fluid, including that from the same inflorescence, and on the stigmas and in tap water, although not in the fluid from cut stems. Baker et al. (1973) analysed the contents of stigmatic exudates of 39 species of various plant families (in California and Costa Rica) and summarised previous accounts. They found amino acids POLLINATION OF ARUM MACULATUM ST in exudates from 38 of the 39 species as well as lipids and other substances. They interpreted the function of the exudates as, in some plants, nutritional for the pollinating insects and, in others, as direct aids in the pollination process i.e. as sites for pollen germination and as a sticky fluid to make pollen adhere to the insects. They, and Eisikowitch er al. (1990), studying the highly specialised Asclepias syriaca, found that pollen germinated in exudate of up to 20% sucrose but not in higher concentrations. Eisikowitch et al. (1990) showed that the tubes burst in concentrations of 5% or less. The concentrations required for pollen germination, therefore, are much lower than those found in normal nectar, which acts as a food reward (Corbet 1978). It seems that the stigmatic fluid in A. maculatum is not nectar in any recognised sense, but that its function is as a trap for pollen grains, as a Psychoda crawls over the inflorescence, and as a site for pollen germination. It may also help pollen to adhere to a Psychoda if some is brushed on to it in the spathe chamber. Fluid is present around the stigmas of other members of the Araceae (a large quantity in some), and, in some, appears to act as a food source for visiting pollinators, though very little study has been done (Bown 1988). In Arum hygrophilum in Israel, Koach (1985) reported that Psychoda cinerea Banks males are trapped and may live longer inside Arum because of high humidity maintained by the stigmatic fluid (5% sucrose) and possibly from nutrition. THE IMPORTANCE OF THE TRAP MECHANISM IN POLLINATION Despite the elaborate trap structure and apparent effectiveness of the mechanism in Arum maculatum, the importance of it for pollination in the species was brought seriously into question by Schmucker (1925), whose experiments were reported by Prime (1960). Schmucker (1925) removed the spadix and/or the spathe from plants of A. maculatum, but found no effect on fruit or seed set from any of these mutilations (Table 1). The only detrimental effect on fruit set that he found was from removal of the upper whorl of hairs in addition to the spathe and spadix, and this he attributed to a wound response. Meeuse (1978) studied several members of the Araceae and found that damage to the male flowers had an inhibitory effect on flowering (perhaps hormonal). He did not refer to Schmucker (1925), but his observations back up the possibility of a wound influencing fruit set. Although Schmucker’s (1925) conclusions were most unexpected, no further work on this appears to have been done. The implication from Schmucker’s work was that the trap mechanism was a vestige from the species’ evolutionary past, important perhaps in other members of the genus, but not in A. maculatum. This investigation was set up to see whether we could duplicate and elaborate on Schmucker’s experiments. Three aspects of the trap mechanism were studied: 1. whether various treatments to the trap mechanism had an effect on numbers of insects caught; 2. what effect the treatments and the numbers of insects caught had on seed and fruit set of the inflorescences; and 3. how many pollen grains an individual Psychoda may carry. TABLE 1. NUMBER OF FRUITS AND SEEDS SET BY ARUM MACULATUM IN SCHMUCKER’S EXPERIMENTS (from Schmucker 1925) % setting Mean number of Sample size seed fruits per spike Open pollinated 47 38 15 Spathe removed 31 32 12 Spadix removed 33 43 14 Spathe and spadix removed 43 26 10 Spathe, spadix and hairs removed 29 17 5 338 A. J. LACK AND A. DIAZ METHODS Inflorescences of Arum maculatum were treated in the following ways: 1. Spathe removed 2. Spadix removed 3. Spathe and spadix removed 4. Hole cut in the spathe chamber wall 5. Open-pollinated (control) It was found that earwigs sometimes entered the chamber through the hole cut for treatment 4; all inflorescences in this treatment were checked two days after flowering for any damage, and damaged inflorescences were not used. All treatments were made before 07.00 hours and before the spathe unfurled. Treated inflorescences were at least 1-5 m apart, and were labelled with plastic tags. Sample sizes varied mainly owing to problems of diseased, damaged or otherwise unusable inflorescences and:are given in the results. Treatments 1, 2 and 5 were carried out on inflorescences in site A which opened before 11.00 hours on 13 May 1988. The insects from each inflorescence were collected after 22.30 hours the same day, by placing a cotton wool plug soaked in ethyl acetate over the entrance to the chamber before extracting the insects. This ensured that the majority of the insects that the spike would catch had been caught. Further flower spikes were chosen at random between 24 April and 8 May 1988 and subjected to one of the five treatments. They were then left to mature undisturbed over the next nine weeks. Their condition was checked regularly and any inflorescence that was damaged or diseased was discarded from the sample groups. The maturing fruits were collected during June and July as they began to turn yellow and mature, but before any frugivores (mainly blackbirds, Snow & Snow 1988) ate them. The number of seeds, fruits and aborted fruits (unswollen ovaries) were recorded. A direct measure of the effect of number of insects caught on fruiting success was made between 25 April and 12 May 1989. Insects were collected from open pollinated inflorescences towards the end of the female phase, on the second morning of spathe opening between 07.00 and 08.30 hours. Each study inflorescence was checked to see that the anthers had not dehisced and no insect had escaped. Inflorescences were chosen to show a range of numbers of insects caught. The insects were collected by placing a glass sample tube over a hole cut in the spathe chamber wall. No ethyl acetate was used in case of an effect on fruit maturation. The insects were then counted and the inflorescence left to develop. Checks were made after two days that intruders had not entered through the hole and damaged the inflorescence and further checks were made regularly for any damage. Numbers of fruits and seeds set were measured as described above. In 1990, to investigate whether numbers of insect visitors limited fruit set, 28 spikes in site E, which were left open to insects, were artificially cross-pollinated through holes in the spathe chamber with pollen from three other plants, as described in the self-incompatibility test. 15 individual Psychoda phalaenoides were caught as they entered an Arum spathe chamber, in site E. The number of pollen grains that they were carrying was counted. RESULTS Removal of the spathe or removal of the spadix resulted in fewer insects being caught than in intact inflorescences (Table 2, both comparisons P<0.001, Kolmogorov-Smirnov test). There was no difference between these two treatments. In site A, the percentage of spikes that matured any seed (no fruits were set without seeds) TABLE 2. NUMBERS OF INTACT INFLORESCENCES OF ARUM MACULATUM AND THOSE WITH SPATHE OR SPADIX REMOVED WHICH HAD CAUGHT PARTICULAR NUMBERS OF PSYCHODA APPROXIMATELY TWELVE HOURS AFTER OPENING Sample Numbers of Psychoda trapped size 0 1 2 3 4 m) 6 7 8 >8 Intact 25 3 2 I 0 2 2 3 2 Lng edt Spathe removed 18 13 3 2 = Spadix removed 24 19 l POLLINATION OF ARUM MACULATUM 339 TABLE 3. PERCENTAGE OF SPIKES OF ARUM MACULATUM THAT SET ANY SEED AND THE MEAN NUMBER OF SEEDS SET IN EACH OF FIVE TREATMENTS Site A Sites B, C, D All sites Sample No. setting Sample No. setting Mean no. size seed (%) size seed (%) seeds* Open pollinated 143 57 (40) 111 72 (65) 33-0 Spathe removed 68 12 (18) 48 4 (8) 23-4 Spadix removed a7 10 (18) 38 8 (21) 33-1 Spathe and spadix removed 30 0 Zz 1 (4) 16 Hole in chamber wall 50 10 (20) 18 8 (44) 13-1 * excluding those which set no seed differed considerably between the test groups (Table 3). Those that had been manipulated in any way all had a smaller proportion of spikes setting seed than open pollinated inflorescences (all P<0.01, Nx” tests). There were no significant differences between any of the treatments except for those from which both spathe and spadix were removed. Only one of these set any seed, significantly less than other treatments (all P<0.05, “y? tests). The percentage of open pollinated inflorescences which set seed was greater in sites B, C and D, pooled, than in site A (P<0.001, Nx? test) but differences between manipulated and open pollinated inflorescences were similar to those at site A. Of those spikes in site A that matured at least one fruit (Table 3), there was no significant difference in the number of seeds set by spikes with either the spathe or spadix removed compared with the open pollinated group. It seems that, if it is fertilised at all, all potential fruits are likely to be fertilised. Those spikes with a hole in the chamber wall, however, set fewer seed (P<0.002, modified t-tests, not assuming equal population variances). Pooled results from sites B, C and D showed almost exactly the same pattern as site A. Table 4 shows the results obtained from all four sites together for percentage of potential fruits matured (sample sizes were too small for meaningful comparisons from site A separately). Removal of the spathe and cutting a hole in the chamber wall reduced percentage fruit set (P<0.01, Kolmogorov-Smirnov test) but removal of the spadix did not. Fruiting success did not appear to be related to number of insects caught (Table 5). 82% of the inflorescences that caught any insects set seed and it seems that just one pollen-bearing insect was necessary for full fruit set to occur. Of the 28 artificially cross-pollinated inflorescences, 27 (96%) set fruit. The numbers of A. maculatum pollen grains recorded on individual Psychoda were (in increasing erder)i0,.0; 2,56, 70, 84,93, 101; 111.1247 127,133, 2150, >150, >150. The first two of these fifteen and, perhaps, the third (20%) were probably visiting their first Arum inflorescence, whereas the others had clearly come from another one. TABLE 4. NUMBERS OF INFLORESCENCES OF ARUM MACULATUM IN VARIOUS TREATMENTS (PERCENTAGES IN BRACKETS) WHICH MATURED PARTICULAR PROPORTIONS OF POTENTIAL FRUITS Those which matured no fruits excluded from these results. Percentage of potential fruits matured Sample size <20 20-40 40-60 60-80 80-99 100 Open pollinated 129 3412) 11 (9) 16 (12) 2217) 58 (45) 19(15) Spathe removed 16 Ly (6) SG) 4 (25) 3 (19) 0 3 (19) Spadix removed 16 0 1 (6) 212) 4 (24) 8 (47) 1 (6) Hole in chamber wall 19 8 (42) 4 (21) 3 (16) 0 Diy 2 Git) 340 A. J. LACK AND A. DIAZ TABLE 5. NUMBERS OF FRUITS AND SEEDS SET BY PLANTS OF ARUM MACULATUM WHICH HAD CAUGHT PARTICULAR NUMBERS OF PSYCHODA PHALAENOIDES Number of Sample No. setting Mean no. of Mean % P. phalaenoides caught size any seed seeds set* fruit set* 0 1 0 — — 1 6 5 31 65 2 3 1 47 100 3 - 3 34 60 - 5 5 25 66 5 2 1 54 100 6 2 2 19 57 7 0 _ — — 8 3 2 28 71 >8 8 8 31 72 * excluding those which set no fruits or seeds DISCUSSION In all sites, less than 100% of inflorescences set fruit, and in site A in 1988 only 40%. Some inflorescences in all study sites caught no insects and others caught one or very few, and the failure to fruit was related to the availability of Psychoda phalaenoides. This was further confirmed by the fact that all but one of the artificially crossed inflorescences set fruit. All the evidence from this investigation leads us to the conclusion that fruit set, in any one year at least, is limited by the numbers of pollinating insects. Fruits set in one year can, however, affect fruit set in the next year (a well-known phenomenon in fruit trees), so a definitive test for what limits fruit set would require following the same plants through two seasons. Possible limitation of fruit set by pollinator availability is recorded in a few species, but only definitively (i.e. as described above) in one, another aroid, Arisaema triphylla (Zimmerman 1988). Prime (1960) stated that, from preliminary observations, about 10% of Arum maculatum tubers died each year, but that flowering had no apparent effect on the probability of death. Any disruption to the trap mechanism led to many fewer insects being caught (frequently none) and to lower numbers of inflorescences setting any fruit or seed. In fact, because of the precision of the trap mechanism and the function of all parts of the flower spike and spathe, it was surprising that inflorescences that had parts removed ever caught any insects. On those from which the spathe was removed the spadix was still, clearly, an attractant and one Psychoda was observed crawling around the cut edge of the spathe before slipping down and being trapped. On those from which the spadix was removed it is possible that the flies were attracted by the pale colour of the spathe since they appeared to be attracted by light surfaces (personal observation). Although most spikes which had their spathe, spadix or both removed did not set any seed, of those that set at least one, there was very little difference in numbers of fruit set between the mutilated and intact inflorescences. Only those with the hole cut in the chamber wall set many fewer fruits. It seems that one pollen-bearing Psychoda may well be adequate to fertilise all the female flowers successfully so long as it has travelled from another clone; an individual fly can certainly carry enough pollen grains. Since the flies are normally trapped in the chamber for 18-24 hours, and move about during this time, there is plenty of opportunity to deposit the pollen on all the female flowers. In those chambers in which a hole was cut, the flies normally found their way out very quickly and flew off, so it is not surprising to find big reductions in fruit and seed set in these plants, since the flies will not have crawled over all the female flowers. Psychoda species live as adults for up to seven days (P. Withers, pers. comm. 1990) and this means that a significant proportion of flies trapped in any one inflorescence (perhaps up to 25%) are likely to be first-time visitors; our results for numbers of pollen grains carried confirm this. Others may come from another inflorescence in the same clone, so the pollen that they are carrying is not viable. These are likely to be the reasons for the lack of any fruit set by some inflorescences which trapped small numbers of insects. Pollinator availability seems to be the main limiting factor in fruit set in this study, but other Sere wey 8 OO 20 an ae i i i ee oo Pa a eee nepememnenee that ete POLLINATION OF ARUM MACULATUM 34] factors may also be important. Some decay of fruiting spikes was noted, particularly in 1988 which was a wet, cool summer, but these were not included in the results. Resource limitation, often important in limiting fruit set (Stephenson 1981), did not appear to play any part in this study, though observations over two years would be needed to confirm this. Predation was not important in our study areas but Snow & Snow (1988) noted that many flower spikes of A. maculatum in their Chiltern study sites were eaten by deer before setting any fruit. The results presented here are in clear contradiction of the results of Schmucker (1925). By chance, it appears that our site A was similar to his site in the overall proportion of open pollinated spikes setting fruit (38% of his spikes set fruit, 40% of ours) and, therefore, is likely to be comparable in pollinator availability and associated factors. This suggests that there are differences in the experimental methods used. Schmucker (1925) was not totally clear about his methods but stated that the treatments were made on spathes that were just about to open. He probably waited until just as the spathes opened to do the tests since this is likely to cause the least damage. We know from our study, however, that this stage may well be too late to prevent pollination. The spadix may smell before the spathe opens and those spathes which had only opened a fraction were sometimes found to have caught insects, though this was only noticeable if the chamber was cut open. We must, therefore, contest Schmucker’s conclusion that the trap mechanism is not important for fruiting in A. maculatum, since our results show clearly that all parts are important. It does seem that, for fruit set, it is not important how may insects are caught and that just one Psychoda phalaenoides, bearing pollen from another plant, may be adequate for full seed set, but this is only part of the pollination story. The other part, successful dissemination of pollen, is much harder to study and we can only make some suggestions about this in A. maculatum. Since pollinating insects are trapped by the Arum inflorescences for some hours, it is likely that almost all of the pollen that is taken out of one inflorescence by any one Psychoda will be deposited on the next one it visits — pollen carry-over will be minimal. One trapped Psychoda will probably mean that a significant number of seeds are fathered on only one other plant, or even none if the next inflorescence it visits is part of the same clone. The more Psychoda that are trapped, therefore, the more inflorescences will receive pollen from that individual plant and the more successful that plant will be as a father. This is true in most plants (Bertin 1988) and it suggests that successful trapping of as many Psychoda as possible will be important to ensure pollen dispersal. ACKNOWLEDGMENTS We are most grateful to Ken Howells for help with the sectioning, Graham Barrett for help with the amino acid analysis, Phil Withers for information about Psychoda species and Amots Dafni for searching out the Israeli work. REFERENCES Baker, H. G., BAKER, I. & Opter, P. A. (1973). Stigmatic exudates and pollination, in BRANTJES, N. B. M. & LINSKENS, H. F., eds. Pollination and dispersal, pp. 47-60. University of Nijmegen, The Netherlands. BertTIN, R. I. (1988). Paternity in plants, in Loverr Doust, J. & Lovetr Doust, L., eds. Plant reproductive ecology, pp. 30-59. Oxford. Bown, D. (1988). Aroids. London. Corset, S. A. (1978). Bec visits and the nectar of Echium vulgare L. and Sinapis alba L. Ecol. Ent. 3: 25-37. Dormer, K. J. (1960). The truth about pollination in Arum. New Phytol. 59: 298-301. ErsikowiTcH, D., Kevan, P. G. & LAcHANCcE, M.-A. (1990). The nectar-inhabiting yeasts and their effect on pollen germination in common milkweed, Asclepias syriaca L. Israel J. Bot. 39: 217-225. FAEGRI, K. & VAN DER PUL, L. (1979). The principles of pollination ecology. Oxford. KNOLL, F. (1926). Die Arum-Blutenstande und ihre Besucher (Insekten und Blumen IV). AbjA. zool.-bot. Ges. Wien 12: 379-481. Koacu, J. (1985). Bio-ecological studies of flowering and pollination in Israeli Araceae. Unpublished Ph.D. thesis, University of Tel-Aviv (in Hebrew, summary in English). Lack, A. J. (1982). Competition for pollinators in the ecology of Centaurea scabiosa L. and Centaurea nigra L. II. Observations on nectar production. New Phytol. 91: 309-320. MeeuseE, B. J. D. (1961). The story of pollination. New York. 342 A. J. LACK AND A. DIAZ MeeusE, B. J. D. (1978). The physiology of some sapromyophilous flowers, in RicHarps, A. J., ed. The pollination of flowers by insects, pp. 97-104. London. PrimE, C. T. (1960). Lords and Ladies. London. Prime, C. T. (1980). Arum L., in TuTin, T. G. et al. eds. Flora Europaea 5: 269-271. Cambridge. Proctor, M. & YEO, P. (1973). The pollination of flowers. London. SCHMUCKER, T. (1925). Beitrage zur Biologie und Physiologie von Arum maculatum. Flora 18: 460-475. Snow, B. & SNow, D. (1988). Birds and berries. Calton, Staffs. STEPHENSON, A. G. (1981). Flower and fruit abortion: proximate causes and ultimate functions. Ann. Rev. Ecol. Syst. 12: 253-279. Yarwoop, A. (1989). Manual methods of protein sequencing, in FINDLAy, J. B. C. & Geisow, M. J., eds. Protein sequencing, a practical approach, pp. 119-145. Oxford. ZIMMERMAN, M. (1988). Nectar production, flowering phenology, and strategies for pollination, in Lovett Doust, J. & Lovetr Doust, L., eds. Plant reproductive ecology, pp. 157-178. Oxford. & F (Accepted December 1990) Watsonia, 18, 343-350 (1991) 343 The identity of Euphrasia officinalis L. and its nomenclatural implications A. J. SILVERSIDE Dept. of Biology, Paisley College, Paisley, Renfrewshire, PAI 2BE ABSTRACT The treatment of the name Euphrasia officinalis L. (Scrophulariaceae) as a nomen ambiguum is rejected. Examination of the lectotype material shows that it matches E. rostkoviana Hayne subsp. fennica (Kihlman) Karlsson. The name E. officinalis takes priority and the following infraspecific taxa are recognised: E. officinalis subsp. officinalis, E. officinalis subsp. rostkoviana (Hayne) F. Townsend, E. officinalis subsp. monticola Silverside nom. nov. (= E. montana Jordan) and E. officinalis subsp. anglica (Pugsl.) Silverside comb. et stat. nov. The taxonomic treatment of other species in Euphrasia series Euphrasia is discussed and the implications for the infrageneric arrangement in Euphrasia are noted. INTRODUCTION The name Euphrasia officinalis L. has been generally treated as a nomen ambiguum by those who recognise an aggregate of numerous species, despite the existence of original Linnaean material. There are two sheets of Euphrasia, as the genus is now circumscribed, in the Linnaean Herbarium (LINN). One represents the distinctive Italian species, FE. tricuspidata L., the other (sheet 759.2), consisting of three specimens, represents E. officinalis. Two of the specimens on the latter sheet have been generally considered to be referable to the taxon currently known as E. rostkoviana Hayne, while the third, of Russian origin (see below), has been regarded as E. nemorosa (Pers.) Wallr. (including E. curta (Fries) Wettst.) (Townsend 1867; Pugsley 1930), or as E. stricta J. P. Wolff ex J. F. Lehm. (Yeo 1972). Sell & Yeo (1970) partially lectotypified E. officinalis by equating it with the element regarded as E. rostkoviana, but nevertheless rejected the name E. officinalis as a nomen ambiguum, a view subsequently reiterated by Yeo (1972, 1978). This rejection of the name E. officinalis was presumably based on Article 69 of the [International Code of Botanical Nomenclature, which formerly stated, ““A name is to be rejected if it is used in different senses and has so become a long-persistent source of error”’ (Stafleu et al. 1972). For those who took a broad view of E. officinalis and who would consider the two elements of the Linnaean sheet as representing variants of a single species, the application of the name was clear. It could be used for the aggregate species without further qualification. For those who recognised a number of species within the Linnaean concept, the name E. officinalis was not to be used for any of the segregates. While this interpretation of Article 69 was perhaps questionable, the result of this interpretation has been unambiguous and convenient. Unfortunately, from this viewpoint, such a treatment is no longer tenable. Article 69 was altered at the International Botanical Congress in Leningrad in 1975 (Stafleu et al. 1978) and a name may now be rejected under this article only if “it has been widely and persistently used for a taxon or taxa not including its type” (Article 69.1) (Greuter et al. 1988). Further, a name may not be summarily rejected under this article; a name becomes formally rejected only after consideration by the General Committee established by the International Association for Plant Taxonomy and its placement on the list of nomina rejicienda (Articles 69.1, 69.2). To date, E. officinalis has not been so treated, and it is clear that there is no basis for doing so. While it is true that some early literature used the name in a sense that excluded the E. rostkoviana group, such misapplication has hardly been widespread or persistent. Virtually all use of E. officinalis has been in a broad sense that includes the type material. To interpret Article 69 in any other way would require the rejection of the names of all taxa that have been subdivided. The term nomen ambiguum has no formal meaning 344 A. J. SILVERSIDE under the /.C.B.N. and, in its usual meaning, it can hardly be applied to a name of undisputed modern application and for which there is accessible and reasonably good type material. It follows that the name E. officinalis is valid and would have priority over the name E. rostkoviana if they were shown to refer to the same taxon. This situation has already been briefly pointed out by Barker (1982), but seems otherwise to have been generally overlooked or ignored. I have been as guilty as anyone for long maintaining a treatment that I knew to be invalid, but in the course of determining material and in preparing a semi-popular account of the British diploid taxa (Silverside 1990), I have increasingly come to doubt the validity of maintaining FE. anglica Pugsl. as a separate species from E. rostkoviana. It has been clear that a new combination cannot be published until the nomenclature of the E. rostkoviana group has been revised, and a re-examination of the Linnaean material has seemed prudent. It should be noted that although I have referred above to the publication of Sell & Yeo (1970) in relation to the lectotypification of E. officinalis, they did not, in fact, indicate a specimen, either in their publication or, apparently, at LINN. While there can be no reasonable doubt as to which of Linnaeus’s specimens they considered as representing E. rostkoviana, I consider the unequivocal lectotypification to have been provided by Yeo (1978, p. 237), when he excluded the specimen with seven pairs of branches and eglandular foliar hairs. While this still leaves a choice of two specimens, I believe that these represent a single collection, as discussed below, and that they count as “‘small herbaceous plants’’, so being jointly acceptable as the type under Article 9.1. EXAMINATION OF THE LINNAEAN MATERIAL OF E. OFFICINALIS Although the two specimens that constitute the lectotype of E. officinalis have been declared to represent the taxon generally known as E. rostkoviana, it should be borne in mind that Yeo (1972, 1978) incorporated two taxa of eastern Scandinavia and northern Russia, FE. fennica Kihlman and E. onegensis Cajander, within his concept of E. rostkoviana subsp. rostkoviana. These two taxa have been variously treated in Scandinavian literature, with recent viewpoints being to treat them both as named varieties of E. rostkoviana (Jalas 1977) or to treat E. fennica as a subspecies of E. rostkoviana, and E. onegensis as a taxon of uncertain status, perhaps closer to E. hirtella Jordan ex Reuter (Karlsson 1982). The Linnaean specimens were examined with these taxonomic and nomenclatural implications in mind. As has been stated by previous workers, the Linnaean sheet 759.2 consists of three specimens. The left- and right-hand specimens bear long, glandular hairs while the central specimen is apparently eglandular, possesses seven pairs of branches and is clearly the specimen excluded from the lectotype by Yeo (1978). The following descriptions were necessarily made without risk of damage to the material and measurements are in some cases approximate or insufficient to allow ranges to be quoted. THE CENTRAL SPECIMEN Plant exceeding 18 cm in height (specimen curved in pressing), erect, with seven pairs of ascending branches, the lowest almost equalling the main stem; secondary branches absent; flowering commencing at node 12. Foliage with no trace of original colour but drying darker than the other two specimens on the sheet; undersides of leaves apparently paler than the uppersides; cauline leaves missing or fragmentary, with all teeth apparently antrorse and acutely triangular. Lower cauline internodes c. 10-12 mm; uppermost cauline internode 14 mm, its subtending leaf approximately 10 mm in length; lowest floral internode 11 mm, its subtending floral leaf c. 6 mm. Floral leaves broadly ovate to trullate, with mostly five pairs of antrorse, finely acute to aristate teeth. Foliage and calyces sparingly setose, lacking stalked glands. Calyx teeth linear-triangular, aristate. Corolla length (to tip of upper lip) c. 6-5 mm; lower lips with rather narrow, spreading lobes. Mature capsules c. 5-0 mm in length, about equalling calyx and subtending floral leaf; apex emarginate, ciliate; width 1-9 mm. The symbol € is written on the sheet next to this specimen, which, following Savage (1945), indicates that the specimen is probably of Russian origin. Pugsley (1930) referred to the identifications of this specimen as E. nemorosa but considered that it was more probably E. curta. As the specimen is only sparingly setose, he presumably had in mind aT EUPHRASIA OFFICINALIS AND ITS NOMENCLATURE 345 the subglabrous variants that have been recognised as E. curta var. glabrescens Wettst., though Pugsley did not, himself, accept this taxon. Yeo (1971) considered that there is no difference between E. curta var. glabrescens and E. nemorosa and that typical, Scandinavian E. curta is also best regarded as a hairy variant of E. nemorosa, though some populations in Britain, the Faeroes and Iceland he recognised as a separate species, EF. ostenfeldii (Pugsl.) Yeo. The Linnaean specimen is certainly not E. ostenfeldii and there seems no basis to identify it with E. nemorosa var. curta Fries, as reinstated and illustrated by Hartl (1972). There remains the choice between E. nemorosa and E. stricta, which appear to be the only species that should be considered. The specimen certainly bears a strong resemblance to E. nemorosa, particularly to the rather large, strongly branched variants sometimes met on waste ground. The corolla size indicates E. nemorosa, being considerably smaller than would be expected in E. stricta. However, the finely acute to aristate and apparently uniformly antrorse toothing of the leaves indicates E. stricta and the relatively narrow capsules perhaps also support this view. The leaves of E. stricta are commonly markedly paler beneath and there is perhaps some indication that this was true of the Linnaean specimen. Both taxa extend into the Soviet Union and since both tend to be outcrossing species and have rather similar habitat requirements, the possibility that the specimen came from a hybrid population cannot be ruled out. Karlsson (1984) notes the ready formation of hybrid swarms between the two species on the island of Gotland, Sweden. While, on balance, the specimen seems nearer to E. stricta, it is not, in my opinion, reliably identifiable, and amply illustrates the difficulty of dealing with single plants. Fortunately, its identity no longer has any nomenclatural implication. THE LEFT-HAND SPECIMEN Plant 28-2 cm in height, erect, with three pairs of branches in upper half; branches straight, ascending at about 30° from vertical, distinctly shorter than main stem; secondary branches absent; flowering apparently commencing at node 9. Foliage with no trace of original colour, drying light brown; cauline leaves missing. Lower cauline internodes long, exceeding 3 cm; lowest floral internode 1-2 cm. Lower floral leaves missing; middle and upper floral leaves reaching maximum length of 6 mm, broadly rhomboid-ovate, with five pairs of teeth on examinable leaves; toothing finely acute to aristate; all teeth antrorse. Foliage with some setose hairs; foliage, calyces and upper parts of stem also clothed in long-stalked, waved glandular hairs; lengths of glandular hairs variable, reaching 0-5 (—0-7) mm; stalks multicellular (to 4-celled), transparent; glandular heads globose to ellipsoid, transparent or now brown. Calyx teeth linear-triangular, + aristate. Corolla length (to tip of upper lip, pressed) at least 8 mm; lower lip exceeding upper, with spreading lobes, width at least 5:5 mm; no glands noted on exposed part of corolla tube. Capsules variable in size, longest 5-2 mm, equalling or exceeding floral leaves, shorter than to equalling to exceeding calyx, width 2-1 mm; apex emarginate, ciliate. THE RIGHT-HAND SPECIMEN Plant 29-3 cm in height, erect, with two pairs of upcurved branches emerging at an angle of about 45° from the mid-part of the stem and distinctly shorter than the main stem; secondary branches absent; flowering probably commencing at node 9, but specimen has three intercalary nodes below this, now bare. Cauline leaves missing. Lower cauline internodes c. 2:5 cm; lowest floral internode 1-4 cm. Lower floral leaves to 9-5 mm in length, broadly rhomboid-ovate, with seven pairs of finely acute teeth, the lowest pair patent, the rest antrorse; floral leaves diminishing in size upwards; upper floral leaves with six pairs of teeth, all antrorse. Calyx teeth linear-triangular, + aristate. Corollas appear equal in size to those of the left-hand specimen, no accurate measurements being possible. No mature capsules are present. Foliage colour and details of setose and glandular indumentum are as described for the left-hand specimen. DISCUSSION OF THE LEFT- AND RIGHT-HAND SPECIMENS No indication of the origin of the specimens is given, but the general facies of the two specimens would support their being from a single collection. They differ somewhat in their branching, but branches tend to be rather flexuous and variable in the E. rostkoviana group, to which the specimens undoubtedly belong, and I consider these differences to be unimportant and no more than would be expected within a single population. So far as it is possible to compare foliage and corolla characters, 346 A J..STIEVERSIDE there is a close similarity between the two plants. In the following discussion, I assume that the two specimens are of common origin. In the absence of any indication to the contrary, they would seem most likely to be Swedish, a view for which there is support from the fine detail of their morphology, as discussed below. At first sight, the specimens match E. rostkoviana closely. They are more branched than most material of E. fennica that I have seen but, on the other hand, they match the photograph of the lectotype material of E. fennica, published by Jalas & Kukkonen (1973), extremely well. The other possibility cited above, E. onegensis, can apparently be discounted. Little information was provided with its original description (Cajander 1901) and I know the taxon only from the photograph of lectotype material, again published by Jalas & Kukkonen (1973). However, this photograph shows a distinctive plant of aestival habit, with markedly small corollas, low flowering nodes, and large upper floral leaves, giving the developing plants a capitate appearance (much as in well grown E. frigida Pugsl.), and with leaves with few pairs of rather blunt teeth. Karlsson (1982) has investigated Finnish records of E. onegensis and has shown that they are referable to E. fennica, leaving E. onegensis as a taxon currently known only in the north-western part of the Soviet Union and needing further study. The Linnaean specimens clearly differ from the lectotype of E. onegensis and from Karlsson’s redescription in such features as corolla size, lowest flowering node and shape and toothing of the floral leaves. The differences between E. rostkoviana, particularly with regard to British and Irish material, and E. fennica seem, at first, so slight, that Yeo’s (1972) inclusion of the latter taxon under E. rostkoviana subsp. rostkoviana has appeared, to me, fully justified. However, Karlsson’s (1982) meticulous study of the complex in Sweden forces a reassessment of these taxa. Karlsson has shown that E. fennica, which he recognises as a subspecies of E. rostkoviana, is widespread in eastern Sweden, Finland and the adjacent part of the Soviet Union, often on relatively poor soils. E. rostkoviana (sensu stricto), by contrast, is largely restricted to two calcareous regions of Sweden, and is a recent and casual introduction in Finland. As well as in its slender, usually autumnal habit, E. fennica differs from Scandinavian and continental FE. rostkoviana in its floral leaves, with more numerous (between five and eight) pairs of finer, narrower teeth and in the narrower mid-lobes of its lower corolla lips. Even in Karlsson’s figures, there is only limited evidence of disjunction in these characters, but he argues convincingly for recognition of E. fennica at subspecific rank, and even that evolutionary differentiation of E. fennica from E. rostkoviana s.s. occurred at an earlier time than the differentiation of E. rostkoviana subsp. montana from the latter taxon. The opinion of a respected authority on the genus, based on detailed work in his own area, cannot be lightly disregarded and I accept his conclusions here. Linnaeus’s specimens fit E. fennica in habit and the floral leaves of the right-hand specimen, with their seven pairs of finely acute teeth, clearly fall within the range of E. fennica and outside the range of most or all E. rostkoviana from elsewhere. The single measurement of a width of a lower corolla lip (equalling or exceeding 5-5 mm, left-hand specimen) would infer a mid-lobe width unlikely to exceed 3 mm, which, while not conclusive, again strongly indicates E. fennica. I conclude that the lectotype material of E. officinalis matches the taxon currently known as E. rostkoviana subsp. fennica (Kihlman) Karlsson. It may be noted that Karlsson found that FE. fennica, in Sweden, occurs in greatest abundance in the vicinity of Uppsala, an area in which Linnaeus collected extensively (Blunt 1971) and hence the most likely origin of his material. NOMENCLATURE Since the name E. officinalis L. takes priority over both E. rostkoviana Hayne and E. fennica Kihlman, the following nomenclatural arrangement is required. Euphrasia officinalis L., Sp. Pl. 604 (1753). Lectotype: Sheet 759.2, Herb. Linnaeus (LINN), excluding specimen with seven pairs of branches (Yeo 1978). 1) E. officinalis L. subsp. officinalis E.. fennica Kihlman, in Mela, Suomen Koulukasvio, 4th ed. , 247 (1899). EUPHRASIA OFFICINALIS AND ITS NOMENCLATURE 347 E. rostkoviana Hayne var. fennica (Kihlman) Jalas in Ann. Bot. Fenn. 14: 191 (1977), E. rostkoviana Hayne subsp. fennica (Kihlman) Karlsson in Vaxtekologiska Studier 15: 42 (1982). Fuller synonymies are provided by Jalas & Kukkonen (1973) and Karlsson (1982). Distribution: Eastern Fennoscandia and adjacent parts of the Soviet Union. As inferred in the foregoing discussion, some British and Irish populations resemble this subspecies, and EF. fennica was reported from Exmoor and Galway by Bucknall (1917). Pugsley (1930) doubted these identifications, but mentioned other Irish material that resembled it. Material I have seen matches subsp. rostkoviana in the floral leaves with four to five pairs of rather coarse teeth and is undoubtedly referable to that subspecies, though showing some differentiation from continental populations. 2) E. officinalis L. subsp. rostkoviana (Hayne) F. Townsend, inJ. Bot., Lond. 22: 165 (1884). (As E. officinalis L. *E. Rostkoviana, but with indication that the rank of subspecies was intended.) LeEcTotyPE: not designated. The plate accompanying Hayne’s description shows a plant possessing floral leaves with five pairs of coarse teeth, consistent with the taxonomic views adopted in this paper. E. rostkoviana Hayne, Getreue Darstellung und Beschreibung der in der Arzneykunde Gebrdauch- lichen Gewdchse 9: t.7 (1825). Distribution: throughout most of Europe, including Britain and Ireland; rare in Scandinavia. It is pleasing that what I take to be the correct combination at subspecific rank maintains the continuity of use of the familiar epithet, rostkoviana. However, there is a potential problem in the existence of the combination Euphrasia officinalis L. A) E. pratensis Fries, Novitiae florae Suecicae, edit. altera 198 (1828), which could be regarded as invalidly published (.C. B.N. Articles 32.1b and 33.4) but which may also be regarded as legitimate under Article 24.4. It is clear in the context of Fries’ account that his use of upper case lettering was to denote a concept above that of variety, but which he clearly stated he did not wish to distinguish as separate species. His concept seems to match that of the subspecies but is not explicitly stated. Under Articles 24.4 and 35.2, he can thus be taken to have validly published an infraspecific taxon of undefined rank, under which he lists, as a synonym, E. rostkoviana (as ‘“‘Rostkowiana’’). He separated his “‘A) E. pratensis” from his other subdivision, ““B) E. montana’’, by the former being glandular hairy. As well as his very brief description of pratensis and citation of E. rostkoviana, he also referred to the pre-Linnaean description of Haller (1745). Reference to Haller’s work (p. 240) shows that under ‘“‘Euphrasia officinarum’”’ he listed two taxa, “‘Euphrasia ramosa, pratensis, flore albo” and ‘“‘Euphrasia minus ramosa, flore ex caeruleo purpurascente’’. The first of these would appear to correspond to Fries’ pratensis, while the second must then correspond to Fries’ parallel reference to Haller under ““B) E. montana’’. In fact, this is of little help in interpreting the application of Fries’ names. Although Fries included E. rostkoviana in his concept of “‘A) E. pratensis’’, it seems clear that he must also have included the glandular component of the complex that Karlsson (1976) groups under E. stricta and which Yeo (1972, 1978) splits between E. stricta (including E. brevipila Burnat & Gremli ex Gremli) and E. arctica Lange ex Rostrup subsp. tenuis (Brenner) Yeo. Populations of this complex are widespread and locally abundant in Scandinavia and Fries could hardly have overlooked these attractive and conspicuous plants. Taking the whole of Fries’ protologue, there is no justification, at present, for equating his pratensis with E. rostkoviana and I consider his “‘A) E. pratensis” to be anomen dubium. Should a future lectotypification of pratensis be based on material referable to E. rostkoviana, Fries’ combination, even if validly published, would still not, itself, have any status in questions of priority (Article 35.2). Nevertheless, the epithet pratensis was widely taken up, numerous citations being given by Wettstein (1896). Although it is possible that a valid combination at subspecific rank exists and predates Townsend’s combination, the earliest such relevant use of the epithet pratensis appears to be E. Rostkoviana Hayne subsp. pratensis Ascherson & Graebner, Flora des nordostdeutschen Flachlandes (ausser Ostpreussen): 644 (1899), which is invalid under Article 26.1 and would, in any case, become a later homonym if transferred to E. officinalis (Article 64.4). 3) E. officinalis L. subsp. monticola Silverside, nom. nov., pro Euphrasia montana Jordan, Pugillus plantarum novarum praesertim gallicarum 132 (1852); non E. officinalis L. var. montana (Fries) 348 A. J. SILVERSIDE Fries, Summa vegetabilium scandinaviae 19 (1845). Type: in herb. Jordan, inaccessible and not seen by me; see Yeo (1978). E. rostkoviana L. subsp. montana (Jordan) Wettst. in Denkschr. Akad. Wiss., Wien 70: 319 (1901). E. rostkoviana L. subvar. montana (Jordan) Hartl, J/lustrierte Flora von Mitteleuropa, 2nd ed. 6(1,5): 349 (1972). Non E. montana Phillipi, Plantas nuevas Chilenas 116 (1896). Distribution: scattered throughout much of central and northern Europe, including Britain, mainly in upland areas and typically in damp meadows and pastures; undoubtedly declining through drainage and changes in land-use. Article 64.4 of the /.C.B.N. prohibits the use of the same epithet for different infraspecific taxa, not based on the same type, within the same species, even if they are of different rank. The use of the name E. officinalis in a broad sense has resulted in a host of infraspecific taxon names and, in this case, the existence of E. officinalis var. montana prevents any such combination based on E. montana Jordan. Fries’ var. montana is validated by indirect reference to his earlier description (Fries 1828), under E. officinalis B) E. montana, even if this latter construction is not itself taken to be validly published, for reasons discussed above in relation to his E. officinalis A) E. pratensis. A new epithet has been required and I have chosen an epithet that in both form and meaning is close to the original and should minimise the inconvenience of the change of name. In his account of E. rostkoviana, Wettstein (1896) mentioned E. uliginosa Ducommun. Although Wettstein recognised E. montana separately, a variant of E. rostkoviana from a damp habitat has seemed to need further investigation as a possible synonym and source of epithet. However, although I have seen neither the original description nor any original material of E. uliginosa, it is clear from the accounts of the genus in Reuter (1861) and Ducommun (1869) that E. uliginosa was regarded as a small-flowered, late-flowering taxon, which both authors justifiably maintained separately from E. montana. The illustration of this taxon in Hayek & Hegi (1913: fig. 54) is referable to subsp. rostkoviana and in the second edition of this work (Hartl 1972: fig. 175), the caption is duly altered to “‘subvar. Rostkoviana’’. 4) E. officinalis L. subsp. anglica (Pugsl.) Silverside, comb. et stat. nov. LectotyPe: Box Hill, Surrey, 22nd September 1920, Pugsley 440 (BM) (Pugsley 1930; Yeo 1978). Euphrasia anglica Pugsley in J. Bot., Lond. 67: 225 (1929). Distribution: Britain, Ireland and perhaps also in the neighbourhood of Rouen, France (Yeo 1978), usually in damp, heathy, grazed grassland. Subsp. anglica is at its most distinct in south-western England, where it occurs in the apparent absence of subsp. rostkoviana. Essentially a plant adapted to withstand grazing, it differs from subsp. rostkoviana in its shorter lower-internodes and more basal branching, its smaller corollas and in its frequently more rounded and rather darker leaves. As emphasised by Pugsley (1930), the rather larger floral leaves, decreasing less in size upwards, are characteristic of subsp. anglica, and contribute to the distinctive appearance of many populations. However, when one considers plants from other parts of western Britain and from Ireland, these characters become less reliable. Much Welsh material, in particular, is intermediate in nature and can be named only with difficulty (and then perhaps arbitrarily). Herbarium material of more upright, narrower-leaved plants has sometimes been named as E. anglica var. gracilescens Pugsl., though such specimens do not match the type, from Myrtleberry Cleave, N. Devon, W. C. Barton 277 (BM). While subsp. anglica is recognisable northwards to southern Scotland, with Mrs O. M. Stewart having found small, but quite characteristic material at a cluster of sites in Galloway, it is clear that in much of its range, subsp. anglica is not fully differentiated from subsp. rostkoviana. While it is likely that, as elsewhere in the genus, difficulties arise principally through hybridisation following breakdown of habitat barriers through man’s activities, the two taxa are clearly so closely allied that it is best to consider_E. anglica as a localised derivative of subsp. rostkoviana. Subspecific status of E. anglica appears most appropriate. STATUS OF RELATED TAXA Species concept in Euphrasia is currently a matter for some debate. Hartl’s (1972) account of the genus in central Europe treats at lower rank some taxa recognised as species by Yeo (1972). EUPHRASIA OFFICINALIS AND ITS NOMENCLATURE 349 Karlsson (1976) proposed rather more sweeping revisions. As pointed out by Sell & Yeo (1970), while there is a biological argument for accepting a greatly reduced number of species, this would lead to a cumbersome infraspecific nomenclature. While I agree broadly with Karlsson’s view that the number of taxa given specific status could be greatly reduced, it is also clear that any reduction should be based on careful consideration of taxa over the whole of their ranges and should aim to clarify relationships rather than be based on superficial similarities. Against this background of debate, I have given some consideration to the status of two other glandular species, the diploid, E. vigursti Davey, and the presumed diploid, E. campestris Jordan. E. vigursii is endemic to south-western England, where it is a strikingly attractive plant characteristic of Agrostis curtisii Kerguélen heathlands. There is good reason to believe that it has originated by hybridisation between E. officinalis subsp. anglica and the tetraploid species E. micrantha Reichenb. (Yeo 1956). Yeo (1972) treated E. campestris as a subspecies of E. rostkoviana, occurring in dry grassland from Belgium to Italy. In his account, Yeo (1978) accepted a somewhat wider distribution and regarded E. campestris (still as a subspecies) as being of polytopic origin, derived in some cases directly from subsp. rostkoviana and in other cases through introgression from tetraploid species, including E. stricta. Accordingly, he regarded it as one end of a range of variation, defined rather arbitrarily by small leaves and late-flowering habit. While I readily admit that Dr Yeo has examined considerably more material than I have done, I do not entirely agree with his views. E. campestris has occasionally been reported in Britain, such records having been treated by Pugsley (1930) as referring to hybrids between E. anglica or E. rostkoviana and tetraploid species such as E. nemorosa, a conclusion with which I agree. I have recently compared a distinctive collection by T. G. Evans from a limestone site in the Wye Valley (herb. T. G. Evans) with a range of E. campestris material and, while I concluded that the Wye Valley collection, along with other British candidates for “‘campestris’’, should be regarded as E. officinalis X nemorosa (the latter probably as var. calcarea Pugsl.), I feel the same could be said for a number of continental specimens. There is a nucleus of material characterised by commencing to flower at a high node, usually at node twelve or above, rather short upper internodes, rather numerous, stiff, straight branches emerging high on the stem and held at an angle of around 30° from vertical, distinctly small floral leaves (range not noted but probably rarely exceeding 7 mm) with noticeably acute to aristate toothing and flowers arranged in strict pairs. While I have not seen type material, I take this nucleus to represent the true FE. campestris, a distinctive taxon that, as suggested by Yeo (1972) in his earlier account, has probably arisen by hybridisation with E. stricta. The two taxa, E. vigursii and E. campestris, are, therefore, comparable in origin and should be similarly treated. While I would not argue with their being treated as subspecies of E. officinalis, they are inherently different in nature from the other taxa here so treated, and, for the present, I prefer to retain them at specific rank. The remaining British species of series Euphrasia is E. rivularis Pugsl., endemic to mountain flushes in Wales and the Lake District. As I have suggested elsewhere (Silverside 1990), this may also be of hybrid origin, perhaps originally derived from E. officinalis subsp. rostkoviana crossing with E. micrantha, but with further adaptation to its restricted habitat. While its distinctiveness may be obscured locally by hybridisation with E. officinalis subsp. rostkoviana, 1 consider it a good species in the current context of Euphrasia taxonomy. INFRAGENERIC CLASSIFICATION The genus Euphrasia is typified by E. officinalis. Barker (1982) gives useful discussion. Article 22.1 of the /.C.B.N. now requires that any subdivision of a genus containing the type species of that genus takes the same name as the genus. This means that all generic subdivisions containing E. officinalis must take the epithet Euphrasia, without author citation. The following changes are required from the treatments by Pugsley (1930) and Yeo (1978). Details of typification are taken from Yeo (1978). Section Euphrasia Pugsley: sect. Semicalcaratae Bentham (but see Sell & Yeo (1970) regarding author citation). Yeo: sect. Euphrasia. 350 A. J. SILVERSIDE Subsection Euphrasia Pugsley: subsect. Ciliatae Jorgensen (Lectotype: E. scottica Wettst.) Yeo: subsect. Ciliatae Jorgensen. Series Euphrasia Pugsley: series Hirtellae Pugsley (Type: E. hirtella Jordan ex Reuter) Yeo: series Grandiflorae Wettst. (Lectotype: E. rostkoviana Hayne) ACKNOWLEDGMENTS I am most grateful to the staff of the libraries and herbaria, particularly E, BM and LINN, who have provided assistance and access to their collections, to Arthur Chater for some nomenclatural advice, and also to those collectors who have sent me material and greatly enhanced my experience of the E. officinalis group. REFERENCES Barker, W. R. (1982). Taxonomic studies in Euphrasia L. A revised infrageneric classification, and a revision of the genus in Australia. J. Adelaide Bot. Gard. 5: 1-304. Bunt, W. (1971). The compleat naturalist. A life of Linnaeus. London. BucKNALL, C. (1917). British Euphrasie. J. Bot., Lond. 55 (Suppl.): 1-29. CAJANDER, A. K. (1901). Euphrasia onegensis n. micrsp. Meddel. Soc. pro Fauna et Flora Fennica 1900-1901: 101-102. DucommMun, J. C. (1869). Taschenbuch fiir den schweizerischen Botaniker. Solothurn. Fries, E. M. (1828). Novitiae florae Suecicae, edit. altera. Lund. GREUTER, W.., et al. (1988). International code of botanical nomenclature adopted by the Fourteenth International Botanical Congress, Berlin, July-August 1987. KOnigstein. HA. er, A. (1745). Flora Ienensis, 3rd ed. Jena. Hart_, D. (1972). Euphrasia, in HartL, D. & WaGENITZ, G., eds. G. Hegi illustrierte Flora von Mitteleuropa, 2nd ed. 6(1,5): 335-373. Munich. Hayek, A. von, & Hea, G. (1913). Euphrasia, in Hea, G., ed. /llustrierte Flora von Mittel-Europa, 6(1): 83— 100. Munich. JaLas, J. (1977). New nomenclatural combinations in Euphrasia, Rumex, Silene and Stachys. Ann. Bot. Fennici. 14: 191-192. JALAS, J. & KUKKONEN, I. (1973). Typification of the taxa of Euphrasia (Scrophulariaceae) described by Finnish botanists. Ann. Bot. Fennici. 10: 27-42. Karisson, T. (1976). Euphrasia in Sweden: hybridization, parallelism and species concept. Bot. Notiser. 129: 49-60. Karisson, T. (1982). Euphrasia rostkoviana i Sverige. Vaxtekologiska Studier. 15. Uppsala. Karisson, T. (1984). Early-flowering taxa of Euphrasia (Scrophulariaceae) on Gotland, Sweden. Nord. J. Bot. 4: 303-326. PucsLey, H. W. (1930). A revision of the British Euphrasie. J. Linn. Soc. Lond. (Bot.) 48: 467-544. Reuter, G. F. (1861). Catalogue des plantes vasculaires qui croissent naturellement aux environs de Geneve. Geneva. SAVAGE, S. (1945). A catalogue of the Linnaean Herbarium. London. SELL, P. D. & Yeo. P. F. (1970). A revision of the North American species of Euphrasia L. (Scrophulariaceae). Bot. J. Linn. Soc. 63: 189-234. SILVERSIDE, A. J. (1990). A guide to eyebrights (Euphrasia). Il The E. rostkoviana group. Wild Flower Mag. 418: 31-34. STAFLEU, F. A., et al. (1972). International code of botanical nomenclature adopted by the Eleventh International Botanical Congress, Seattle, August 1969. Utrecht. STAFLEU, F. A., et al. (1978). International code of botanical nomenclature adopted by the Twelfth International Botanical Congress, Leningrad, July 1975, Utrecht. TOWNSEND, F. (1897). Monograph of the British species of Euphrasia. J. Bot., Lond. 35: 321-336, 395-406, 417- 426, 465-477. WETTSTEIN, R. VON (1896). Monographie der Gattung Euphrasia. Leipzig. Yeo, P. F. (1956). Hybridisation between diploid and tetraploid species of Euphrasia. Watsonia 3: 253-269. Yeo, P. F. (1971). Revisional notes in Euphrasia. Bot. J. Linn. Soc. 64: 353-361. Yeo, P. F. (1972). Euphrasia L., in Tutin, T. G. et al., eds. Flora Europaea 3: 257-266. Cambridge. Yeo, P. F. (1978). A taxonomic revision of Euphrasia in Europe. Bot. J. Linn. Soc. 77: 223-334. (Accepted February 1991) Watsonia, 18, 351-358 (1991) 351 Polygonum lichiangense W. Smith: rejected as a naturalized British species Ay P.oCONOLLY 25 Brocks Hill Drive, Oadby, Leicester, LE2 5RE ABSTRACT Despite published records suggesting that Polygonum lichiangense W. Smith (Polygonaceae) has occurred as a naturalized alien in Britain, it is now realized that these are erroneous and arise from confusion with P. polystachyum Meissner var. pubescens Meissner (in Wales) and P. campanulatum Hook.f. (in Cornwall). The bases for these misconceptions are explored, and the characters distinguishing P. lichiangense from the two other taxa described. INTRODUCTION It has been alleged, mistakenly, on at least two occasions, that Polygonum lichiangense W. Smith has occurred as a naturalized garden escape in Britain (Lousley & Kent 1981; Murphy 1988). It would seem useful therefore to outline the reasons for these mistakes and to indicate some of the salient features which distinguish this species from those easily confused with it. Polygonum lichiangense was described in 1914 from material collected in August 1910 by George Forrest (6296, E) in Western China (Lichiang Range, Yunnan) on his second expedition (Smith 1914; Cox 1945) (Fig. 1). Although once cultivated in the Royal Botanic Garden, Edinburgh, but now lost (D. G. Long pers. comm., 1989), it is still grown by D. McClintock at Platt in Kent, froma root obtained many years ago from the Royal Botanic Garden, Edinburgh, bearing the Forrest number 6070 (raised from seed collected in July 1910 from a locality somewhat to the south of F 6296). According to Kent (Lousley & Kent 1981, p. 81) it has been “‘cultivated in British gardens for about 60 years’’. It was in cultivation at the Royal Botanic Gardens, Kew, in 1952 (K, C46). A second, inadvertant, introduction happened when a “plant came up in soil in which plants of Primula sonchifolia were imported from Burma in 1933”’. Grown by Tom Hay, it gained a Royal Horticultural Society Award of Merit in 1933 (Chittenden 1934, p. 306; Cowan 1952, p. 213). The alleged occurrence of Polygonum lichiangense in Wales and in Cornwall, stems from confusion with two other species of Polygonum widely naturalized in the British Isles: firstly with P. polystachyum Meissner to which Smith (1914) considered his P. lichiangense was closely related, and secondly with P. campanulatum Hook.f. to which species Steward (1930) transferred the Chinese plant as a variety — var. lichiangense (W. Smith) Steward. This latter (misleading) assignment has been followed in, for example, the Supplements to the Dictionary of Gardening (Stearn 1956, p. 299; 1969, p. 467) and in the European Garden Flora (Akeroyd 1989, p. 128). ALLEGED NATURALIZATION OF POLYGONUM LICHIANGENSE The first alleged naturalization of P. lichiangense was in the 1970s from Pembs. (v.c. 45) (cf. Lousley & Kent 1981), but Ellis (1983) indicated that this record, as well as one from Merioneth (v.c. 48), was erroneous. They arose from confusion with the ‘hairy’ variant of P. polystachyum, var. pubescens Meissner, to which these records refer (Conolly 1977). The second alleged occurrence was an E. Cornwall (v.c. 2) plant reported (though not determined) by Murphy (1988, p. 24) as ““Polygonum campanulatum var. lichiangense’’, a taxon “thought to be not previously recorded as a garden escape’’. It was growing beside a stream near St Ann’s Chapel (GR 20/40.70). This stand has now been re-examined (Oct. 1989), and, although pale S52 A. P. CONOLLY f Bteesececiinon gomnsnnnononnsreosemnsanioninnne Ficure 1. The lectotype of Polygonum lichiangense W. Smith, Forrest 6296 (E), from eastern flank of Lichiang Range, Yunnan, China. The specimen shows the long ochreae (central stem) and the woody base. Eee a eaten STATUS OF POLYGONUM LICHIANGENSE 353 in colour, agrees with the usual variant naturalized in this country, P. campanulatum var. fulvidum Hook.f., and is clearly distinct from genuine P. lichiangense as represented by the lectotype Forrest 6296 (E) and by specimens of Forrest 6070 (E) and by the clone in McClintock’s garden at Platt, Kent (LTR). The differences, however, are not clearly explained in the more readily obtained literature. REASONS FOR MISIDENTIFICATION When one looks at the original description of P. lichiangense (Smith 1914) it is easy to see how the first of these misunderstandings arose. Smith emphasised the setose stipules (ochreae), as well as the dense grey tomentose undersurface of the leaves, as distinguishing the Lichiang plant from the closely related P. polystachyum; but he failed to point out, or more likely was ignorant of, the pubescent variant P. polystachyum var. pubescens in which the ochreae are indeed conspicuously setose, bearing buff hairs, unlike the glabrous ones in the more usual (in Britain) morph. In addition to pubescent stems and petioles, the leaves of this hairy variant are clothed in a velvety pubescence, especially on the undersurface; and, although this last character can not strictly be termed a ‘tomentum’, it is apparent that a lack of knowledge of this var. pubescens could readily lead to its being misidentified as P. lichiangense. This could happen using the key in Docks and Knotweeds of the British Isles (Lousley & Kent 1981, p. 20). The basis for the confusion with P. campanulatum is more complex, and follows from a sequence of selective quoting of characters without reference to full diagnoses. It originates primarily with Steward (1930) who seemed to consider that those same features highlighted by Smith (setose stipules and grey tomentum) as the main distinction of P. lichiangense from the allied P. polystachyum, merited instead, placing it ‘“‘very close to P. campanulatum var. fulvidum’’. He thus distinguished the Lichiang plant from this latter variety by only three characters: the grey (as opposed to fulvous) tomentum on the under-face of the leaf-blades, the ochreae being ‘‘coarsely brown-strigose”’, and the “pubescence of the upper surfaces of the leaf-blades set in basal glands”’ (a character which in fact also applies equally to P. polystachyum var. pubescens). Steward thus implied the quite false assumption that, in all other characters (including those of the perianth), var. lichiangense agreed with P. campanulatum. Subsequently, although a full and accurate description for P. lichiangense as a species was published in the Dictionary of Gardening (Synge 1956, p. 1628), in the later Supplements (Stearn 1956, p.299; 1969, p. 467) Steward’s reduction of P. lichiangense to a variety of P. campanulatum was taken up again. This is the case too in the recent European Garden Flora (Akeroyd 1989). Both authors select merely the colour of the flowers (white), and of the indumentum (greyish or with white hairs), as the only two features to distinguish the var. lichiangense from typical P. campanulatum. The possible consequent assumption, that in all other respects this variety is identical with P. campanulatum, can explain the mis-naming of the Cornish plant. For here, in deep shade, most flowers (Oct. 1989) were indeed white, and only a few showed the usual pink hue. Here too, the tomentum under the leaves had retained the grey-white colour typical of younger leaves and only the oldest showed the mature ‘fulvous’ buff colour. This, a change with age, is considered a normal sequence (R. J. Murphy pers. comm., 1989). Indeed the distinction of var. fulvidum from the type variety has now been abandoned, and Hara et al. (1982, p. 172) have reduced var. fulvidum to a synonym of P. campanulatum var. campanulatum. DISTINGUISHING CHARACTERS OF P. LICHIANGENSE Examination of the lectotype of P. lichiangense (Forrest 6296, E), of Forrest 6070 (E), and herbarium (LTR) and fresh material from Platt, Kent (Forrest 6070), as well as reference to Smith’s full description (1914) reveal a number of features which distinguish this species both from P. polystachyum var. pubescens and from P. campanulatum: in particular the form of the perianth, the shape, size and colour of the ochreae, and the detailed character of the indumentum of the leaf- blades. These are discussed below and uphold specific distinction for P. lichiangense — an opinion with which Hong (pers. comm., 1990) concurs. 354 A}: P: CONOLLY PERIANTH In P. lichiangense the cup-shaped perianth closely resembles that of P. polystachyum with two narrow outer, and three broad inner members all “cleft nearly to the base’ (Smith 1914 for P. lichiangense;, Hooker 1886 for P. polystachyum). In P. campanulatum, in contrast, the perianth is campanulate, i.e. bell-shaped, with the members only free in their upper part; the lower portion is tubular. Steward (1930) made no mention of this character in relation to his var. lichiangense; nor indeed did later authors. Smith’s (1914) diagnosis makes it clear that the perianth members are like those of P. polystachyum. I can see little difference in the form of the perianth of P. lichiangense (Forrest 6296, 6070) and that of P. polystachyum, apart from a slight difference in size. The perianth members in P. lichiangense Forrest 6296 (type) are larger (c.4 to 4:5 mm or even 5 mm), those of Forrest 6070 (E) somewhat shorter (c.3 mm to 3-5 mm, a few to 4 mm); in P. polystachyum they range 3-0 to 3-5 mm, occasionally to 4 mm, but often less than 3 mm. This size difference is emphasised by Hong (pers. comm., 1989). The very different perianth of P. campanulatum measures 3-5-4 mm. (All measurements were made on dry herbarium material.) OCHREAE (STIPULES) Smith’s emphasis on the setose stipules as distinguishing his P. lichiangense from P. polystachyum only tallies with the more widespread (in Britain) essentially glabrous variant of the latter, in which the ochreae are without hairs; but in the var. pubescens they bear buff hairs precisely as in P. lichiangense. In both these species the ochreae are membranous, very long, narrowly lanceolate, with long attenuate tips and are brown/buff in colour. In P. polystachyum they may reach 4 or even 5 cm, in P. lichiangense often no more than 3 cm; they are mostly long-persistent in both species. This is in contrast to P. campanulatum where the much shorter distal portion of the ochreae is shed early, leaving a loose, broad sheath of c.1 cm in length with a more or less truncate margin; this sheath is white (or colourless) when fresh and beset with off-white hairs. Further shedding reduces the ochreae to a mere 1-2 mm collar and a conspicuous ring of hairs (but cf. Akeroyd (1989) who states “ochreae c.1 mm’’). In P. polystachyum the distal part of the ochreae are eventually also shed, leaving an unevenly truncate sheath, which, even if pale when young, soon becomes buff. LEAF-BLADE INDUMENTUM Steward’s (1930) reference to ‘tomentum’ on the leaf-blade lower (under) surface in P. campanula- tum and in var. lichiangense, fails to distinguish the marked difference in the detailed character of the indumentum in these two taxa. In P. campanulatum the so-called ‘tomentum’ is merely a thin powdery layer — a weft of minute tangled threads too fine to distinguish individually except under high magnification (Fig. 2C). In P. lichiangense (Fig. 2A), in contrast, the indumentum forms a thick, woolly pile of crisped, coarse hairs readily discernible individually under a low magnification, and closely resembling that found in P. weyrichii. This tomentum is less dense in some specimens of Forrest 6070. The pubescent form of P. polystachyum differs again: there is no true tomentum, but a pubescence, usually dense, of individual, somewhat crooked, bent hairs oriented acroscopically, giving a velvety feel (Fig. 2E). The hairs on the upper leaf-blade surface of P. lichiangense (Fig. 2B) have bulbous often coloured (glandular) bases set in pedestal-like raised multicellular supports. This feature is close to that found in P. polystachyum var. pubescens (Fig. 2F). Steward (1930) emphasised the “pubescence of the upper surfaces of the leaf-blades set in basal glands” as distinguishing his var. lichiangense from the type P. campanulatum. And indeed, the equivalent hairs in P. campanulatum do differ (Fig. 2D) for as well as being much coarser and stiffer, they are not glandular, and lack the pedestal-like base: only some having slightly raised green supporting cells. As in P. polystachyum var. pubescens these hairs are restricted to parallel bands between the prominent lateral veins. The angle of divergence of these with the mid-rib, some 45°, is in marked contrast with the broad angle (c.90°) seen in P. polystachyum. No such conspicuously parallel lateral veins are evident in P. lichiangense, the hairs being un-zoned. Under very high magnification (SEM) the surface ornamentation on these hairs differs in detail for the three taxa concerned (Figs. 3A, B, and C). . In P. campanulatum coarse bristle-like hairs resembling those of the upper leaf-blade surface cover the prominent veins on the under surface (Fig. 3D). STATUS OF POLYGONUM LICHIANGENSE 200 oe, . if at > Pi ? . 7 Ne ‘ i" r — Me ae YF Bes 1 Wy os a VELA AL A UF Sal } Gs — — r . = ie is Ze 2 4 Py as Sey oe EM cS ont Laon a8 a? ge AP FiGureE 2. SEM photograph of leaf-blade surfaces to show indumentum in Polygonum lichiangense (A,B); P. campanulatum (C,D); and P. polystachyum var. pubescens (E,F). A,C,E abaxial surface; B,D,F adaxial surface. Same magnifications for abaxial surfaces; ditto for adaxial surfaces. Note: A, P. lichiangense crisped hairs obscuring abaxial epidermis; C, P. campanulatum fine tomentum of tangled threads; E, P. polystachyum var. pubescens isolated hairs not obscuring epidermis. B,F, P. lichiangense and P. polystachyum var. pubescens hairs with swollen bases on raised supporting pedestal. D, P. campanulatum bristle-like hairs with parallel orientation, on raised supporting cells. Source of material for SEM: P. lichiangense: (Forrest 6070) garden, Platt, Kent. P. campanulatum: St Ann’s Chapel, E. Cornwall (GR 20/40.70). P. polystachyum var. pubescens: Rydal Water, Cumbria. 356 A. P. CONOLLY FicurE 3. SEM photographs to show detail of ornamentation on hairs of adaxial leaf-blade surface: A, P. lichiangense; B, P. campanulatum; C, P. polystachyum var. pubescens. Note: ornamentation elongated, wavy in A; short, mound-like in B; and elongated spirally in C. D, abaxial surface of leaf-blade of P. campanulatum to show prominent hairs of vein. A, B, C, same magnification. Material: as in Fig. 2. FURTHER FLORAL FEATURES Another major difference between these taxa is that both P. polystachyum and P. campanulatum have dimorphic flowers: separate plants are either long-styled or short-styled; P. lichiangense, on the other hand, is apparently homostylous (but material from only three collections has been seen). The styles in P. polystachyum measure up to 1-0 (1-2) mm and 0-4 (0-5) mm respectively, and the stigmas are small and capitate or peltate. P. campanulatum is much the same. In P. lichiangense the three stigmas are sessile (no measurable length of style), and are obconical (funnel-shaped), a shape approaching that seen in P. weyrichii and some (but not all) other members of Section Aconogonon. This agrees with Kral’s (1985) illustration of the obconical sessile stigmas in P. alpinum, which he contrasted with the long styles of P. polystachyum — a contrast which would have been more dramatic had he illustrated a genuine long-styled flower and not the short-styled morph. Hong (pers. comm., 1989) has confirmed this feature in P. lichiangense. P. lichiangense is also distinguished by the anther colour which is pale creamy-fawn. The anthers in the other two species become very dark, almost black, but start violet-purple (P. campanulatum), or reddish-purple (P. polystachyum). The pollen appears white in both these species, in P. lichiangense almost transparent with a bluish tinge. Smith (1914) did not observe mature fruit on his material of P. lichiangense. Material examined STATUS OF POLYGONUM LICHIANGENSE 857 (Platt, Kent, LTR) suggests a trigonous (or somewhat flattened), light-brown, shiny nut (achene) which does not protrude beyond the faded perianth. This fruit is not dissimilar to that formed (and then only extremely rarely from my observations) in P. polystachyum; although Akeroyd (1989) indicated ‘‘Fruit much longer than perianth”. P. campanulatum also has an included nut, not ‘exserted’ as Steward (1930) wrote, nor truly winged when fully mature. Nor is the fruiting perianth consistently fleshy as Hara (1966, p. 632) implies; any thickening of the perianth seems to be a rare late-season development, and quite unlike P. molle whose spherical black fruit formed by the thick fleshy perianth around the enclosed nut, is shed as a unit. Such a ‘baccate’ fruit (Hooker 1886, p. 50) is a feature peculiar to P. molle s.1. There are, of course, several other features distinguishing P. lichiangense which can be seen on perusal of herbarium material. Thus the strongly developed woody root-stock, without apparent sign of rhizomatous off-sets contrasts with P. polystachyum where these are abundant, and with P. campanulatum which has extensive prostrate stems. The overall shape and form of the leaves in P. lichiangense differs too, and is only approached by narrower-leaved examples of P. campanula- tum, or by the small leaves on the inflorescence of P. polystachyum. The veining is quite unlike the prominent parallel form of the other two species. The form of the inflorescence is closer to that of P. polystachyum. CONCLUSION On several counts, P. lichiangense can be maintained as a species, clearly distinct from P. campanulatum, but less so from P. polystachyum var. pubescens, an opinion confirmed by Hong (pers. comm., 1989) and upheld by his investigation of pollen characters (Hong & Hedberg 1990). As yet there is no evidence that genuine P. lichiangense has escaped from cultivation in Britain. The assignment to a variety of P. campanulatum should be abandoned. TAXONOMY AND NOMENCLATURE The three species here discussed as species of Polygonum s.|. are commonly placed in the sect. Aconogonon Meissner. Hara (1966) followed Reichenbach (1837) who raised the section to a genus, Aconogonum. All three have valid names in that genus were this taxonomic split to be adopted (Hara 1966; Kral 1969; Sojak 1974). Furthermore all three have also been transferred to another segregate genus, Persicaria (Ronse Decraene & Akeroyd 1988; Gross 1913; Greuter & Raus (1989); and invalidly by Trehane (1989). The elevation of P. polystachyum to a separate genus (Kral 1985) is not accepted here. The following specimens of P. lichiangense have been seen: 1) Yunnan, West China. Lat. 27°40’N. Aug. 1910. Forrest 6296 (E). Type fide W. W. Smith (Lectotype selected by S.-P. Hong). ‘‘Flowers creamy-white’’. (Isotype BM). 2) Yunnan, West China. Lat. 27°30'N. July 1910. Forrest 6070 (E). “Polygonum lichiangense W. W. Sm. var. Flowers creamy yellow”’. 3) Forrest 6070 ex Hort. R. B. G. Edin. 1911 (E). ‘Polygonum aff. P. polystachyum Wall. var. vel Spi nove. 4) & 5). Forrest 6070 ex Hort. R. B. G. Edin. via Garden at Bracken Hill, Kent. 29 Oct 1975, D. McClintock (LTR). ditto, 27 Sept. 1975, D. McClintock (LTR). 6) Royal Botanic Gardens, Kew. Cultivated. Aug. 1952. C46 (K). 7) West China. Oct. 1933. McLaren 231 (K). ACKNOWLEDGMENTS I am indebted to D. G. Long of the Royal Botanic Garden, Edinburgh, for the loan of the type (and other sheets) of Forrest’s original gatherings of P. lichiangense, to D. McClintock who first drew my attention to this plant, provided herbarium material (now LTR) and fresh leaves for the S.E.M., to G. McTurk, J. Bailey, I. Riddell and S. Ogden for technical help with S.E.M. and photography. I 358 A. P. CONOLLY record my thanks to Miss R. J. Murphy for taking me to the Cornish plant, to the National Museum of Wales (per R. G. Ellis) for the loan of the negative of the type specimen, and to S.-P. Hong (Uppsala) for his comments from his unpublished thesis; also to the Natural History Museum, London, for facilities, and to Prof. C. A. Stace for taxonomic guidance. REFERENCES AKEROYD, J. R. (1989). Polygonum Linnaeus, in WALTERS, S. M. et al., eds. The European garden flora 3: 125— 128. Cambridge. CHITTENDEN, F. J., ed. (1934). Plants to which awards have been made 193344. J. R. Hort. Soc. 59: 300-307. Cono ty, A. P. (1977). The distribution and history in the British Isles of some alien species of Polygonum and Reynoutria. Watsonia 11: 291-311. Cowan, J. M., ed. (1952). The journeys and plant introductions of George Forrest. Oxford. Cox, E. H. M. (1945). Plant hunting in China. London. Eris, R. G. (1983). Flowering plants of Wales. Cardiff. GREUTER,. W. & Raus, TH., eds. (1989). Med-checklist notulae, 15. Willdenowia 19: 41. Gross, H. (1913). Remarques sur les Polygonées de Asie Orientale. Bull. Acad. Int. Géogr. Bot. 23: 7-32. Hara, H. (1966). The flora of eastern Himalaya. Tokyo. Hara, H., Cuater, A. O. & Witiiams, L. H. J. (1982). An enumeration of the flowering plants of Nepal 3: 172. London. Hong, S.-P. & HepBeErRG, O. (1990). Parallel evolution of aperture numbers and arrangement in the genera Koenigia, Persicaria and Aconogonon (Polygonaceae). Grana 29: 177-184. Hooker, J. D. (1886). Flora of British India 5: 50. Ashford. KrAL, M. (1969). Aconogonon polystachyum comb. n. Preslia (Praha) 41: 258-260. KRAL, M. (1985). Rubrivena, a new genus of Polygonaceae. Preslia (Praha) 57: 63-67. LousLey, J. E. & Kent, D. H. (1981). Docks and Knotweeds of the British Isles. B.S.B.I1. Handbook No. 3. London. Murpny, R. J. (1988). Plant notes and records, mainly 1987. Botanical Cornwall Newsletter 2: 11-29. REICHENBACH, H. G. L. (1837). Handbuch des natiirlichen Pflanzensystems, 236. Dresden & Leipzig. Ronse DecrAENE, L-P. & AkeEROYD, J. R. (1988). Generic limits in Polygonum and related genera (Polygonaceae) on the basis of floral characters. Bot. J. Linn. Soc. 98: 321-371. SmiTH, W. W. (1914). Diagnoses specierum novarum in herbario Hort. Regii Botanici Edinburgensis cognitum LI-CII (Species Chinensis). Notes R. Bot. Gard. Edin. 8: 173-212. SosAK, J. (1974). Bemerkung zur Gattung Truellum Houtt. (Polygonaceae). Preslia (Praha) 46: 139-156. STEARN, W. T. (1956). Polygonum in SynGe, P. M., ed. Supplement to the Dictionary of Gardening: 298-299. Oxford. STEARN, W. T. (1969). Polygonum in SyncE, P. M., ed. Supplement to the Dictionary of Gardening, 2: 466-468. Oxford. STEWARD, A. N. (1930). The Polygoneae of Eastern Asia. Contr. Gray Herb. Harv. 88: 1-129. SyNnGE, P. M., ed. (1956). Polygonum, in Dictionary of Gardening, 2nd ed., 3: 1627-1629. Oxford. TREHANE, P. (1989). Index Hortensis, 1. Wimborne. (Accepted February 1991) eae ) Aas a a We Watsonia, 18, 359-367 (1991) 359 Further observations on Stachys germanica L. At. DUNN Flat 2, Sandford Mount, Charlbury, Oxford, OX7 3TL ABSTRACT Observations on the life cycle and ecology of Stachys germanica L. (Labiatae) were made over a period of six years (seven flowering seasons) between June 1984 and October 1990, particular attention being paid to the role of predators. The hybridization of S. germanica in cultivation in private and botanic gardens is noted, and present and proposed conservation measures are discussed. INTRODUCTION Stachys germanica is widely distributed throughout western, central and southern Europe and is at the northern limit of its range in Britain; it extends southwards through the Mediterranean to North Africa, eastwards to southern central Russia, and into the Orient (Moore 1987). The earliest British record, published in Johnson’s revised edition of Gerard’s Herball (Gerard 1633), was by Leonard Buckner, a London apothecary, who found it in 1632 “wilde in Oxfordshire in the field ioyning Witney Parke [This deer park was laid out by the Bishop of Winchester in the 13th century and remained recognisable as a walled enclosure until late in the 17th century. The land is now farmed or built upon.] a mile from the Towne’’. Undisputed 18th, 19th and 20th century records are from N. Hants (v.c. 12), W. Kent (v.c. 16), S. Lincs (v.c. 53), Northants (v.c. 32) and Oxon (v.c. 23), but it is likely to have been more widespread in parts of southern Britain than these records suggest (Marren 1988). Most of the numerous localities recorded by Druce (1886, 1927) were from the oolitic limestone country of western Oxon, to which region the few remaining sites are now confined. S. germanica is an endangered species, protected under the Wildlife and Countryside Act, 1981 (Perring & Farrell 1983). POST 1983 DISTRIBUTION AND POPULATIONS Since 1983 S. germanica has been recorded from five sites. Of these only two produced flowering plants in seven consecutive seasons (1984-1990), while at another site a single plant was seen only in 1987. In 1990, flowering plants were recorded at three sites. All are within the Oxfordshire distribution area shown by Marren (1988), and for convenience the site references he used are adopted here. SITE A (ALSO KNOWN AS SITE X) Fifty-six flowering stems were recorded at this new site, a green lane-bordered by arable fields, in June 1984 (Dunn, 1987a). Long-buried seed germinated after scrub clearance and hedge cutting between 11 February and 4 March 1983. (This date, recorded in the farm diary of the Estate concerned, is the correct one. A verbally given date, November/December 1982, was incorrect.) Although plants have flowered every year since then, largely because open ground conditions have been maintained, the colony has lost much of its vigour since 1985. The majority of plants have produced only one flowering stem instead of up to seven stems. They have also been smaller, on average. It was incorrectly stated (Marren 1988) that ‘‘. . . in 1986 the stems were picked before it [S. germanica] could set seed.” The theft of these seed-bearing stems in late autumn 1986 has probably affected population numbers since then. There was no germination in the dry spring and summer of 1989. The 33 flowering stems recorded in 1990 were poor, and only five very small plants 360 A. J. DUNN TABLE 1. STACHYS GERMANICA (SITE A) COMPARISON BETWEEN NUMBER OF OVER WINTERING ROSETTES AND SUBSEQUENT NUMBER OF FLOWERING STEMS Che: f.s. O.r. f.s. O.r. iS. oe f.s. 1983/84 1984 1984/85 1985 1985/86 1986 1986/87 1987 — 56 20 60 116 56 133 116 O.r. ESS O.F. is. o.r. f.s. or: 1987/88 1988 1988/89! 1989 1988/90 1990 =1990/91 81 62 185 67 41 33 5 o.r. — overwintering rosettes f.s. — flowering stems ‘A mild winter in which 30-40% of overwintering rosettes, some very small, were partially or wholly destroyed by snails and/or slugs. were found to have germinated after rain in September and early October ended a second year of drought. Table 1 shows the fluctuations in numbers of overwintering rosettes and subsequent flowering stems between 1984 and 1990/91. SITE B For several decades S. germanica has appeared erratically at this site, an old bridleway bordered by hedges and bounded on either side by arable or other farm crops. Between the wars this lane was much wider and had no hedge on the southern side. A retired local farm worker recalled driving a horse and wagon along it and seeing a few villagers grazing house cows on the verges, which, until comparatively recently, were cut at regular intervals by council workmen. In 1985, 43 fine flowering stems were recorded. No plants were seen in 1986 and 1987, by which time the site had become densely overgrown with Prunus spinosa, Rosa and Rubus spp., Clematis vitalba, coarse grasses, herbs and mosses. In March 1988 the invading vegetation was cut down and burnt. The ground was then raked hard to clear it of dead plant litter and mosses, and to expose as much bare soil as possible. Buried seed germinated in late spring and by mid-June many seedlings and young plants of S. germanica had developed. That autumn c. 100 rosettes of various sizes were recorded. All survived the following mild winter, producing in the hot summer of 1989 — at least three weeks earlier than normally —c. 100 flowering stems. Further along the lane, where there had been simultaneous clearance work and burning but no raking, another 25 flowering stems were located among dense tall stands of Artemisia vulgaris, Cirsium vulgare, Dipsacus fullonum subsp. sylvestris, sprouts of Crataegus monogyna, Cornus sanguinea, Euonymus europaeus, Prunus spinosa and encroaching layers of Clematis vitalba. By autumn 1989, c. 80 new rosettes were found at the main site, which measures approx. 7 X 1-5 m. In November a long section of the verge to the west was cleared by scrub-cutter. Raking and burning followed at the end of that year, and again in March 1990. In the second hot summer of 1990, c. 130 flowering stems were recorded, all but seven of these being at the main site. By early August all but a few were found to have been completely stripped of their seeds by small mammals (see Predation). Only ten small overwintering plants were found in the late autumn of 1990. SITE C This site has been fully described by Marren (1988). Table 2 shows how widely S. germanica populations have fluctuated between 1984 and 1990. Of the 103 flowering stems borne by the 55 plants in 1989, 20 were found, on 1 August, to have been severed at or a little above ground level, and by mid-October this number had risen to 34. This site is difficult to search for overwintering rosettes on account of standing crops. eer OBSERVATIONS ON STACHYS GERMANICA 361 TABLE 2. CHANGES IN POPULATION SIZE OF S. GERMANICA AT SITE C, 1984-1990 Year No. of plants No. of flowering stems 1984 c. 100 no record 1985 1986 1-40 no record 1987 1988 17 22 (plus c. 18 pairs of fi. later- als) 1989 55 103 (plus 42 pairs of fl. laterals) 1990 1 1 SITE D Although S. germanica has been known for many years at this site, numbers are declining. Only one flowering plant was recorded in 1987, none in 1988, and the single plant found in 1989 was growing in a small patch of bare earth at the bottom of this limestone grassland bank. No plants were seen in 1990. TETRAD SP 4224 In 1987 one flowering stem was found in this tetrad, on the edge of a roadside ditch which had been cleaned out by workmen. No plants have appeared since. THE GARDEN SITE This is not a wild site but is included on account of its interest. As reported elsewhere (Dunn 1987b), S. germanica seeds were introduced accidentally into this west Oxfordshire garden with imported local topsoil more than twenty-five years ago. Although flowering plants were not noticed until the mid-1970s, they have appeared almost every year since then, no doubt because flower beds are continually disturbed by digging and weeding. The 1987 plants were exceptionally fine, bearing many pairs of flowering laterals on tall spikes. Apart from the one hybrid plant found and removed in the late summer of that year (see below), all the plants in this garden colony remain pure. ECOLOGY This section on the ecology of S. germanica is concerned with observations on the plant’s life cycle, pollination, predation, and certain environmental factors. THE LIFE CYCLE S. germanica seeds ripen between July and early autumn, according to the season. Germination, which takes place almost exclusively around the parent plants, is erratic, commencing in late spring and, if conditions are favourable, continuing into early autumn. This accounts for the great variation in size of over-wintering rosettes, which at Site A in September 1987, for instance, measured between 2 X 1-5 cm and 24 X 19 cm in diameter. For three successive autumns at this site all new rosettes were marked and labelled, an experiment which showed that some overwintered twice before flowering. Out of an original group of 81, 24 did so in 1987/88 and again in 1988/89, the largest plant measuring 26 x 22 cm by the second autumn. A few were suspected of overwintering a third time, but as several labels were displaced by rabbits this could not be proved. Healthy, well-grown rosettes, which by late autumn have formed between four and ten medium-to-large leaves, usually overwinter successfully. Under-developed plants, or those measuring less than c. 2 cm in diameter tend to die, survive but fail to flower or produce only very small flowering stems in the following summer. The silky-hairy appearance of the plants is most attractive in May and early June, when the plants are young. Growth is quite rapid during early summer, and by mid-June to mid-July mature plants can measure as much as 104 cm, or as little as 10 cm (Site A, 1985). Vigorous plants may produce up 362 A. J. DUNN to seven stems apiece, each stem bearing one or more pairs of laterals. Small, late-developing plants may not flower until September or early October. Strictly speaking, S. germanica is a biennial, forming a rosette in the first year and dying after flowering in the second. Occasionally a robust plant will produce several shoots at the base of its stem, or stems, in late summer. This appears to be simply the result of the plant’s vigour, in response to late summer/early autumn rains after drought, or after damage. Strong shoots will develop into fine flowering stems the following year, after which the plant dies. During the period of study, no such plant was seen to repeat this behaviour. Dead plants will remain upright, or inclined, for many months. The seeds, four to each calyx, are not easily dislodged. Some have been found unshed after several months, a few even after a year. Deferred seed dispersal, erratic germination and delayed flowering are all strategies which extend the life cycle and assist the survival of the species. POLLINATION The following species of bumblebee were observed pollinating S. germanica flowers at Sites A and B: Bombus hortorum (L.), B. lapidarius (L.), B. pascuorum Scopoli, B. pratorum (L.) and B. terrestris (L.). It was noticed that the smallest of these, B. pascuorum, worked the flowers more thoroughly. Conspicuous numbers of the hoverfly Episyrphus balteatus (Degeer) visited Site A flowers in August 1987; other hoverflies seen here in 1989 were Platycheirus albimanus (F.) and Syritta pipiens (L.). The pollen beetle Meligethes aeneus (F.), which eats as well as distributes pollen, has occasionally been seen crawling in and out of the flowers. PREDATION Minimal to serious damage and destruction have been observed at all stages of growth in S. germanica, the principal predators being small mammals (mice and voles), snails and/or slugs, and the larvae of a micro-species of Tortrix moth. Rabbits have caused slight damage to plants at one site; and rabbits, hares or deer are suspected of causing more extensive damage at another. Mice and voles The most interesting, but most devastating, destruction occurred at Site B between 15 July and 7 August 1990. On the latter date all but a few of c. 130 stems (including laterals) were found stripped bare. Only the leaves remained. This proved to be the work of small mammals. The evidence lay in the shredded remains of S. germanica lying below and around the plants, two of these mats of plant material measuring 27 X 23 cm and 17 X 15 cm. A quantity of the material collected for closer examination contained, in addition to pieces of stem and small, whole leaves, a great quantity of calyces which had been detached from the whorls individually or in small sections. Each calyx had been split open from the base almost to the apex, or torn in half lengthwise. All were void of seeds, many of which would not have been ripe. Mixed in with these remains were seedless wings of the fruits of Heracleum sphondylium and broken, empty capsules of Silene latifolia and S. vulgaris, and, significantly, the droppings of mice and/or voles. Nearby, similarly shredded heads of Centaurea scabiosa littered the ground. Finally, the microscope revealed the presence of fragments of husks of S. germanica seeds. To establish the identity of the rodents, three Longworth live-traps were set beside the plants on ten occasions, usually on alternate evenings, between 11 and 26 August. Fifteen long-tailed field or wood mice (Apodemus sylvaticus (L.)) and four bank voles (Clethrionomys glareolus Schr.) were captured, and on twelve occasions mice or voles entered the traps, ate some or all of the bait, deposited droppings, and escaped. Thirty-one captives and escapees are accounted for by the fact that on one occasion two long-tailed field mice were found in one trap. All captives were released at the edge of a wood 8 km away. One dead long-tailed field mouse was also found on the path beside the site. It was learnt from discussions with a member of staff of the Department of Zoology, University of Oxford, that unusually high numbers of small mammals were captured in his study area in 1989 and 1990. These increases in population are attributed to two successive mild winters and fine summers (1988/89 and 1989/90) which no doubt reduced mortality rates and increased breeding levels. Other factors would have affected the numbers and behaviour of these small mammals at Site B. The crops (barley and rape) at either side of the bridleway were harvested by mid-July and the fields ploughed : : | OBSERVATIONS ON STACHYS GERMANICA 363 shortly afterwards; hedgerow fruits and berries were unripe; and the only adjacent cover, once the fields were bare, would have been in these hedgerows and verges. Food and moisture were probably scarce. Out of a potential 100,000 or more seeds only a few hundred were saved. Similar destruction, on a minor scale, was first observed at Site A early in October 1985. Here a Longworth live-trap set close to a hole near two small de-headed S. germanica plants captured a long-tailed field mouse. Close by, in June, seeds of Viola hirta L. were found to have been similarly raided, the cream-coloured seed-coats lying in little heaps around the plants. Small mammals tunnelling through the soil have disturbed seedlings and young plants, detaching their roots and raising them up. A group of 30 seedlings at Site B (1989) failed to survive such disturbance; others, pushed down into firm soil, were saved. Other mammals At Site A rabbits have occasionally forced their way into a fenced-off area, disturbing seedlings and young plants and burying rosettes under scuffed-up soil. In May 1990, at Site B, the tops of four or five young plants were found lying on the ground below. Rabbits were almost certainly responsible, and further damage was prevented by placing Prunus spinosa trimmings around the plants and into the runs between the hedge and the site. At Site C, in 1989, 34 flowering stems were found to have been severed from about a dozen plants, some of which had borne up to five stems each. None of the missing stems was traced, except on one occasion, when two, freshly broken off, were found lying beside a plant. Stems of such plants growing among second-year kale had been broken off several cm from ground level, while others in open ground had been severed at the base, as though bitten off. This site is frequented by rabbits, whose numbers have greatly increased on this farm in recent years (c. 500 were killed after harvest in 1990). Hares are seen occasionally, and deer are known to pass through the site. Deer droppings — much less plentiful than those of rabbits — were of fallow or muntjac. Snails and/or slugs At Site A, in the autumn of 1988, 185 rosettes of various sizes, some very small, were individually numbered (with labels), measured and recorded. Only 67 survived to flower the following summer. Of the remaining 118, 75 were lost and 43 either failed to grow beyond the vegetative stage or developed into small, non-flowering plants. It was evident that between 30 and 40 per cent had been partially or wholly eaten away by slugs and/or snails during the mild winter. Live snails collected at the site included Candidula gigaxii (Pfeiffer), Cepaea hortensis (Miller), Monacha cantiana (Montagu), Pomatias elegans (Miller), Trichia striolata (Pfeiffer) and Vitrina pellucida (Miller); empty shells were of Cernuella virgata (da Costa), Cochlodina laminata (Montagu) and Trichia hispida (L.). One unfortunate plant which first overwintered in 1987/88 failed to develop in 1988 and in late August of that year had most of its leaves shorn off by a verge-cutting machine. It subsequently formed four large new leaves, only to have these eaten away by snails or slugs during the following winter (a live specimen of Vitrina pellucida was found on the underside of one of its leaves). By mid- May 1989 only fragments of the leaves and midribs remained, and a few weeks later the plant died. Tortrix moth larvae At Site A, in May 1985, the leaves and developing shoots of a few S. germanica plants were damaged by a small, dingy-brown caterpillar, only one of which was found on each plant. All were destroyed except one, which was removed and allowed to pupate in captivity. The emergent imago was identified as Cnephasia interjectana (Haw.), a microspecies of leaf-rolling Tortrix moth. While the effect on leaves was not serious, damage to the growing shoots was severe, resulting in stunted spikes and the consequent loss of many whorls of flowers and seeds. Damage by this Tortrix moth has been noted in up to seven plants at Site A each year of the study, but has not been observed at other sites. SOME ENVIRONMENTAL INFLUENCES The oolitic limestone soil of these sites varies from a light, almost sandy cornbrash to medium, stony loam or marl and supports a varied and sometimes rich flora. Where this soil combines with an open, sunny habitat S. germanica thrives best; whereas in situations which are overshadowed, by 364 A. J. DUNN hedgerow trees for example, it tends to grow less vigorously and flowers later. It is a species which competes poorly with other plants, soon becoming overwhelmed by grasses, perennial herbs and shrubs, especially where these form a close sward or scrub conditions. Bare ground is essential for the germination of its seeds and the formation of overwintering rosettes. Where the soil is kept open deliberately, or is disturbed in the course of crop cultivation, it maintains its presence more successfully, and more regularly. Conversely, at Site A, for example, in places where single or small groups of plants first appeared in 1984 and it has not been possible to preserve open ground, S. germanica has not reappeared. The composition and number of associated plants varies at different sites. In late May 1988, for instance, during clearance work by hand of invading vegetation in an area at Site A measuring approx. 5 x 2m, no less than 46 other species were found to be growing in close proximity to plants of S. germanica. Among these were Arrhenatherum elatius, Bromus erectus, Cirsium vulgare, Clinopodium vulgare, Centaurea nigra, C. scabiosa, Galium mollugo, G. verum, Mercurialis perennis, Plantago lanceolata, Poa pratensis, Primula veris, Ranunculus repens, Silene latifolia, Stachys sylvatica, Tamus communis, Torilis japonica, Viola hirta and V. riviniana. Also here were sprouting stumps of Acer campestre, Clematis vitalba, Crataegus monogyna, Prunus spinosa, Sorbus torminalis and Viburnum lantana. In addition, the following mosses covered parts of the soil: Barbula unguiculata Hedw., Bryum sp., Eurhynchium praelongum (Hedw.) Hook., Fissidens taxifolius Hedw., Phascum cuspidatum Hedw. and Pseudoscleropodium purum (Hedw.) Fleisch. Extremes of weather can affect S. germanica adversely. Very small plants may be lifted from the soil in severe frosts; leaves, seedlings or small plants may decaye in prolonged wet weather in autumn and winter; and summer drought can cause stress, inhibit growth and prevent germination. A summary of weather at Oxford between 1984 and 1990, extracted from records compiled by the Radcliffe Meteorological Station, School of Geography, University of Oxford, is shown in Table 3. There is also the human factor. In 1987 and 1988, seeded plants at Site A, an S.S.S.I., were torn to pieces and scattered by a hedge- and verge-cutting machine following a breakdown in communi- cations and instructions on the part of the Estate concerned. Flowering stems have occasionally been picked by unknown persons, and, as mentioned previously, seed-bearing stems have been stolen. HYBRIDIZATION In 1987, suspected hybridization between S. germanica and S. byzantina C. Koch (syn. S. lanata Jacq.) occurred at the Garden Site. The hybrid plant, which grew beside a stand of pure S. germanica in a flower border, was different in several respects: the stem and leaves were densely felted rather than silky-hairy; the leaves were of a slightly different shape, and entire instead of crenate; and at the base of the stem were several fresh shoots and leaves. The plant was removed, potted, and taken away. In 1988 it produced three flowering stems and fresh basal shoots and leaves. At the request of Dr S. R. J. Woodell, Department of Plant Sciences, University of Oxford, a few pollen-bearing flowers were collected from this plant and also from S. byzantina and S. germanica. The results of the pollen stain tests (Table 4) confirm that the plant was of hybrid origin. The hybrid plant produced one flowering stem in 1989 and one in 1990. Each year it has formed basal shoots and leaves, indicating a perennial rather than a biennial habit. About 120 seeds were extracted from the original hybrid, 55 of which were sown in a seed tray the following spring. The largest twelve of the 25 young plants which germinated were potted on and by October 1988 were well developed. During the hot summer of 1989, in unavoidable circumstances, ten perished through lack of water. One of the remaining two produced a stunted stem bearing only a few flowers, and the other failed to develop beyond the vegetative stage. By autumn both had recovered, put on fresh growth and overwintered successfully in 1989/90. In the summer of 1990 one bore four short flowering stems, and, by autumn had produced a dense mass of basal leaves; the other bore one tall flowering stem and no basal shoots or leaves. All appeared to carry viable seed. In 1987 and 1988 visits were made to University botanic gardens at Bristol, Cambridge and Oxford to examine S. germanica in cultivation in systematic beds. It was interesting to find that in all cases the stands labelled ‘Stachys germanica’ contained plants unlike the wild species. Some were strikingly different, their appearance suggesting that S. alpina and S. byzantina were involved in the a eS Sale OBSERVATIONS ON STACHYS GERMANICA 365 TABLE 3. WEATHER SUMMARY AT OXFORD: 1984-1990 (by permission of the Radcliffe Meteorological Station, School of Geography, University of Oxford) Bright Mean air Rainfall sunshine Year temp. (°C) (mm) (hours) 1984 10-2 608-5 1574-2 1985 9-2 662-0 1483-2 1986 9-2 704-3 1617-8 1987 9-4 592-6 1372-6 1988 9-9 571-1 1401-6 1989 11-1 598-1 1868-1 1990 11-1 473-0 1875-7 LONG TERM MEAN READINGS: Air temperature (°C) (1815-1980): 9-5 Rainfall (mm) (1767-1980): 642-5 Bright sunshine (days) (1881—1980):approx. 1480-0 Ground frost (days) (109 years): 103-7 Snow lying (days) (64 years): 11-4 Ground frost (days) 113 114 116 102 104 94 96 Snow lying (days) 5 Zi 14 16 Notes 1983/84 winter slightly warmer and wetter than average. Unsettled spring, with frequent ground frosts; driest and sunniest April since 1912. Notably warm, dry, sunny summer, with below aver- age rainfall. Mild wet autumn/ early winter. Jan.—Mar. cold, with frequent snow and hard frosts. Changeable spring; heaviest daily rainfall in May on record. June—Aug. wettest since 1971. Dry, cool autumn. Dec. mild and wet with some severe frosts. Year dominated by westerly airst- reams. Feb. extremely cold. Spring cold and wet; April coldest since 1881. Cooler and wetter summer than average. Dry, cold autumn. Mild early winter. Some unusually cold weather at beginning and end of year. Chan- geable spring. Average summer, with less sunshine. Wettest Oct. since 1960. Beginning and end of year remark- ably mild. Very wet Jan. (100 mm). Unsettled spring with less than average sunshine. Cool, rather wet summer. Last five months bright and very dry. Second consecutive mild winter (1988/89). One of the warmest and sunniest years of the century. (Nationally the warmest year since 1659.) Rainfall well below aver- age. Dec. wettest month. Third successive mild winter (1989/ 90); warmest since 1815, wettest since 1767. Spring warm, sunny and dry. Hot, dry summer (max. temp. 35-1°C on 3 Aug. highest ever recorded). Warm autumn with below average rainfall. 366 A. J. DUNN TABLE 4. NUMBERS OF POLLEN GRAINS IN SAMPLES OF THREE STACHYS TAXA Species Well stained Collapsed or non-stained Stachys germanica 265 22 213 2D) 283 66 224 22 S. byzantina Za 1) 215 23 224 v4 184 41 hybrid 196 230 170 126 164 100 148 166 hybridization which had clearly taken place; both these related species grew with S. germanica in all the beds, together with S. affinis Bunge (China, Japan), S. grandiflora (Willd.) Benth. (Caucasus) and S. officinalis at Cambridge, and S. officinalis at Oxford. Also of interest were specimens of S. germanica seen in three private gardens in Oxon, all plants having been raised from wild seed of uncertain provenance. In one case, where the introduction had taken place about twenty-five years ago, the plants were no longer pure, their appearance suggesting hybridization with S. byzantina, which grew in another part of the garden. Seed collected from these plants by the owner and sent to a seed bank were later destroyed as impure. In the second case, where plants had been established for about nine years, several of many very fine stems exhibited features, including a semi-prostrate habit, which were uncharacteristic of the true species and more typical of S. byzantina. In the third case, where seed had been introduced only four years previously, the plants were pure. In view of the ease with which S. germanica hybridizes it was strongly recommended that in the event of reintroductions into the wild only seed gathered directly from wild stock should be used. Although hybridization of S. germanica and S. palustris L. has been reported from France, Germany and Hungary, and with S. sylvatica L. from Hungary (Stace 1975), there is no record of such hybridization in Britain. However, a watch is being kept at one site where S. germanica and S. sylvatica grow in close proximity. CONSERVATION AND RECOVERY PLAN In September 1989 the B.S.B.I. Rare Plants Translocation Panel agreed that S$. germanica plants could be raised from seed and reinstated, at the rosette stage, in the wild, as part of a recovery programme. About 100 seeds were collected from each of three sites, labelled and then refrigerated. In the following spring/early summer the seeds were sown in three separate containers, the subsequent germination rate being 25, 34 and 50 per cent. The young plants were later transferred to individual pots, and after late September/early October rains ended the long drought of 1990, a total of 112 leafy rosettes were planted at the parent sites. These reintroductions will help to strengthen populations, assist continuity of flowering, and increase seed banks in the soil. Past and present efforts to warden sites, protect plants and initiate site clearance could not have been sustained without the co-operation of a few local botanists and, in 1988 and 1989, several members of staff of the N.C.C. The aims of the immediate future for these three sites are to continue the management work and maintain the interest and support of site owners. In the long term, it is recommended that a few old sites which are still suitable for supporting populations should be opened up, and appropriately managed, within the next five years. This would help to make the future of S. germanica more secure and possibly change its status from threatened to rare. It would be one example of a successful recovery plan. However, if these measures fail, consideration should OBSERVATIONS ON STACHYS GERMANICA 367 be given to the creation of one or two carefully selected sites in places where the species is known to be native. In the meantime, these recent observations have shown that it is not only land management practices that continue to threaten the survival of S. germanica in Britain, but also climatic and animal factors. ACKNOWLEDGMENTS I wish to thank the British Ecological Society, the Nature Conservancy Council, and The Wild Heritage for grants received in support of this study; Miss Lynne Farrell, N.C.C. Chief Scientist’s Directorate Specialist for lowland heaths and rare plants, for helpful criticism and advice; Dr Michael P. Kerney (Mollusca Section) and Mr. Len Ellis (Bryophyte Section), Natural History Museum, for identifying, respectively, snails and mosses; Mr Chris O’Toole, The University Museum, Oxford, for similar assistance with bumblebees and other insects; and the many botanist friends with whom I have shared interesting discussions and management tasks on behalf of Stachys germanica. REFERENCES Druce, G. C. (1886). The flora of Oxfordshire. Oxford. Druce, G. C. (1927). The flora of Oxfordshire, 2nd ed. Oxford. Dunn, A. J. (1987a). Observations on Stachys germanica L. at a new site in Oxfordshire. Watsonia 16: 430-431. Dunn, A. J. (1987b). Stachys germanica L. in an Oxfordshire garden. B.S.B.I. News 47: 29. GERARD, J. (1633). The herball or generall historie of plantes. Revised by T. JOHNSON. London. MarreENn, P. M. (1988). The past and present distribution of Stachys germanica L. in Britain. Watsonia 17: 59-68. Moorg, D. M. (1987). Labiatae, in CLAPHAM, A. R., TuTIn, T. G. & Moore, D. M. Flora of the British Isles, 3rd ed. Cambridge. PERRING, F. H. & FARRELL, L. (1983). British red data books: 1. Vascular plants, 2nd ed. Lincoln. Stace, C. A. (1975). Hybridization and the flora of the British Isles. London. (Accepted January 1991) . i \, byneatier biW wif bas jena yousvesencD oie! tend Wid 9.9.4 Stove one t a - au De 7 csoivba bue gain Loigised ol .,2inaiey fim atitwod basiwol 3 nor lmutet loodesé stvdqoyilly ie esd bas (eotes2 so2diipy (imeavin J 24% 2leoTO ent WM jarom dmoe .ylovitvoqest ,ggivitasbhi: ot us tainatod yaam ac) bes sepasent waco has tiw sonetzizss welegiz 102 ,b oO, ryztwue bo iledsd ne etea? tieecandor bore sue gainers bona oved mosh fiwe ety - Ldo* LAT Live trtated 5 So ir grew Wi S 44 _ to oso) Utabmsipgtat) 3 pranadiislor: vi) Sethe Sak i ae ray! a i ati abe 7 Joo sorte and Wings Or boodantyendl 988 ae mal brit. boat, eid acnincte cy | + - a tinal hat? Ake @reta ani ger gl ipsa IG hei pee wrth eo ee Oa? 7 eo |4 bth a ee an ee er eae 58 Git pecaneoy tate ltG » (GPRE OR Se eae ; ratte pan whoky 25 styl %, awn Navas an ge . ‘ vy Mii fs icy oF WPS, ee ie * Poe vu mad? {aaessq vs Jay _ aie Bat cio MU, ee aT. Yas MAVRAI Bulg ¢ r 7 ae ye GF petrgliemstor NERS TH ‘Toe Heono) aie) dein’ st lo sali oA bre Holte cihrdyet ACkRT) ‘ 4 Cd nv 4 : Lay te a omer 1 (evi Cwai tn, thro edits eciry ; un “het e ek *.* r ry iL. hes beco report ie - 9 : eé i] ed ry A ity thé wikd. i part dag : ’ 43% : C cel vee refige ) hi fate conta ta enlsom #ere letes te he lone Qfoveht of 98 ‘ ie, Wi tery io stra ah ee i * a 7 : ; ia vis lenranee aaa me . mi tt (SSK ane 1509 ; Giadi' uhiect cm Mage n. OOF vwrtierd, it te log (tin osh i 4togea ai? hap oc 7 al ® ; ti it “vod to rat BS Watsonia, 18, 369-379 (1991) 369 The distribution of hybrid Marram Grass, < Calammophila baltica (Fligge) Brand, in the British Isles J. R. RIHAN Department of Biology, The University, Southampton, SO9 5NH and A. J. GRAY Institute of Terrestrial Ecology, Furzebrook Research Station, Wareham, Dorset, BH20 5AS ABSTRACT The geographical distribution of < Calammophila baltica (Fligge) Brand (= x Ammocalamagrostis baltica (Schrad.) P. Fourn.), in the British Isles is described, and aspects of its ecological distribution and limits are discussed, especially in relation to Ammophila arenaria (L.) Link. INTRODUCTION The hybrid Marram Grass, X Calammophila baltica (Fligge) Brand (= X Ammocalamagrostis baltica (Schrad.) P. Fourn.), is a naturally occurring intergeneric hybrid, derived from Ammophila arenaria (L.) Link and Calamagrostis epigejos (L.) Roth (Westergaard 1943). It is rare in Britain, being reported only from the East Anglian coast and around Holy Island, Northumberland. It is found on sand dunes coexisting with one of its progenitor species, A. arenaria. On the coasts of mainland Europe the hybrid is found in three varieties depending on the relative contributions of the genomes of the progenitor species (Westergaard 1943). The British populations have all been identified as belonging to X C. baltica var. subarenaria (Hubbard 1975), the variety to which the A. arenaria genome makes the greatest contribution. Ammophila arenaria, Marram Grass, is found on all coastal dune systems in Europe and North Africa, from the Mediterranean to about 62° N. (Tutin 1980), including the whole of the British Isles. It is occasionally found inland and on the sandy shores of lakes (Westhoff et al. 1970). Doing (1985) describes A. arenaria as occurring from Norway to western Spain, and A. arenaria var. arundinacea on Mediterranean coasts. It has also been transplanted to the Pacific coast of the North American continent (Pohl 1968). In the British Isles Marram is found wherever a suitable habitat occurs, usually in coastal sand dune systems. It is common on the coasts of both East Anglia and Northumberland. Marram is a very important component of the sand dune successional system, and largely controls the rate of dune growth and sand stabilisation. As such it is widely planted in order to protect beaches threatened by erosion. Calamagrostis epigejos, Bush Grass, is found in damp and disturbed areas throughout Europe, though it is less common in the south-west. It may be found on sand dunes around Baltic and northern European coasts (Westergaard 1943; Doing 1985). In Britain this species appears to be more rare than in former years, which may be a reflection of the changes in agricultural practices over the past 100 years. It is less common than it was in East Anglia, and is now absent in Cheviot (v.c. 68). x Calammophila baltica, Hybrid Marram, appears to be confined to north-western coasts of Europe, from Dunkirk in the south (van den Berghen 1964) to somewhat south of 60° N. (Hultén 1971). It is common in parts of southern Sweden and Norway, in Poland, Russia, Denmark and the Netherlands (Doing 1985), where, together with marram grass, it is widely planted for sand dune 370 J. R. RIHAN AND A. J. GRAY conservation. Indeed in many places, because of its apparent vigour, it is preferred over marram for this use. The history of the origin and spread of Hybrid Marram in Britain has been complicated by the failure of early authors to recognise it as a separate taxon. The earliest reliable dates appear to be 1871 in Northumberland (Richardson 1872; Trimen 1872) and 1885 for East Anglia (Ellis 1960). At the latter site, however, it is possible that a hybrid individual was found over 80 years earlier (Ellis 1960). When the present study began, Hybrid Marram was thought to occur in Northumberland, on Holy Island and Ross Links, and at Alnmouth (Swan, pers. comm.); and in East Anglia, at Holme- next-the-Sea on the north coast, and from Waxham to Sizewell on the east (Ellis 1960). Much of the East Anglian distribution has been altered by the transplanting of marram from Winterton for dune rebuilding works, particularly after the tidal surge of 1953 (Ellis 1960; Taylor & Marsden 1983). It seems unlikely that the hybrid was knowingly chosen for transplanting, but its larger size and general vigour would encourage the men collecting to take it, rather than A. arenaria, when they were growing side by side. There is a record for the hybrid on Handa Island, off the Sutherland coast, one of the few places in Britain where A. arenaria and C. epigejos occur together (Heslop-Harrison & Heslop-Harrison 1938). This has never been confirmed despite several intensive searches which have been made for it over the last 50 years. It is still reported as being present (though rare) on the island in the most recent Flora of Sutherland (Kenworthy 1979). Hybrid Marram is also known to have been accidentally transplanted, as a consequence of taking marram for dune conservation purposes from Winterton, to Milford on Sea, Hampshire. It is known that marram has been exported to many parts of the country (and, indeed, the world) (E. A. Ellis, pers. comm.) from Winterton, and it seemed highly likely that other such populations may exist. In Britain, Hybrid Marram is generally described as a foredune plant (Robertson 1955) or evena coloniser of bare beach sand (Ellis 1960). METHODS DISTRIBUTION All the sites where X C. baltica had been recorded (records taken from Perring & Sell (1978), Swan, pers. comm., and E. A. Ellis (1960 & pers. comm.)) were visited between August 1979 and August 1982. Adjacent beaches, identified as sandy on large scale Ordnance Survey maps were also searched. A similar survey was carried out in East Anglia and Northumberland in 1987. Handa Island had also been visited in 1978, prior to the commencement of this study. The beaches were reached from suitable points inland, and approximately one hour spent on each area searched. The results were recorded on a visual estimate of abundance; one record was made for each area surveyed. ECOLOGICAL LIMITS In order to determine the ecological limits of x C. baltica a survey was carried out in both East Anglia and Northumberland. Ten 2 m X 2 m quadrats were taken at random within 50 m x 50 m areas selected to represent each successional stage of the dune systems. Ten areas were surveyed in East Anglia, all in the Winterton/Horsey dunes, and four in Northumberland, at Ross Links and Budle Bay; this represents the greater heterogeneity of the East Anglian dune system. A number of variables were recorded, including the percentage cover of each species, the length of the longest (vegetative) tiller, and the number of flowering tillers. The cover abundance of x C. baltica and A. arenaria were compared for evidence of the apparent coexistence of the two species, and the performance data were examined for competitive effects between them. The performance data of x C. baltica only were examined for evidence of differential distribution of the species across the different dune stages. RESULTS AND DISCUSSION THE NORTHUMBERLAND POPULATIONS The North Northumberland coastline was searched from Berwick upon Tweed, in the north, to DISTRIBUTION OF x CALAMMOPHILA BALTICA 37] Berwick upon Tweed a\o Oo 4 2 Holy Island s) 8 Ross Links @. slo 7 maul) alg) col .coneti 16 bree) 5 pedi : oa Arar J — = ax ped “T wrwnmlisnigish. * = aah, vy vate 2 sunk A Myers (2801) a ! ound .TTe-4TF seek A iush wh Ye a of sav mo Ot ivi ee is , rojorsonte seabed \ rand thirtst a b y LTS He ‘oucdnibs Sauhrutian oe ely eqnadink nae ie? uw nits Yes wen Sin Sanat ye acai ABILT) abst at, MP sleing cwol beth seat) + 6 wand Ai wih ah es . 108 é 5, pax lier s ‘ether! fi yo wore, Adal). Y 4 iibV. Ob} raph Wh mannmess opel ies gi lo qpaicos AT AeeeT} A ay A ne © Pit ah AioerooMt “DbAoe~ ob STEN an ith apomed (6PC11.a A oe va AS 4 | ’ nobrot th ? hos 1)? )o SF eodkod oncne) (O08 on 4 | : rey py t nique, | (eT) OT ie p ot. Beet? £29) moors! sav (EOL). Mi. ROE rl eran tinct: . a. Seseiies. ReGe 3K We Ae > BF ai? — Toa? se alent oh —— ate Gidiptae 4%) voltediad = i . : P — . wis A Ma> i> ey Os rt “+ paint aigalibts ah = *< . 7 —. ch: habit to fi het Garren, ‘oar eon : . ‘ > ‘ “ nS ond J ley CE 2 é = iil * e ™ “@ s - J 4 4 rE ReOt, 40 —_- a " oe “a 4 %~ e! A ' J S 2 AGS. “a abe = 4 /- : ' i ¥ sa & = , ~~. r. ° ee rannih Sat 4 — : ie ssc Te Tey ral} J R “_—T> " dea! . I-A oes ai MT Tiel tmis organs wizee ‘avtse# xi bse ah ne Y ‘ov. ts Pa .i } * ’ } e ’ tp er Watsonia, 18, 387-390 (1991) 387 Orchis purpurea Hudson in the Avon Gorge, Bristol A. J. WILLIS Department of Animal and Plant Sciences, The University, Sheffield, S10 2TN M. H. MARTIN Department of Botany, The University, Bristol, BS8 1UG and K. B. TAYLOR 28 Berkeley Road, Bristol, BS6 7PJ ABSTRACT The discovery of a single flowering specimen of Orchis purpurea Hudson (Orchidaceae) in Leigh Woods, Avon Gorge, Bristol, in May 1990 is reported. The main features of this Carboniferous limestone habitat are briefly described and a short account of the associated vegetation is given. The occurrence of the Lady Orchid in this site is discussed with reference to its distribution in Britain and on the Continent and to its phenology and modes of dispersal. INTRODUCTION A single specimen of the Lady Orchid (Orchis purpurea Hudson) was found (KBT, conf. MHM, AJW) on the Carboniferous limestone of Leigh Woods in the Avon Gorge, Bristol, v.c. 6, on 6 May 1990. The plant was well grown, with five large leaves and a smaller one, and an inflorescence 42 cm tall bearing about 30 flowers, nearly all fully open when first seen. A single flower was taken on 9 May as a voucher specimen (herb. AJW). The labellum was flushed pink, with distinct reddish- purple spots throughout. The lateral lobes of the labellum were quite well developed, the large median lobe bearing a small but distinct central tooth between its two lobules. Of the two variants of the species recognized by Rose (1948) in Kent, the Leigh Woods plant resembled the western one more closely than the eastern one, having a fairly dense flowering spike and short ovary, as well as heavy spotting on the lip. HABITAT The site was a wooded slope in Leigh Woods on the Somerset (W.) side of the Avon Gorge, where the soil is deeper and moister than on the more precipitous eastern city side (Gloucestershire). The angle of slope in the general area was about 20° and the aspect approximately south-east. The major contributor to the canopy was Fraxinus excelsior, these small trees being mostly 3-5 m tall (with some up to 6 m) and 3-8 cm in diameter (at breast height). Although the canopy was fairly complete, there were some breaks, with sunflecks reaching the ground flora. The soil was moderately moist and quite deep for this woodland, ranging from 28 to 52 cm, with a mean value (10 estimates, by use of a thin steel probe) of 37 cm. Fragments of limestone were present but there was only a little exposed rock in the general vicinity. When dry, the surface soil colour was a very grey brown, becoming blackish-brown when wet. This indicated organic enrichment, the surface 388 A. J. WILLIS, M. H. MARTIN AND K. B. TAYLOR containing a substantial amount of leaf mould. Below this enriched layer was a reddish silty B horizon. With respect to soil texture, a sample when wet moulded easily into fragile ‘fingers’ indicative of an organic rich silt loam in the distinct A horizon. In the surface soil the pH (direct readings after dampening with deionized water) was 6-7—7-0 and in the sub-surface (at about 2 cm) was 7-0-7-1. VEGETATION Although there were no very large trees at the site, the tree layer was substantial, with abundant Fraxinus excelsiar, frequent Corylus avellana and occasional Acer pseudoplatanus and Ulmus glabra. Nearby were Acer campestre, Crataegus monogyna, Rubus fruticosus agg., Sambucus nigra and Viburnum opulus. The ground flora was moderately open, and quite strongly dominated by Mercurialis perennis. Also present were Arum maculatum, Hedera helix and Hyacinthoides non- scripta, together with the ferns Dryopteris filix-mas and Asplenium scolopendrium. The quite thick moss carpet included Brachythecium rutabulum, Eurhynchium swartzii, Plagiomnium undulatum and Thamnobryum alopecurum, with other mosses nearby characteristic of soil and rocks of these woods (Willis 1964). Additional species in the vicinity included Brachypodium sylvaticum, Carex sylvatica, Circaea lutetiana, Deschampsia cespitosa, Festuca gigantea, Geranium robertianum, Lathraea squamaria (on Corylus), Melica uniflora, Taraxacum officinale agg., Urtica dioica and Viola riviniana. THE DISTRIBUTION OF O. PURPUREA In Britain this orchid has a distinctly south-eastern distribution (Summerhayes 1951; Perring & Walters 1962). It is now known almost only from the chalk in Kent, where it grows in scrub coppice under moderate light and shelter from extreme dryness and wind, and in chalk escarpment beechwoods in fairly deep shade, mostly on the lower, south-facing slopes (Rose 1948). It was recorded from N. Essex in 1738 (Hall 1936), from a site apparently on chalky boulder clay in Walter Belchamp Parish (the date of 1935 for this record implied by Rose (1948) and Lang (1980) is misleading). Orchis purpurea was also formerly known in Surrey where the last record is for 1959 (Leslie 1987). In W. Sussex, where the orchid was always very rare, the most recent published record is also for 1959 (a good year for flowering), on chalk at Chanctonbury Ring (Hall 1980). However, in 1976 a single flowering specimen was found in a site nearby, but unfortunately this was completely destroyed by cattle several years later (Mrs M. Briggs, pers. comm.). In 1961 a flowering specimen was found on the chalk in southern Oxfordshire under beech with Mercurialis perennis. It was last seen three years later but in 1986 seven plants (three flowering) were found in a cleared and disturbed area in the same vicinity (Kemp 1987). The colony is still flourishing, flowering well in 1990 (R. J. Kemp, pers. comm.). There is a further record of a single plant from the Lower Old Red Sandstone in Herefordshire in 1967 “under hazels in hazel coppice at edge of beech wood on calcareous subsoil” but this plant has not reappeared (Whitehead 1976). This is the most westerly site for the plant in Britain, Leigh Woods being a little to the east. Virtually all of the British records refer to chalk, the Leigh Woods plant appearing to be the only occurrence on limestone. Orchis purpurea has a wide distribution in continental Europe where it occurs typically on calcareous base-rich soils, including limestone, as for example in France and Italy. It extends southwards and westwards from Denmark to Germany, France and Spain, and is abundant in the mountains of central Italy. Orchis purpurea, classified by Summerhayes (1951) as a member of the southern Eurasian geographical element, is present in many countries bordering the Mediterranean including Algeria and Tunisia. In the east it extends to Greece, Crimea, Caucasus and Turkey. Ellenberg (1988) gives O. purpurea as a member of the alliance Quercetalia pubescenti-petraeae of xerothermic mixed Oak woods in Central Europe and for its ‘ecological indicator values’ lists half- shade, warmth indicator, suboceanic, moderate dampness, and mostly on chalk and limestone. ORCHIS PURPUREA IN THE AVON GORGE 389 DISCUSSION Orchis purpurea may be a recent arrival in Leigh Woods, but could possibly have been present, especially if in only a vegetative state, for many years. Whatever the case, it has every appearance of a native, and is probably not a recent transplant. Leigh Woods are undoubtedly ancient (Rackham 1982), with a wealth of ancient woodland indicator species (Lovatt 1989), and with a rich limestone flora containing many rarities (White 1912; Hope-Simpson & Willis 1955; Willis 1962). The conditions in Leigh Woods are closely similar to the ‘ecological indicator values’ for O. purpurea given by Ellenberg (1988); also the dominance of the ground flora by Mercurialis perennis fully accords with the statement by Rose (1948) that this species is “the most constant and characteristic associate of O. purpurea in Britain’’. Rose (1948) regards O. purpurea as well ‘at home’ in the mountains of central Italy where it is the most abundant woodland orchid, the winters and springs are cool and wet and the summers extremely dry and sunny. These particular conditions, according to Rose (1948), under which the tubers dry out in the summer, may lead to good seasons for this species. The two last summers (1988, 1989) were quite hot and in 1989 very dry; these climatic conditions, coupled with the very mild winters in these years, may well have favoured the plant in Leigh Woods, leading to its flowering. A further factor may have been the favourable growth conditions for early developing species of the woodland floor in 1990 (leaves of O. purpurea may appear above ground in February), enabling good growth to be made before the tree canopy expanded. However, Rose (1948) considers that conditions in the previous year are those which have the most influence on the performance of the plant in the following year. Certainly the high vigour shown by flowering spikes of O. purpurea in Kent in 1990 is in full accord with the view that drying of the tuber in summer is beneficial. Although O. purpurea is now largely restricted to Kent in Britain it is possible that the very small seeds may be carried long distances — even across the Channel (Summerhayes 1951); indeed it may be a fairly recent immigrant from the Continent (Rose 1948), although known in Britain since 1666. The origin of the Leigh Woods plant from seed unwittingly carried by a visitor (well-travelled botanists come to the Woods) is not impossible. Another possibility is from wind-borne seed from a batch of orchid species, including O. purpurea, grown in 1980-1982 (O. purpurea was known to flower and possibly fruit) in the University of Bristol Botanic Garden, North Road, Leigh Woods, from material brought from France. However, scrutiny of photographs of the flowers taken in 1980 of the garden-grown plant shows several differences in shape and markings of the labellum from the Leigh Woods plant, and the specimen of French origin seems to match the eastern rather than the western variant of the species. The existence of single plants in sites is known, as, for example, in Herefordshire and (originally) in Oxfordshire. It is well established that only a small proportion of plants in a population of O. purpurea usually flower in any one year, but no purely vegetative specimens have been seen in Leigh Woods. If the plant persists here it will be a remarkable addition to an already notable flora. ACKNOWLEDGMENTS We are indebted to C. D. Preston for information about the distribution of O. purpurea in Britain, to R. J. Kemp for details of its performance in Oxfordshire, and to Mrs M. Briggs with respect to Sussex. The support of the Leverhulme Trust (to A.J.W.) is gratefully acknowledged. REFERENCES ELLENBERG, H. (1988). Vegetation ecology of Central Europe, 4th ed. Cambridge. HALL, P. C. (1980). Sussex plant atlas. Brighton. HALL, P. M. (1936). Plant Records. Rep. botl Soc. Exch. Club Br. Isl. 11: 41. Hope-Simpson, J. F. & Wiuis, A. J. (1955). Vegetation, in WHITTARD, W. F. & McInngs, C. M., eds. Bristol and its adjoining counties, pp. 91-109. Bristol. Kemp, R. J. (1987). Reappearance of Orchis purpurea Hudson in Oxfordshire. Watsonia 16: 435-436. Lana, D. (1980). Orchids of Britain. Oxford. 390 A. J. WILLIS, M. H. MARTIN AND K. B. TAYLOR Lesuigz, A. C. (1987). Flora of Surrey — Supplement and checklist. Guildford. Lovatr, C. M. (1989). The historical ecology of Leigh Woods, Bristol. Proc. Bristol Nat. Soc. 47: 3-19. PERRING, F. H. & WALTERS, S. M. eds. (1962). Atlas of the British flora. London. RAcKHAM, O. (1982). The Avon Gorge and Leigh Woods, in BELL, M. & Limprey, S. eds. Archacolosea aspects of woodland ecology. Oxford. Rose, F. (1948). Orchis purpurea Hudson, in Biological Flora of the British Isles. J. Ecol. 36: 366-377. SUMMERHAYES, V. S. (1951). Wild orchids of Britain. London. Wuite, J. W. (1912). The flora of Bristol. Bristol. WHITEHEAD, L. E. (1976). Plants of Herefordshire. Hereford. WIiLuis, A. J. (1962). Rare plants of the Bristol district. Rep. Ann. Conf. 1961 and 1962, South Western Nat. Union. 4 pp. WILLIs, A. J. (1964). Bryophytes of Leigh Woods, Somerset. Proc. Bristol Nat. Soc. 30: 451-454. « (Accepted September 1990) ~ Watsonia, 18, 391-393 (1991) 391 Differential pollination efficiency within a hybrid swarm between Dactylorhiza purpurella (1. & T. A. Stephenson) So6 and D. fuchsii (Druce) So6 E. HAZELDON, T. NAISBITT and A. J. RICHARDS Biology Department, The University, Newcastle-upon-Tyne, NEI 7RU ABSTRACT Pollination efficiency, as estimated by frequency of pollintum removal, was studied in a mixed population of Dactylorhiza purpurella (T. & T. A. Stephenson) So6, D. fuchsii (Druce) S06 and their hybrid (Orchidaceae). Plants of D. purpurella were visited more frequently by vectors than were those of D. fuchsii or the hybrid. This may restrict levels of hybridization. INTRODUCTION Hybridization has frequently been reported between diploid (2n = 40) Dactylorhiza fuchsii (Druce) Soo and tetraploid (2n = 80) marsh orchids sometimes classed within D. majalis (Reichenb.) P. F. Hunt & Summerhayes (Bateman & Denholm 1983). Lord & Richards (1977) found backcrossing in a Co. Durham population hybrid between D. fuchsii and D. purpurella (D. majalis subsp. purpurella (T. & T. A. Stephenson) D. M. Moore & So6), such that presumptive backcrosses had aneuploid chromosome numbers. They suggested that the degree of contribution to the hybrid made by D. purpurella could be estimated by the chromosome number. Complex hybrids within such hybrid swarms appear to be fairly fertile, suggesting that D. fuchsii may itself be ancestrally polyploid. Dactylorchids only set seed after pollinia are transferred from the anther of one flower into the stigmatic cavity of another as the result of an insect visit. There appears to be no published _information concerning visitors or rewards for D. majalis agg., although we have frequently observed and photographed Bombus spp., especially the queens of B. terrestris, visiting the flowers of D. purpurella. D. fuchsii is visited by B. lapidarius, B. terrestris and Apis mellifera (Dafni & Woodell 1986). Other visitors including various diptera and lepidoptera do not transport pollinia. Those authors record an average fruit set for D. fuchsii of 54%, but note that the lower (earlier) flowers set more than twice as many capsules as the upper flowers on the spike. Grant (1981), for instance, provides many examples of potentially interfertile sympatric species- pairs whose specific identity is largely preserved by ethological isolation. Hybridization following the breakdown of ethological isolation is particularly frequent in the Orchidaceae (review in Dafni (1986)), notably in Orchis and Ophrys. Very little attention seems to have been paid to the pollination of hybrids, perhaps because they are frequently sterile. Although it has been suggested that fertile hybrids may be maladapted to successful pollination in comparison to their parents (e.g. Dafni 1986), and thus by implication that this maladaptation reinforces the genetic isolation of their parents, only Grant (1952) provides any data to this effect. In the present study, pollination efficiency, judged by rates of pollinium removal, is studied in a hybrid swarm between D. purpurella and D. fuchsii. METHODS Every plant in a mixed colony of D. purpurella and D. fuchsii at Malham Tarn Fen, N.W. Yorks (v.c. 64) (GR 34/887.672) was non-destructively sampled. A hybrid index was formulated as 392 E. HAZELDON, T. NAISBITT AND A. J. RICHARDS TABLE 1. CHARACTER STATES USED IN THE HYBRID INDEX Score 0 2 Leaf spotting zero, Or punctate near apex larger spots throughout Labellum colour red-purple pale lilac Labellum pattern loops spots Ratio spur length/width at base <4-0 >5-0 Ratio length labellum mid-lobe/length labellum <0-2 >0-4 follows. Plants were assessed for five characters by which D. purpurella and D. fuchsii differ (Table 1), based on Heslop-Harrison (1957) and Lord & Richards (1977). Plants judged to be intermediate for a character were scored one. Using this simple index, it is considered that plants with a total score of 0 are morphologically typical D. purpurella, and those scoring 10 are typical D. fuchsii. To estimate pollen fertility, pollinia were squashed in acetocarmine and examined at X 400. Tetrad cells with regular nucleus formation were scored as fertile, and those with absent or abnormal nucleus formation were scored infertile. Percentage fertility was based on samples of over 50 pollen cells. To estimate the frequency of pollintum removal, the position of each open flower on the spike and the number of pollinia removed from it was noted. RESULTS AND CONCLUSIONS 1. Most plants in the population are D. purpurella, or hybrid. From hybrid index scores, it is likely that only between one and three plants are referable to D. fuchsii (Table 2). 2. Most plants show a high level of apparent pollen fertility, and consequently this character is of little use in assessing hybrid status. However the relatively low apparent fertility of plants scoring 6 suggest that these might be F, hybrids (Table 2). 3. There is a progressive reduction in the proportion of pollinia removed from flowers as the hybrid index score increases (Table 1). There is a significantly greater proportion of plants from which some pollinia have been removed for plants scoring 0-3 (plants resembling D. purpurella in floral characters) than from those with higher scores (D. fuchsii and hybrids) (Table 3). 4. There is no overall significant difference in flowering time or vigour as assessed by the number of open flowers, including the bottom-most, between plants of different hybrid index scores (Table 3). It is therefore concluded that differences in levels of pollintum removal are not due to the size or maturity of the flowering spikes, but result from pollinator choice. TABLE 2. HYBRID INDEX SCORES, AVERAGE POLLEN FERTI- LITY, AND AVERAGE PERCENTAGE POLLINIUM REMOVAL OF PLANTS SAMPLED, + STANDARD DEVIATION Hybrid index Average pollen Average percentage No. of score fertility pollinium removal plants 0 84-6+19-6 36:6+ 6-7 22 l 91-5+ 8-1 39-4+ 5:8 14 2 89-94+11-3 29-0+ 5-6 16 3 89-7+ 7-7 28-5+ 3-1 14 4 94-1+22-4 12-5+15-8 4 3 83-5+18-6 17-4+17-0 5 6 57:-9+ 9-9 0 3 7 92-34 3-2 8-5+12-0 2 8 33:3 0 l 9 87-5 0 l 10 78:8 0 | eee POLLINATION EFFICIENCY IN A DACTYLORCHID HYBRID SWARM 393 TABLE 3. POLLINIUM REMOVAL AND MEAN NUMBER OF FLOWERS OPEN FOR PLANTS WITH DIFFERENT CLASSES OF HYBRID INDEX SCORES Pollintum Hybrid index score removal 0-3 4-6 7-10 No. of plants All intact 22 i) 4 33 Some removed 43 5 1 49 Chi-square (df = 2) = 6-33, p<0-05 Average number 2-88 3-42 3-0 of flowers open 5. It is concluded that D. purpurella-like plants in this population are more efficiently visited by insects than hybrids. This may restrict levels of hybridization. From a very small sample, it seems that the minority species D. fuchsii may also be poorly visited, perhaps because it presents a different, unusual, search-image to visitors. It is possible that hybrids also suffer by presenting a different search-image to visitors in comparison with the dominant D. purpurella. However, it is noteworthy that although Bombus spp. which visit D. purpurella were commonly present, few if any Apis, the major pollinator of D. fuchsii (Dafni & Woodell 1986) were observed. REFERENCES BATEMAN, R. M. & DENHOLM, I. (1983). A reappraisal of the British and Irish dactylorchids, 1. The tetraploid marsh-orchids. Watsonia 14: 347-376. Darn, A. (1986). Pollination in Orchis and related genera, in Arpitt1, J., ed. Orchid biology, reviews and perspectives, pp. 81-103. Ithaca. Darni, A. & WoopDELL, S. R. J. (1986). Stigmatic exudate and the pollination of Dactylorhiza fuchsii (Druce) So6. Flora 178: 343-350. GRANT, V. (1952). Isolation and hybridization between Aquilegia formosa and A. pubescens. Aliso 2: 341-360. GRANT, V. (1981). Plant speciation, 2nd ed. New York. Lorp, R. M. & Ricuarps, A. J. (1977). A hybrid swarm between the diploid Dactylorhiza fuchsii (Druce) S06 and the tetraploid D. purpurella (T. & T. A. Steph.) So6 in Durham. Watsonia 11: 205-210. (Accepted December 1990) ie ' 7 ae eee eas af bs - Boe HAE ini! peerig Gi motes id br i 449 re eh oy iat SOP ry J Teer Ses ores A shee qe I feal t+ er niee' inhi eta 0 oni oO fh d IRE S1BS4 36s RST 4% a inside cher west hed cdaty oatini esha aa eSeel af any P : ~_aee ae ‘Sy “er _ co) YO, SIT GF 4 k kan tl wala < ev “6 "7 —28 tz Watsonia, 18, 395-399 (1991) 395 A new dactylorchid hybrid F. HORSMAN 7 Fox Wood Walk, Leeds, LS8 3BP ABSTRACT An account is given of a new hybrid from Cardiganshire (v.c. 46), Dactylorhiza x dinglensis (Wilmott) So6 nothosubsp. robertsii F. Horsman, nothosubsp. nov. (D. majalis (Reichenb.) P. F. Hunt & Summerhayes subsp. cambrensis (R. H. Roberts) R. H. Roberts x D. maculata (L.) S06 subsp. ericetorum (E. F. Linton) P. F. Hunt & Summerhayes). INTRODUCTION In 1987 I visited a locality near Borth (Cards., v.c. 46) which was one of the two sites from which Roberts (1961b) described Dactylorhiza majalis (Reichenb.) P. F. Hunt & Summerhayes subsp. cambrensis (R. H. Roberts) R. H. Roberts and which has the largest known population of that taxon. In 1988 I counted nearly 1500 flowering spikes. This suggests the population has maintained its size since Roberts first saw it 30 years ago (Roberts, pers. comm. ). Ellis (1983) records D. majalis subsp. cambrensis from only four 10-km squares in Wales. In common with Roberts (1961a), the only other dactylorchid I found in the Borth locality in 1987 and 1988 was D. maculata (L.)So6 subsp. ericetorum (E. F. Linton) P. F. Hunt & Summerhayes. In 1987 I observed plants intermediate in morphology between D. mayalis subsp. cambrensis and D. maculata subsp. ericetorum which were presumably hybrids, a conclusion confirmed by Roberts. A further visit was made to the site in 1988 to study the hybrid and its putative parents. The hybrid was noticed on my first brief visit because its spur seemed to be intermediate between the two other taxa present, but only a few such plants were seen. In 1988, however, I found the hybrid to be common. The pale lilac colour of the flower, together with the shape of the lip and its markings, were sufficiently distinctive to enable a reliable estimate of the number of hybrids to be made. The Borth locality consists of wet meadowland and grassland on old stabilised sand dunes, both of which have a rich flora. The dactylorchids are confined to the wet meadowland. The hybrid is common on the two sides of the site which adjoin an old river bed containing a brackish-water marsh. D. maculata subsp. ericetorum is most common on one of these two sides where a wide range of intermediates between it and D. majalis subsp. cambrensis is concentrated. It is intriguing to note that Stephenson & Stephenson (1921), in describing Orchis latifolia from this site, could well have been referring to this hybrid when they stated: ‘““Here are many dark forms along with some lighter ones which may be hybrids, where O. ericetorum is present, but no trace of O. praetermissa’’. The hybrid is confined to the Borth site, exact details of which are witheld for conservation reasons. No other sites are known where D. maculata subsp. ericetorum and D. majalis subsp. cambrensis grow in close proximity. Evidence of the presence of a third dactylorchid was found at the site in 1988. A solitary flowering spike of what Roberts suspects, from my colour photographs, may have been an F; hybrid or backcross derived from D. fuchsii and D. majalis subsp. cambrensis was found. It was growing on the border of the wet meadowland and the grassland of the old sand dunes. As D. fuchsii grows fairly close by, such a find was not surprising. TAXONOMY AND NOMENCLATURE Bateman & Denholm (1983) made D. praetermissa, D. traunsteineri and D. purpurella subspecies of D. majalis. Heslop-Harrison (1954) treated them as species, a view with which I concur. In the 396 F. HORSMAN British Isles D. majalis has three subspecies, subsp. occidentalis (Pugsley) P. D. Sell, subsp. cambrensis (R. H. Roberts) R. H. Roberts and subsp. scotica Nelson. D. majalis subsp. majalis is confined to Continental Europe, where it is also represented by subsp. alpestris (Pugsley) Senghas. Bateman & Denholm (1983) reduced D. majalis subsp. cambrensis to D. majalis subsp. occidentalis var. cambrensis (R. H. Roberts) Bateman & Denholm. The reduction of D. majalis subsp. cambrensis to varietal status is rejected for the following reasons. Bateman & Denholm (1983) state that Roberts (1961b) separated D. majalis subsp. cambrensis from D. majalis subsp. occidentalis primarily by the broader leaves and narrower spurs of the plants in the population of D. majalis subsp. occidentalis measured by Heslop-Harrison (1953), in Co. Clare, v.c. H9, Ireland. D. majalis subsp. cambrensis is not accepted as a subspecies by Bateman & Denholm (1983) because “Unfortunately, this population was atypical of var. occidentalis in these characters”’. Heslop-Harrison (1953) measured the maximum width of the /ongest leaf, and the width of the spur at about aynillimetre from the mouth when flattened. He emphasises that the ‘width’ figure represents, therefore, not the diameter, but approximately half the circumference at this point. Bateman & Denholm (1983) measured the maximum width of the widest sheathing leaf (it must be pointed out that the widest leaf is not necessarily the longest), and the width of the spur at the entrance when flattened. Heslop-Harrison’s and Bateman’s & Denholm’s data are not, therefore, compatible in these two respects. However, Roberts’ (1961a, 1961b) data are compatible with those of Heslop-Harrison (1953). Hybrids involving Dactylorhiza majalis subsp. cambrensis have previously only been reported from Anglesey (v.c. 52) (Ellis 1983). The other putative parents were D. purpurella (T. & T. A. Stephenson) Soo and D. fuchsii (Druce) So6 subsp. fuchsii. This paper reports the occurrence of a third hybrid with D. majalis subsp. cambrensis which is previously undescribed. The name D. X townsendiana (Rouy) Soo has previously been used for hybrids between D. maculata and D. majalis (as defined here). The basionym for this name is Orchis X braunii Halacsy var. townsendianus Rouy in Rouy & Fouc., Fl. Franc. 13: 173 (1912). This in turn is based on Orchis latifolia-maculata Townsend, Fl. Hampshire 2nd ed., 409 (1904), hence the specific epithet. D. majalis does not grow in Hampshire and I believe the plant is D. maculata X praetermissa. The name D. X dinglensis (Wilmott) Sod is, however, almost certainly referable to D. majalis subsp. occidentalis X maculata subsp. ericetorum and that must be called nothosubsp. dinglensis. The new hybrid between D. majalis subsp. cambrensis and D. maculata subsp. ericetorum I call nothosubsp. robertsii after R. H. Roberts, in recognition of his contribution to the study of Dactylorhiza and for all the generous help he has given to the Botanical Society of the British Isles as its Dactylorhiza referee for nearly 20 years. D. majalis subsp. occidentalis is not known to occur in Wales. D. majalis subsp. cambrensis differs from D. majalis subsp. occidentalis in having much longer, narrower and more rigid leaves, and much wider spurs (Roberts 1961b). D. x dinglensis nothosubsp. robertsii differs from D. X dinglensis nothosubsp. dinglensis in having a wider spur with a throat closer in width to D. majalis subsp. cambrensis. MEASUREMENTS In order to quantify the key differences between the hybrid and its parents, one flower from the mid portion of the inflorescence was removed from each of approximately 30 plants of D. majalis subsp. cambrensis, D. maculata subsp. ericetorum and their hybrid in 1989 from the Borth locality. The width of the spur throat was obtained by supporting each flower on a plasticine-type material which enabled the flower to be manoeuvred into position for measurement. A high quality steel rule was gently rested on the mouth of the throat and the width at this point measured through a X 10 magnifying glass. The labellum and spur were then excised from the flower as an integral unit and mounted underside uppermost on transparent glue on white card. The spur length and width were measured as soon as possible after mounting. Spur length measured was equal to character 2 in Fig. la of Bateman & Denholm (1989). Spur width was measured at the point where character 1 meets character 2 in the same figure. The results are tabulated in Table 1. : : ’ ; : A NEW DACTYLORCHID HYBRID 397 TABLE 1. SPUR DIMENSIONS FOR DACTYLORHIZA Xx DINGLENSIS NOTHOSUBSP. ROBERTSII AND ITS PARENTS (N = sample size, S.D. = standard deviation) Spur throat width Spur length Spur width (mm) (mm) (mm) N Mean =‘ S..D. N Mean _ i S..D.. N Mean S.D. D. majalis subsp. cambrensis 30 2:51 0:25 30 7:87 0-73 29 2-61 0-25 D. X dinglensis nothosubsp. robertsii 30 2°18 0-15 28 8-30 =: 0-66 27 2:08 0-35 D. maculata subsp. ericetorum PA | 1-81 0-16 30 6-52 0-90 30 1:19 0-22 DESCRIPTION Dactylorhiza < dinglensis (Wilmott) So6 nothosubsp. robertsii F. Horsman, nothosubsp. nov. Dactylorhiza majalis (Reichenb.) P. F. Hunt & Summerhayes subsp. cambrensis (R. H. Roberts) R. H. Roberts X D. maculata (L.) So6 subsp. ericetorum (E. F. Linton) P. F. Hunt & Summerhayes. Ho.otypus: Near Borth, Cards., v.c. 46, wet meadows, 17 June 1988, F. Horsman V88.58.1 (NMW). Calcar floris 8-3 mm longum, 2-1 mm latum, crassitudine D. majalis subsp. cambrensi propinquior, utriusque parentis calcare longior, nonnunquamque perspicue maculatum, eius faux 2-2 mm lata, latitudine D. majalis subsp. cambrensis propinquior. Spur of flower 8-3 mm long, 2-1 mm wide, closer in thickness to that of D. majalis subsp. cambrensis and longer than in either parent, sometimes clearly spotted, its throat 2-2 mm wide, closer to that of D. majalis subsp. cambrensis. Roots not examined. Average height c. 30 cm. Stem c. 5 mm in diameter at third lowest expanded leaf node, solid, suffused purplish from between first and second non-sheathing leaves upwards, faint at first but increasing in intensity, ridged from lowest non-sheathing leaf upwards. Basal sheath brown. Expanded sheathing leaves commonly 5, non-sheathing leaves commonly 3, 80% of expanded sheathing leaf nodes commonly in lowest 25% of stem. Expanded leaves — lowest c. 10 X c. 2 cm, becoming longer and narrower up the stem to the fourth (c. 13 X c. 1:25 cm) with the uppermost c. 10 X c. 1 cm; non-sheathing leaves — lowest c. 5-5 0-75 cm, becoming shorter and narrower up the stem to uppermost (c. 2:75 X c. 0-5 cm); lower floral bracts exceeding flowers, lowest c. 2:5 X 0-5 cm; all leaves widest between base and middle and becoming progressively more erect up the stem; leaves grey-green, the upper expanded leaves darker; lowest expanded leaf blunt and keeled, higher expanded leaves becoming acute and very shallowly hooded; lowest non- sheathing leaf tapering from c. one-third length from base; spots on upper surfaces of expanded leaves, maximum diameter on lower leaves c. 2-5 mm and on upper c. 1-5 mm, some of those on lower leaves transversely elongated or + oval, fewer transversely elongated on upper leaves, very high density on middle leaves, distributed towards leaf margins and only c. 10% in upper half of leaf, very few spots on uppermost expanded leaf; upper non-sheathing leaves faintly suffused with purple. Inflorescence c. 20% of total stem length, c. 2 as long as wide, c. 5 flowers/cm, fragrant, cylindrical. Flowers pale lilac; outer lateral perianth segments spreading to drooping, faintly marked with blotches and lines; median outer perianth segment unmarked. Forward edges of inner lateral perianth segments deeper in colour; labellum shape as in Fig. 1, basic pattern of markings — 2 adjacent loops in central part, elongated and narrowed towards mid-lobe, loops + continuous through spur throat and within the upper part of each loop is another elongated loop, but pattern of markings in central part of labellum exhibiting great variability; lateral lobes very slightly reflexed, with few rather faint marks; spur rather longer than ovary, tapering gradually for approximately one-third of its length from base and then more rapidly, with a gentle upward sweep to a point. 398 F. HORSMAN O0:-5cm Ficure 1. Labellum shapes in Dactylorhiza x dinglensis nothosubsp. robertsii and its parents. A. D. majalis subsp. cambrensis, B. D. X dinglensis nothosubsp. robertsii, C. D. maculata subsp. ericetorum. ACKNOWLEDGMENTS I should like to thank R. H. Roberts for confirming my determination of this new taxon, and for the very generous help and encouragement he has given me with my studies of this difficult genus. My thanks are also due to the v.c. Recorder, Arthur Chater, for all his help and encouragement with this paper, and the referee, for his guidance. I should like to thank Albert Henderson for his help with the Latin translation and Jo Nash for his excellent drawings. REFERENCES BATEMAN, R. M. & DENHOLM, I. (1983). A reappraisal of the British and Irish dactylorchids, 1. The tetraploid marsh-orchids. Watsonia 14: 347-376. BATEMAN, R. M. & DENHOLM, I. (1989). Morphometric procedure, taxonomic objectivity and marsh-orchid systematics. Watsonia 17: 449-455. Evus, R. G. (1983). Flowering plants of Wales. Cardiff. Hes_op-Harrison, J. (1953). Studies in Orchis L. Il. Orchis Traunsteineri Saut. in the British Isles. Watsonia 2: 371-391. Hes_op-Harrison, J. (1954). A synopsis of the dactylorchids of the British Isles. Ber. geobot. ForschInst. Riibel 1953: 53-82. A NEW DACTYLORCHID HYBRID 399 Roserts, R. H. (1961a). Studies on Welsh orchids, I. The variation of Dactylorchis purpurella (T. & T. A. Steph.) Vermeul. in North Wales. Watsonia 5: 23-36. Roserts, R. H. (1961b). Studies on Welsh orchids, II. The occurrence of Dactylorchis majalis (Reichb.) Vermeul. in Wales. Watsonia 5: 37-42. STEPHENSON, T. & STEPHENSON, T. A. (1921). Orchis latifolia in Britain. J. Bot., Lond. 59: 1-7. (Accepted December 1990) ; : ie me ta i tlhe Bere 7 nity. aT j ' _ > A Pia ReneS 4 ‘ «iG ; — Ar ps ¥ - et disisH) alnuan stole 1 all aero Gis U re ees ~ 1-2 town 208 A wtintind i abate: d ' D 7 ie ae i Watsonia, 18, 401-417 (1991) 40] Short Notes ANTHYLLIS VULNERARIA L. SUBSP. POLYPHYLLA (DC.) NYMAN, AN ALIEN KIDNEY-VETCH IN BRITAIN Anthyllis vulneraria (Kidney-Vetch) is a most variable species, both in Britain and Ireland and in Europe. Five subspecies have been recognized in these islands (Cullen 1976, 1986; Akeroyd 1988), three of them native: subsp. vulneraria, the most widespread variant, subsp. lapponica (Hyl.) Jalas, in Scotland and northern and western Ireland, mainly in the mountains and near the coast, and subsp. corbieri (Salmon & Travis) Cullen, a rare plant of coastal cliffs and sand-dunes in western Britain. Two other subspecies, subsp. carpatica (Pant.) Nyman and subsp. polyphylla (DC.) Nyman are adventive, both perhaps introduced in forage and amenity plantings. In recent years, subsp. carpatica (present in Britain and Ireland as var. pseudovulneraria (Sag.) Cullen) has expanded its range considerably in Britain, although apparently less so in Ireland (Scannell & Synnott 1987), mainly as a plant of grass and legume mixtures sown on new road verges, embankments and roundabouts. Subsp. polyphylla, on the other hand, is now a scarce and perhaps extinct plant in Britain — it has not been reported from Ireland — although it was collected or recorded in several localities from Dorset to Moray in the 1950s to the 1970s. There are no records after 1973. A. vulneraria L. subsp. polyphylla (DC.) Nyman, Consp. Fl. Eur. 164 (1878) Short-lived perennial. Stems 30-50 cm, erect, pubescent with appressed to patent hairs or more or less glabrous below, glabrous above. Basal and lower cauline leaves inequifoliate, with 3—5 pairs of lateral leaflets; upper cauline leaves subequifoliate, with 3—6(—7) pairs of leaflets. Inflorescences usually several on each stem, some lateral and sessile. Calyx rather narrow, concolorous (i.e. without a red tip), the hairs appressed. Corolla yellow. I have traced the following records: v.c. 9, Dorset. Briantspuddle, parish of Alfpuddle, roadside, 3.7.62, T. Woodisse 1, BM. v.c. 22, Berks. Maidenhead, by A423 at junction of M4 (GR 41/88.78), grass bank on chalk, 12.6.66, I. K. Ferguson 1378, TCD. v.c. 23, Oxon. Near Thame, chalk pit, 7.6.64, J. Rogerson. Intermediate between subsp. polyphylla and subsp. vulneraria, det. J. Cullen. B.R.C. card. v.c. 29, Cambs. Near Burwell (GR 52/575.653), bank of old railway line, 22.6.66, P. D. Sell 66/26, CGE (reported in Crompton & Wells 1990). v.c. 51, Flints. Melinden, 1 km south of Prestatyn (GR 33/169.801), limestone quarry, 29.5.61, R. K. Brummitt, J. Cullen & P. E. Gibbs 61.195, LIV; Trelogan (GR 33/12.80), rough grass, K. S. Kandall, det. J. Cullen. B.R.C. card. v.c. 85, Fife. Tentsmuir, Long Road (near Poles’ Camp), E. Crapper, 26.7.56, STA; near Kinshaldy, edge of Tentsmuir forest (GR 37/48.23), in hayfield, chiefly Lolium, 30.7.65, A. Angus 5176, STA. v.c. 89, E. Perth. c.1/2 mile south of Pitlochry (GR 27/95.57), main road embankment, 12.7.66, M.McC. Webster 10738, BM, RNG. v.c. 95, Moray. Near Randolph’s Leap, Dunphail, road-verge, 3.7.61, M.McC. Webster 5965, ABN, CGE, E, RNG; Elgin, golf course, car-park, 12.8.70, M.McC. Webster 13335, CGE; Whitemire (GR 28/98.54), bank by road, 18.6.73, M.McC. Webster, CGE. v.c. 96, Nairn. Near Kilravock, road verge, 1.8.62, M.McC. Webster, CGE; Auldearh (GR 28/9.5), station yard, 9.6.67, M.McC. Webster 11266, ABN, intermediate between subsp. polyphylla and subsp. vulneraria; Allanburn, Newton Hotel, 27.6.63, M.McC. Webster, CGE. Comparative descriptions of the five British subspecies are given in Akeroyd (1988), and they have been illustrated by Stewart (1987). Perhaps the most obvious feature of subsp. polyphylla is the 402 SHORT NOTES many pairs of lateral leaflets of the basal and lower cauline leaves. The most similar of the other subspecies in Britain is subsp. carpatica, which differs mainly in the inequifoliate cauline leaves and the more swollen calyx with somewhat spreading hairs. The tall, erect habit of both subspecies is distinctive in the field. The erect habit reflects the sometime use of subsp. carpatica and subsp. polyphylla as forage plants that can be cut readily during hay-making. This growth habit is in contrast to the often prostrate or weakly ascending habit of subsp. vu/neraria and subsp. /apponica in natural and semi- natural grassland. A similar pattern of differentiation exists between Medicago sativa L. subsp. sativa (Alfalfa or Lucerne) and subsp. falcata (L.) Arcangeli (native variant), and forage versus native variants of several other legumes, for example Lotus corniculatus L. (Bonnemaison & Jones 1986), Trifolium pratense L. (Williams 1927) and other clovers, and Onobrychis viciifolia Scop. (D. E. Coombe pers, comm.). About half the British records of A. vulneraria subsp. polyphylla are from road verges, which suggests that the plants were sown deliberately or accidentally as part of alegume and grass mixture. I have not observed subsp. polyphylla in the field in Britain, but have frequently observed subsp. carpatica. Typical associates include a tall, erect, fistulose-stemmed variant of T. hybridum L. and robust, erect variants of 7. pratense L. and T. repens L., which suggest the deliberate sowing of legumes derived from forage sources, probably the most readily available seed. The native distribution of A. vulneraria subsp. carpatica var. pseudovulneraria has perhaps been obscured by a history of cultivation (Cullen 1976), but the subspecies is generally central European in its distribution. Subsp. polyphylla is a plant of central and eastern Europe, and it therefore appears that central Europe may be the source of seed used in at least some roadside landscaping. The use of seed from other regions in road verge plantings has been documented in Finland (Suominen 1974), and the practice has attracted some attention from botanists in Britain in recent years, both from the evolutionary (Bonnemaison & Jones 1986) and the practical and ethical (Dony 1989 and pers. comm.) points of view. One of the records of A. vulneraria subsp. polyphylla from Tentsmuir, Fife (v.c. 85). is from a Lolium ley, which suggests either the deliberate sowing of subsp. polyphylla in a seed mixture or that the plant was present as a contaminant. The other Tentsmuir record, the earliest from Britain, is from near to a former Polish Army encampment, which generates a more colourful hypothesis. Many Polish soldiers, including cavalry units, were stationed in Fife during World War II (as a student in St Andrews in the 1970s I often heard Polish spoken in the town) and it is tempting to postulate that subsp. polyphylla was introduced to Tentsmuir with or in the horses’ fodder. Many adventive species in Finland were found to be associated with the activities of Russian soldiers and their horses during, and in the decade following, World War II, and several persisted for some years (Niemi 1969). The peak of records of subsp. polyphylla in the mid-1960s may reflect the use of this plant, or its presence as an impurity, in seed mixtures used on the new verges of an expanding and improved road network in Britain. The record from the bank of the recently constructed M4 motorway at Maidenhead may be significant in this respect. The lack of records from the early 1970s to the present, coupled with an increasing number of records of subsp. carpatica, may indicate that subsp. polyphylla has been replaced in commercial seed mixtures or that the source of these has shifted somewhat. It thus appears to join the list of adventive species that have been only temporarily established in Britain (Salisbury 1961). Nevertheless, the fact that it is a variant of a native species with which it may have hybridized (e.g. the record from Oxon), could mean that traces of its former presence are retained in the gene-pool of A. vulneraria in Britain. However, it is not clear to what extent such processes operate in natural populations. ACKNOWLEDGMENTS I am grateful to C. D. Preston for help in compiling records and for comments on an earlier draft of the manuscript, and to the late Dr J. G. Dony for discussion of the problems posed by the indiscriminate sowing of alien seed on road verges. SHORT NOTES 403 REFERENCES AKEROYD, J. R. (1988). Anthyllis vulneraria L. in Ricu, T. C. G. & Rico, M. D. B. Plant Crib, pp. 45-46. London. BONNEMAISON, F. & JonEs, D. A. (1986). Variation in alien Lotus corniculatus L. 1. Morphological differences between alien and native British plants. Heredity 56: 129-138. Crompton, G. & WELLS, D. (1990). Vascular plant records. Nature Cambs. 32: 77-80. CULLEN, J. (1976). The Anthyllis vulneraria complex: a resumé. Notes Roy. Bot. Gard. Edinb. 35: 1-38. CULLEN, J. (1986). Anthyllis vulneraria L. in the British Isles. Notes Roy. Bot. Gard. Edinb. 43: 277-281. Dony, J. G. in Briccs, M. (1989). Report of Annual General Meeting, 7 May 1988. Watsonia 17: 382-383. Niemi, A. (1969). Influence of the Soviet tenancy on the flora of the Porkala area. Acta Bot. Fennica 84: 1-52. SALISBURY, E. (1961). Weeds and aliens. London. SCANNELL, M. J. P. & Synnott, D. M. (1987). Census catalogue of the Irish flora, 2nd ed. Dublin. STEWART, O. M. (1987). Anthyllis vulneraria L. (Kidney vetch). B.S.B.I. News 46: 12-15. SUOMINEN, J. (1974). Maantieluiskanurmetuksista ja tulokaskasveista. Luonnen Tutkija 78: 12-18. WiuiaMs, R. D. (1927). Red clover investigations. Bulletin of the Welsh Plant Breeding Station H7. Shrewsbury. J. R. AKEROYD 24 The Street, Hindolveston, Norfolk, NR20 5BU BEVAN’S” BITTERCRESS:, (“THE)) SMALL, WHITE-FLOWERED VARIANT:> OF CARDAMINE X FRINGSII WIRTGEN ALSO PRESENT IN BRITAIN In April 1989, D.B. discovered a striking, white-flowered Cardamine (see Bevan 1990) in a flush at Bentley Priory, Middlesex. The plant was robust, uniform and approximately intermediate between C. flexuosa With. and C. pratensis L. (Table 1), with which it was growing. However, as the plant had small, white flowers rather than the larger, pale lilac flowers usually expected of the hybrid in Britain (Jones 1975), and as C. pratensis is very variable (Lovkvist 1956), a hybrid origin of ““Bevan’s Bittercress” could not be assumed. Further investigation of the Bentley Priory populations was required. TABLE 1. MORPHOLOGICAL CHARACTERS OF CARDAMINE TAXA FROM BENTLEY PRIORY Height Branching Flowering stems Upper surface of rosette leaves Petal colour length width Sepal length Mean pollen stainability + S.D. AND OF THE SYNTHESIZED HYBRID Measurements are ranges made on fresh material. C. flexuosa to 30 cm strong below, weak above sparsely hairy below hairy blade white, claw whitish 2:9-4-2 mm 1-1-1-5 mm 1-7—2:2 mm 60% 6% * Not assessed due to slug damage. Bevan’s Bittercress to 45 cm weak below, strong above glabrous glabrous or sparsely hairy when young blade white, claw greenish 5-2-7-3 mm 1-8—2-8 mm 2:2-2:6 mm 0% +0% C. pratensis to c. 30 cm simple below, weak above glabrous glabrous blade purple, pink or white, claw greenish 9-1-14-2 mm 4-5-8-0 mm 3-3—-4-6 mm 62% +9-6% Synthesized hybrid glabrous glabrous or sparsely hairy when young blade white, claw greenish 5-0-7-0 mm 2:0-4-2 mm 2-0-3-0 mm 0% +0% 404 SHORT NOTES FERTILITY Pollen viability was investigated using Alexander’s Stain. At least 100 pollen grains were counted from six flowers of each Cardamine (Table 1). About two-thirds of the pollen grains of C. flexuosa and C. pratensis took up stain, whilst those of Bevan’s Bittercress took up no stain at all. The results suggest that Bevan’s Bittercress is pollen-sterile. Note that pollen grains which take up stain are not necessarily viable. A better test of fertility is to assess seed production, and hence flowers of Bevan’s Bittercress were self- and cross-pollinated by hand. Bevan’s Bittercress stubbornly refused to set any good seed, even when cross-pollinated with C. pratensis. By contrast, seed developed freely in both white- and pink- flowered clones of C. pratensis when cross-pollinated. C. flexuosa is self-compatible and regularly sets good seed. & ~ CHROMOSOME COUNTS C. flexuosa has a chromosome number of 2n = 32, and C. pratensis has reported aneuploid chromosome numbers ranging from 2n = 16 to 2n = 96; the commonest races in Britain are 2n = c. 56 and 2n = c. 30-32 (Hussein 1955; Dale & Elkington 1974). It was hoped that a chromosome count of Bevan’s Bittercress might help clarify its relationships. Chromosomes were counted in root-tips of Cardamine taxa from Bentley Priory, at Leicester University. The chromosome numbers of C. pratensis were 2n = 30 (white-flowered clone) and 2n = 32 (pink-flowered clone); that of C. flexuosa was 2n = c. 30; and that of Bevan’s Bittercress 2n = 32. The results can neither confirm nor rule out a hybrid origin of Bevan’s Bittercress. HYBRIDIZATION EXPERIMENTS If Bevan’s Bittercress arose as a hybrid between local plants of C. pratensis and C. flexuosa, then it should be possible to synthesize it artificially using the same material. Flowers of C. flexuosa were cross-pollinated in bud after removal of the stamens (see Lovkvist 1956) using pollen from white- and pink-flowered clones of C. pratensis. Seed set was erratic and germination even more so. No seed was obtained from crosses with the pink-flowered clone of C. pratensis. Seed resulting from pollination with the white-flowered clone gave rise to 21 seedlings, 17 of which were C. flexuosa probably resulting from pollen con- tamination. Two of the remaining four plants flowered in 1990 and were an almost exact match for Bevan’s Bittercress morphologically and in pollen fertility (Table 1); they also failed to set seed when pollinated by hand. This is strong evidence in favour of a hybrid origin of Bevan’s Bittercress. CONCLUSIONS It is concluded that Bevan’s Bittercress is C. flexuosa X pratensis = C. X fringsii Wirtgen (= C. X haussknechtiana O. E. Schulz). The name C. X fringsii Wirtgen (1899) has priority over the more widely known name C. X haussknechtiana O. E. Schulz (1903) (D. H. Kent, pers. comm. 1991). Bevan’s Bittercress differs from variants of C. x fringsii previously reported in Britain (cf. Jones 1975), but is almost identical with material from the continent (Schulz 1903). Wirtgen describes the petal colour of C. x fringsii as lilac (D. H. Kent pers. comm.), whilst Schulz describes it as white. The two colour variants may be derived from different chromosome races of C. pratensis. Specimens have been placed in herb. T.C.G.R. and herb. D.B. ACKNOWLEDGMENTS We wish to thank J. P. Bailey and J. Wentworth, Leicester University for the chromosome counts, and D. H. Kent and E. Nic Lughadha for help with nomenclature. SHORT NOTES 405 REFERENCES BevaN, D. (1990). Cardamine flexuosa X pratensis at Bentley Priory, Middlesex. London Naturalist 69: Frontispiece. Date, A. & ELkincTon, T. T. (1974). Variation within Cardamine pratensis in England. Watsonia 10: 1-17. Hussein, F. (1955). Chromosome races in Cardamine pratensis in the British Isles. Watsonia 3: 170-174. Jones, B. M. G. (1975). Cardamine L., in Stace, C. A., ed. Hybridization and the flora of the British Isles. London. Lovxvist, B. (1956). The Cardamine pratensis complex. Outlines of its cytogenetics and taxonomy. Symb. Bot. Upsal. 14 (2): 5-131. ScHuLz, O. E. (1903). Monographie der Gattung Cardamine. Bot. Jahrb. 32: 548-549. D. BEVAN* & T. C. G. RICH *3 Queens Road, Bounds Green, London, NII 2QJ THE STATUS OF PULICARIA VULGARIS GAERTNER IN BRITAIN IN 1990 The Small Fleabane, Pulicaria vulgaris Gaertner, is an annual plant associated with traditionally grazed village greens. The historic distribution of the plant and its decline through this century, together with various ecological studies have been described by Hare (1990). The Small Fleabane is sufficiently rare to be listed in the British Red Data Book (Perring & Farrell 1983) and to receive special protection through inclusion on Schedule 8 of the Wildlife and Countryside Act 1981. The Small Fleabane in Britain is confined to southern England. The major strongholds lie in the south-west of the Hampshire basin with scattered outlying populations in the south of the Thames basin. The plant is present in the administrative counties of Dorset, Hampshire and Surrey but in the Watsonian vice-counties of S. Hants. (v.c. 11), N. Hants. (v.c. 12) and Surrey (v.c. 17). In 1990 botanists from the Flora Group of the Hampshire and Isle of Wight Naturalists’ Trust undertook a review of the status of the plant in Britain. All the areas where the plant has been known in recent years were visited, or were subject to reports from local botanists. At each site the number of plants was counted where practicable, or estimated where present in large numbers. At most sites the location and distribution of the plants were mapped both on 1:10000 O.S. base maps and on sketch maps. The notion of a ‘site’ is not entirely appropriate for this species due to the ruderal nature of the preferred habitat and the consequential erratic distribution of plants within a general area from year to year. For convenience sites were defined on an ad hoc basis around the ease of visiting and mapping populations. The British population of Small Fleabane in 1990 was estimated to be some 10,000 plants spread over ten sites. Seven of these ten sites lay within the borders of the New Forest. One site lay in the Avon Valley to the immediate west of the Forest. The remaining two sites lay in the Thames basin, one in Hampshire, the other in Surrey. The seven New Forest sites accounted for about 77% of the British plants. With a notable exception, the Forest sites which supported strong populations were located not on the Crown Lands managed by the Forestry Commission but on the verges and greens of adjacent commons to the north-east and west of the Forest. Additional small populations were found towards the south of the Forest. The Avon Valley site accounted for about 22% of the British plants. This population was counted as a single site although the plant was found over a wide area within a single farm. The majority of plants were found in a trackway with another colony in a smaller nearby track, and three small populations associated with broken ground within hay fields and a garden fruit cage. The populations of the Forest, combined with that of the Avon Valley, account for over 99% of the British plants. The smaller New Forest sites and the Thames basin sites supported a total of fewer than 60 plants in 1990. Amongst the Flora Group visiting various sites were botanists who had known individual populations over a number of years; we also had the benefit of being accompanied by Dr T. Hare. The general impression was that 1990 was a relatively good year for the species with strong 406 SHORT NOTES populations in most of the traditional strongholds. The prolonged drought was thought to be responsible for the drier sites supporting reduced populations but the relatively wet sites supported strong populations. The presence of two generations of plants was noted, one probably associated with germination in spring, the other with germination after the brief summer rains. The action of Forest ponies occasionally pulling up plants was observed. It was thought the ponies were seeking to eat the Small Fleabane but, finding it unpalatable, were discarding it. The absence of the Small Fleabane from one of its three recent Thames Basin localities was not considered significant. This year the site was not subject to its usual level of grazing and thus the open, trampled sward preferred by the plant had become overgrown by vigorous grasses. A return to grazing and subsequent open ground should revive the population. Within the New Forest the 1990 population may be compared to the 1985 population. In 1985 the Nature Conservancy Council co-ordinated a census of rare plants in the New Forest. The population of Small Fleabané in 1985 was estimated to be in excess of 106,000 plants. The dramatic decline in the total numbers to the present 7700 in the Forest can be attributed to the change in management of a common on the north-east of the Forest. In the early 1980s this common was subject to heavy use by commerical vehicles which resulted in gross disturbance to the vegetation and soils. Whilst the ruts associated with this use were advantageous to the Small Fleabane the disturbance was otherwise undesirable. By 1990 this use had ceased and many of the former ruts and hollows had disappeared. The population of Small Fleabane in this locality has fallen from an estimated 100,000 plants in 1985 to an estimated 1200 plants in 1990. Of the 1985 sites only one area was found to have changed to become unsuitable for the Small Fleabane. This was a length of verge that had been excavated to install underground telecommuni- cations equipment. The reinstatement of the verge was such as to destroy the seasonally inundated hollows preferred by Small Fleabane and to replace them by mounded spoil from adjacent ditches. Another of the 1985 sites has also been adversely modified. This is a small green at a road junction that has been planted with an ornamental tree. Given that the Small Fleabane is intolerant of shade it must be expected that this colany will decline as the tree matures. The traditional practices of turning livestock out onto Forest commons currently maintains suitable habitats for the species, yet the future of the commoning economy is uncertain (New Forest Review Group 1988). The acceptance of the threat to the surviving populations from the cessation of traditional grazings must form the basis of future conservation efforts (Whitten 1990). The Small Fleabane populations in the New Forest should not be assumed to be secure. ACKNOWLEDGMENTS The following provided or collated records for the 1990 survey: E. Brooks, F. Burford, G. D. Field, Dr T. Hare, Dr J. R. Moon, J. Ounstead, J. Smith and C. Chatters. Many thanks to the Hampshire Office of the N.C.C. for making available a summary of the 1985 Small Fleabane Survey. REFERENCES Hare, T. (1990). Lesser Fleabane- a plant of seasonal hollows. British Wildlife 2: 77-79. New Forest Review Group. (1988). Report of the New Forest Review Group. Lyndhurst. PERRING, F. H. & FARRELL, L. (1983). British Red Data Book: 1. Vascular Plants, 2nd ed. Lincoln. Wuitten, A. J. (1990). Recovery: A proposed programme for Britain’s protected species. Nature Conservancy Council, C.S.D. Report No. 1089. C. CHATTERS Hampshire and Isle of Wight Naturalists’ Trust, 71 The Hundred, Romsey, Hampshire, SO51I 8BZ CAREX X GAUDINIANA GUTHNICK - A RARITY? This hybrid, between Carex dioica and C. echinata, is known in a number of central and northern European countries but has only twice been reported from Britain and Ireland. The first discovery SHORT NOTES 407 was in 1942 by A. W. Stelfox in a bog near Louisburgh in West Mayo (specimens in BM and K); in 1970 a second site for it was found by I. R. Bonner in Denbighshire. In 1989 Mrs J. A. Green, the present recorder for Denbighshire, who had been trying to establish the locality for this sedge and assess the size of the population, sent specimens to the Natural History Museum for confirmation. These, however, proved inconclusive, and in 1990 Mrs Green invited me to join a party to survey the site, the North Wales Wildlife Trust’s Reserve at Gors Maen Llwyd. Both the reputed parents of C. x gaudiniana Guthnick were locally abundant in the Reserve, but at first we were confused by what came to be known as “‘funny echinata’’, a form of that species with the male florets at the base of the terminal spike extending into a recognisable male spike which might be as much as a centimetre long. I had never consciously noted this character before in C. echinata, and a later examination of the specimens in BM suggested that in Britain it is of rare occurrence. Yet the monographer of Carex, Kiikenthal (1909), evidently regarded it as the norm for C. echinata, for his description (of “Carex stellulata’’) includes the phrase ‘‘spiculae 3-5 gynaecan- drae, terminalis basi in partem CO longe attenuata”’. When we at last came upon C. X gaudiniana, whose very distinct appearance was checked against the precise, if somewhat convoluted, description in Ascherson & Graebner (1904) and perfectly accorded with it, there could be no doubt about our having found the right thing. The true plant forms tight clumps, some 15 cm across, of upright narrow leaves well surmounted by the rigidly erect flowering stems. These resemble tiny tridents, for the terminal male spike, a narrow cylinder up to 12 mm long, is flanked on either side by short, divergent female spikes. Another characteristic is that the male spike frequently has odd female florets embedded in its apex or at points below. Some dozen plants were spotted in a wet grassy area c. 10 m square and ringed by Calluna vulgaris near the middle of the western half of the bog. A pair and a single were seen at two spots in the eastern half. In all three places the hybrid was accompanied by both parents. It is hard to believe that it does not occur elsewhere in Britain and Ireland where C. dioica and C. echinata are found together, and a look-out should be kept for it. REFERENCES ASCHERSON, P. & GRAEBNER, P. (1904). Synopsis der mitteleuropdischen Flora, Band 2: 232-233. Leipzig. KUKENTHAL, G. (1909). Cyperaceae-Caricoideae, in ENGLER, H. G. A. Das Pflanzenreich, 38 (IV .20): 228-230. Leipzig. R. W. Davip 50 Highsett, Cambridge, CB2 I1NZ THE STATUS OF OENOTHERA CAMBRICA ROSTANSKI AND O. NOVAE-SCOTIAE GATES (ONAGRACEAE) In 1977 K. Rostanski described Oenothera cambrica from Pembrey in South Wales, United Kingdom. According to Rostanski (1982; Rostanski & Ellis 1979) this entity is confined to ““Wales, Jersey and southern England’’, occurring there in “‘sand-dunes, sandy sea-shores, railway banks and waste places’. Work during the past decade towards a thorough revision of Oenothera subsect. Oenothera has resulted in the reduction of O. cambrica to the complex synonymy of Oenothera biennis L. This detailed revision of the entire subsection Oenothera is in final preparation for publication in 1991 by W. Dietrich, W. L. Wagner and P. Raven. It supports and amplifies the philosophy that was described by Raven et al. (1979). This taxonomy provides a classification basically consistent with those applied to other groups of plants, and especially with the other sections of Oenothera. The comprehensive work underway for over the past decade has resulted ina classification in which five out-crossing, bivalent-forming species, and eight, essentially clonal, permanent translocation heterozygote species are recognized. In this treatment the permanent translocation heterozygote species are treated rather broadly. They behave essentially like other clonal species, although the mechanism, involving chromosomal translocations, balanced lethals, and self-pollination, is anomalous (for summary see Cleland 1972; Holsinger & Ellstrand 1984). 408 SHORT NOTES Each of these entities is comprised of a few to numerous true-breeding phenotypes that share common genetic and certain related phenotypic characteristics. Oenothera biennis is the most polymorphic and widespread of the eight permanent translocation heterozygote species of subsection Oenothera and consists of a large number of minor phenotypes, of which O. cambrica is one that originated from naturalized populations of O. biennis from North America. Rostanski (1985), after examining a specimen in BM annotated by R. R. Gates as Oenothera novae-scotiae Gates (Cult. at Regent’s Park, London, Gates 51/35), decided that his earlier described O. cambrica represented the same entity as this specimen. He therefore adopted O. novae-scotiae for these plants. Rostanski, however, did not examine the type of O. novae-scotiae from Nova Scotia, Canada (Gates s.n., UC—193440, holotype) indicated in the original publication by Gates (1918). I have examined the holotype and it does not represent O. cambrica (= O. biennis), but rather a quite distinct North American species with subterminal sepal-tips, O. parviflora, which is also naturalized in Europe. Oenothera parviflora is known in Great Britain from scattered collections in and near Glamorgan, Wales (Rostanski 1982). Thus, the names O. cambrica and O. novae-scotiae should not be placed together. Oenothera cambrica represents a phenotype of O. biennis with red pustulate-based hairs, a trait that is widely variable in the indigenous range of this species. Oenothera novae-scotiae represents O. parviflora. European workers who have not adopted the taxonomic philosophy discussed above for subsect. Oenothera should refer to the plants with small flowers, terminal sepal-tips and red pustulate-based hairs from Great Britain as Oenothera cambrica. REFERENCES CLELAND, R. (1972). Oenothera cytogenetics and evolution. London. Gates, R. R. (1918). A new evening primrose. Oenothera novae-scotiae. Trans. Nova Scotia lit. Soc. 14: 141- 145. Ho sinGer, K. E. & ELLSTRAND, N. C. (1984). The evolution and ecology of permanent translocation heterozygotes. Am. Nat. 124: 48-71. RAvEN, P. H., Dietrich, W. & StruBBE, W. (1979). An outline of the systematics of Oenothera subsect. Euoenothera (Onagraceae). Syst. Bot. 4: 242-252. RostTANSKI, K. (1977). Some new taxa in the genus Oenothera L. subgenus Oenothera. Part III. Fragm. flor. geobot. 23: 285-293. ROSTANSKI, K. (1982). The species of Oenothera L. in Britain. Watsonia 14: 1-34. RostaANskI, K. (1985). Zur Gliederung der Subsektion Oenothera (Sektion Oenothera, Oenothera L., Onagraceae). Feddes Repert. 96: 3-14. RosTANSKI, K. & ELuis, G. (1979). Evening Primroses (Oenothera L.) in Wales. Nature Wales 16: 238-249. W. DIETRICH Botanischer Garten der Universitat Diisseldorf, Universitatsstrasse 1, D-4000 Diisseldorf 1, Germany A NEW VARIANT OF OPHRYS APIFERA HUDSON IN BRITAIN On 20 June 1990, during a visit to Oxwich Burrows N.N.R. in Glamorgan, v.c. 41, I found a group of abnormal Bee Orchids (Ophrys apifera Hudson) in the dune slacks. This group included ten abnormal and four normal plants growing in an isolated slack. The abnormal plants were all robust, about 310 mm tall, with an inflorescence 200 mm in length containing about ten florets. Most were fully expanded, about two were in bud and a further one had failed to develop. There were three sheathing stem leaves, and the lowest bract measured 45 mm in length. All the perianth segments were entire, pink in colour with a strongly marked green central vein and two or more paramedian veins. The three inner perianth segments were slightly crinkled, but otherwise resembled the outer perianth segments (Fig. 1). Measurements of length and maximum width of the segments of a specimen floret were as follows: dorsal outer segment 16 X 8-5 mm, lateral outer segments 16-5 X 8mm and 16-5 X 7:5 mm; ventral inner segment 15-5 x 8-5 mm, lateral inner segments 16 * 7-5 mm. The column appeared normal, with well developed pollinia and evidence of self pollination. There is a high frequency of autogamy in Ophrys apifera, which enables SHORT NOTES 409 — Outer perianth segment NAOT NTE PEELS SS isi Cay Inner perianth segment Column Pollinia Outer perianth segment te Inner perianth a segment AETEAETIANE VEN SrA UE mere nee eer 0 10 mm 20 mm 30 mm Figure 1. Abnormal Ophrys apifera from Oxwich Burrows. morphological variants arising through mutation to persist, and sometimes predominate, in certain areas. In his paper on peloria and pseudopeloria, Bateman (1985) identified two categories of peloria: type (a) in which the lateral inner perianth segments are replaced by additional well differentiated labella; and type (b) in which the labellum is replaced by a third, undifferentiated, inner perianth segment. Clearly the peloria demonstrated by the Oxwich plants is not of either category, but comes close to that described by Horsman (1990) for Orchis latifolia L. Horsman proposed a new category, which he called type (c) peloria, where all the perianth segments are identical. The perianth segments of the Oxwich plants are not identical, and form an inner and outer series of three each. The inner segments are morphologically similar, but have no resemblance to either the labellum or the lateral inner perianth segments of normal O. apifera. The Oxwich plants probably merit a subdivision of Horsman’s type (c) peloria. The literature describes four abnormal kinds of O. apifera which have been given subspecific or varietal status. It would be more consistent to treat these as having equal rank, preferably using varietal names. Var. botteronii Chodat (syn. subsp. jurana Ruppert, var. friburgensis Freh.). The two lateral petals resemble pink sepals, with finely hairy margins. Labellar speculum bears two confluent yellow areas. First described in Britain in 1984 in Wiltshire (Laurence 1986). Illustrated by Duperrex (1961), Baumann & Kiinkele (1982), Davies et al. (1983) and Lang (1989). 410 SHORT NOTES Var. bicolor (Naegli) Nelson. The base of the labellum is greenish, lacking any pattern, rest of labellum dark brown. Recorded from Anglesey by Roberts (1985). Illustrated by Baumann & Kiinkele (1982) and Davies et al. (1983). Var. flavescens Rost (syn. var. chlorantha (Hegetschw.) Richter, var. immaculata Breb.). Sepals white, labellum lacking basal red brown pigmentation, appearing sage green. Widespread. Locally frequent in E. Suffolk and E. Sussex. Illustrated by Lang (1980), Baumann & Kiinkele (1982) and Davies et al. (1983). Var. trollii (Hegetschw.) Reichenbach. Labellum long and pointed, often barred brown and yellow. Long known from Gloucestershire, Dorset and several other localities. Illustrated by Summerhayes (1951), Lang (1980) and Baumann & Kiinkele (1982). In addition, a pseudopeloric variant, in which the labellum is replaced by a pink sepaloid segment, was first recorded in E. Sussex in 1924 by C. B. Tahourdin. Illustrated by Tahourdin (1924), Wolley- Dod (1937) and Lang (1980). None of the above corresponds to the Oxwich plants. However, F. Horsman (pers. comm) informs me that Camus & Camus (1929) recorded a similar variant from mainland Europe. ACKNOWLEDGMENTS I am grateful to lan Denholm and Frank Horsman for assistance in the preparation of this note. REFERENCES BATEMAN, R. M. (1985). Peloria and pseudopeloria in British orchids. Watsonia 15: 357-359. BAUMANN, H. & KUNKELE, S. (1982). Die wildwachsenden Orchideen Europas. Stuttgart. BAUMANN, H. & KUNKELE, S. (1986). Die Gattung Ophrys L. -— eine taxonomische Ubersicht. Mitt. BI. Arbeitskr. Heim. Orch. Baden-Wurtt. 18: 320-323. Camus, E. G. & Camus, A. (1929). Iconographie des Orchidées d’ Europe et du Bassin Méditerranéen 2, p. 325. Paris. Davies, P., Davies, J. & Hux ey, A. (1983). Wild orchids of Britain and Europe. London. Durerrex, A. (1961). Orchids of Europe. Blandford. Horsman, F. (1990). Peloria in Dactylorhiza. B.S.B.I. News 55: 16-18. Lana, D. C. (1980). Orchids of Britain. Oxford. LANG, D. C. (1989). A guide to the wild orchids of Great Britain and Ireland, 2nd ed. Oxford. LAURENCE, R. J. (1986). Ophrys apifera Hudson subsp. jurana Ruppert found in Britain. Watsonia 16: 177-8. Roserts, R. H. (1985). Some unusual orchid variants from Anglesey. Watsonia 15: 275-277. SUMMERHAYES, V. S. (1951). Wild orchids of Britain. London. TAHOURDIN, C. B. (1924). Some notes as to British Orchids. Croydon. Wo LLeEy-Dop, A. H. (1937). Flora of Sussex. Hastings. D. C. LANG 20 Ferrers Road, Lewes, Sussex, BN7 1PZ SALIX x SEMINIGRICANS A. & G. CAMUS: A WILLOW HYBRID NEW TO THE BRITISH ISLES On 19 July 1985, as part of a Salix course at Kindrogan Field Centre, Perthshire, a visit was made to the River Tay, near Meikleour. On the west bank of the river, immediately north of Kinclaven Castle, three willow bushes were seen on either side of the bridge that crosses the Tay at this point. Of the two growing north of the bridge one was fully grown, about 9 m tall, with a trunk about 15 cm diam. leaning out over the river; the second was smaller and less mature, and the third, to the south of the bridge, was clearly juvenile, erect, about 2 m tall, and had almost certainly arisen vegetatively SHORT NOTES 411 FicurE 1. Salix X seminigricans A. & G. Camus A: leaves; B: stipules; C: catkins; D: 9 flower; E: tip of flowering shoot. Scale bar = 1 cm (A, C, E) or 0-6 mm (B, D). from a broken-off branch carried downstream from the largest bush. The three were morphologi- cally indistinguishable. At first sight it was clear that the willow was a viminalis-hybrid: it had the acuminate leaves characteristic of such hybrids; but the relatively sparse indumentum on the underside of the leaves, the subglabrous, greenish shoots, and the small, poorly developed stipules ruled out the more usual viminalis-hybrids, namely S. X smithiana Willd. (S. cinerea L. X S. viminalis L.) and S. X sericans 412 SHORT NOTES Tausch ex Kerner (S. caprea L. X S. viminalis L.), and suggested that it was much more likely to be S. myrsinifolia Salisb. x S. viminalis L. (S. x seminigricans A. et G. Camus), a hybrid not previously recorded for the British Isles, and evidentiy rare on the Continent. Salix myrsinifolia was plentiful in the area, and S. viminalis was seen on the banks of the R. Isla nearby. No catkins were available at the time of the visit, and, without these, a final decision on the identity of the willow could not be made. Fortunately Mr Jeremy Heath (of Colchester and Essex Museum) had, at some personal risk, climbed out over the river, and had obtained good specimens from the largest bush, which, on his return to Colchester, he successfully rooted and cultivated, so that, by the spring of 1990, he was able to supply me with excellent flowering material. The catkins confirm my earlier, tentative identification. I am satisfied that the hybrid is S. myrsinifolia x S. viminalis, and since it may be found elsewhere in Scotland or northern England, a full description and illustration (Fig. 1) may be useful. Salix < seminigricans A. & G. Camus, Class. Saules d’Europe 2: 130 (1905). S. nigricans Sm. X S. viminalis L.; Schmalh. in Bot. Zeit. 33:574 (1875); Trautv., Increm. Fl. Phaenog. Ross. 3:698 (1883); Guerke, Pl. Europ. 2:27 (1897). S. myrsinifolia Salisb. X S. viminalis L. Type: U.S.S.R., near Leningrad, Schmalhausen (?KW, LE, P) Spreading shrub or small tree to about 9 m high; twigs glabrous, rather lustrous olive-brown; buds brown, glabrescent; current year’s shoots greenish, subglabrous. Leaves lanceolate, acuminate, 5— 15 x1-3-3 cm, rather glossy mid-green above (blackening slightly when dried), greyish and thinly adpressed-pubescent below, nervation distinctly impressed above, midrib and lateral nerves prominent below, margins indistinctly undulate-crisped, entire or subentire; petiole to 1-5 cm long; thinly pubescent; stipules developed only on vigorous growths, small, acute or acuminate, usually less than 3 mm long, submembranous, thinly pubescent, margins glandular. Female catkins appearing in March before the leaves, rather crowded towards the tips of the branches, sessile or subsessile, erect or spreading, cylindrical, blunt, 2-3 cm long, 0-7—-0-8 cm diam.; bracts sericeous, lanceolate, rather inconspicuous, usually less than 1 cm long; flowers crowded; catkin-scales ovate, obtuse or shortly acute, about 2 mm long, 0-8—1 mm wide, fuscous on the upper half, rather densely sericeous; nectary solitary, oblong, about 0-2 mm long, shorter than the pubescent pedicel; ovary narrowly ovoid, tapering to apex, about 2-2-5 mm long, 0-7 mm wide, closely adpressed-sericeous; style distinct, about 0-5 mm long; stigmas cleft almost to the base into 4 filiform arms about 1 mm long. Male flowers not seen. Scotland: Mid-Perth (v.c. 88), west bank of R. Tay immediately north and south of the bridge at Kinclaven Castle; leaves collected 19 July 1985, J. J. Heath & R. D. Meikle s.n.; flowers from plants cultivated at Colchester by J. J. Heath, comm. 6 March 1990. Europe: U.S.S.R., Leningrad area. R. D. MEIKLE Ranscombe Lodge, Wootton Courtenay, Minehead, Somerset, TA24 8RA THE TYPIFICATION OF RANUNCULUS BACHII WIRTG. In June 1844, the Prussian botanist Philipp Wirtgen discovered some unusual stands of Batrachian Ranunculi in the River Sayn and adjacent mill-races between Sayn and Isenburg, near Koblenz. He provisionally named these plants Ranunculus bachii (Wirtgen 1845), subsequently described them as Batrachium bachii (Wirtgen 1846a) and, later in the same year, provided the combination Ranunculus bachii (Wirtgen 1846b). Wirtgen’s herbarium is at Munster (MSTR) and he took his Dr. phil. at Bonn (BONN). We have borrowed Wirtgen material of this locality from both of these herbaria. It can be divided into two kinds, one of which has no floating leaves and small flowers, the other a different habit, floating leaves and larger flowers. The former fits Wirtgen’s original description and that of Webster (1990) and we select one of the sheets of this kind, “‘Ranunculus fluitans B bachii Wtg. In der Sayn bis Isenburg, lgt. com. Wirtgen’’ (MSTR) as the lectotype of Batrachium bachii Wirtgen. It 1s SHORT NOTES 413 morphologically intermediate between R. fluitans and R. trichophyllus. The other kind seems to be indistinguishable from R. x kelchoensis S. Webster (1990) (R. fluitans x peltatus). There is a possible problem with these hybrids. We do not know what range of morphology would be shown by the hybrid R. aquatilis x fluitans. Cook (1966, 1975) says that some forms of this hybrid are indistinguishable from, and comprise part of R. x bachii. On the other hand if they had floating leaves they might be indistinguishable from R. x kelchoensis. The sensible thing to do is to put all plants into these two taxa, using binomials, until such time as a R. aquatilis x fluitans can be morphologically recognised. REFERENCES Cook, C. D. K. (1966). A monographic study of Ranunculus subgenus Batrachium (DC.) A. Gray. Mitt. bot. StSamml., Miinch. 6: 57-237. Cook, C. D. K. (1970). Hybridization in the evolution of Batrachium. Taxon 19: 161-166. WEBSTER, S. D. (1990). Three natural hybrids in Ranunculus L. subgenus Batrachium (DC.) A. Gray. Watsonia 18: 139-146. WIRTGEN, P. [W.] (1845). Zweiter Nachtrag zur Flora preuss. Rheinlande. Verh. naturh. Ver. preuss. Rheinl. 2: 22-32. WIRTGEN, P. [W.] (1846a). Batrachium bachii Wirtg.: eine neue Pflanzenspecies. Verh. naturh. Ver. preuss. Rheinl. 3: 8-10. WIRTGEN, P. [W.] (1846b). Dritter Nachtrag zu dem Prodromus der Flora der preuss. Rheinlande. Verh. naturh. Ver. preuss. Rheinl. 3: 33-45. P. D. SELL* & S. D. WEBSTER *Botany School, Downing Street, Cambridge, CB2 3EA THREE NEW GRASS COMBINATIONS Three names of grasses used in New Flora of the British Isles (Stace 1991) have not been validly published before now. They are new combinations necessitated by the adoption of the generic circumscriptions in Genera Graminum (Clayton & Renvoize 1986). All three are casuals on tips and waste ground, the first two occasional, the last rare. 1. Helictotrichon neesii (Steudel) Stace, comb. nov. Amphibromus neesii Steudel, Syn. Fl. Glum. 1: 328 (1854) 2. Ceratochloa staminea (E. Desv.) Stace, comb. nov. Bromus stamineus E. Desv. in C. Gay, Fl. Chil. 6: 440 (1854) Bromus valdivianus Philippi in Linnaea 29: 101 (1857) 3. Leptochioa muelleri (Benth.) Stace, comb. nov. Diplachne muelleri Benth., Fl. Austral. 7: 619 (1878) REFERENCES Crayton, W. D. & RENvoIZzE, S. A. (1986). Genera Graminum. London. STACE, C. A. (1991). New Flora of the British Isles. London. C. A. STACE Department of Botany, University of Leicester, Leicester, LE] 7RH 414 SHORT NOTES FESTUCA ARUNDINACEA SCHREB. VAR. STRICTIOR (HACK.) K. RICHT. Hackel (1882) was the author of F. arundinacea Schreb. subvar. strictior which was raised to a variety by K. Richter. A taxon of the description given by Hackel (1882) can be recognised in the field but is seldom found as an isolated plant and is more often seen within large or small colonies of F. arundinacea. This taxon has not been widely accepted and few records exist. Hackel (1882) described subvar. strictior with “leaf blades narrower (3-5 mm broad) and shorter (than F. arundinacea), stiff becoming sub-convolute on drying . . . panicle shorter (up to 15 cm long), strict, scarcely nodding. . . spikelets as of F. arundinacea’. Having examined a considerable number of small specimens of F. arundinacea, I concluded that var. strictior could be more fully described as follows: height 50-90 cm, panicle length 6-15 cm, upper cauline blade (1-5—) 4-10 cm x 1-3-5 mm, spikelet 8-10 mm, lemma (5-5—) 6-8 mm. Druce (1931) collected the plant at Budleigh Salterton, S. Devon (v.c. 3) and Townsend (1953) collected it from ‘‘a marshy field’ near Barbers Bridge station, W. Gloucs. (v.c. 34); both collections were determined by Howarth. Kent (1975) recorded subvar. strictior from Hounslow Heath, Middlesex (v.c. 21) in the late “forties” in the company of C. E. Hubbard, who at the time treated it as a subvariety of no importance. This taxon was not listed in Hubbard (1954). Webster (1978) returned var. strictior to modern recognition with a site on the banks of the R. Spey at Dandaleith, Moray (v.c. 95), 1974, E and on newly sown verges at Cluanie Dam in Glen Moriston, Easterness (v.c. 96), 1974, E; both of these records were determined by C. E. Hubbard. In Cambs. (v.c. 29) F. arundinacea demonstrates its preference for calcareous clay. With Balsham as a centre, it is seen on the road verges to the south of Woodhall Farm, along the Balsham-Linton road. Large colonies are found at intervals along the B1052 in a north-easterly direction as far as Dullingham and east to Kirtling. The soil of this area is a chalky boulder clay of the Hanslope series over a chalky till. The soil survey of England & Wales (1983) describes it as slowly permeable calcareous clayey soil. Most of the soil samples would be pH 7-5-8-0. It is heavy land and its road verges have been subject to much over-riding of farm machinery and other wheeled traffic, which has compacted the soil to cause impeded drainage and allow for a high level of moisture retention. F. arundinacea is almost absent from the soils of the Middle Chalk in Cambs. (v.c. 29). In a survey of road verges from the south of Thriplow on the A505, east to Babraham, over The Gog and Magog Hills to the Beech Woods and down to Fulbourn, only three plants of var. strictior were seen in isolation, just west of Duxford and two plants near Pampisford Hall. Plants corresponding to var. strictior are found on the boulder clay in Cambridgeshire, in association with the typical variety. They are seldom seen in isolation, as quoted above at Duxford and Pampisford, but as a single culm within a clump or on the open margin of a colony in which it is frequently over-shadowed by the taller culms of typical F. arundinacea. The smaller variant is also found near the margin of road verges and a closer look sometimes reveals that the plant has been damaged by traffic running over the verge. Within any colony of F. arundinacea of 50-80 flowering stems, there are plants with short, narrow upper cauline blades of 4-10 cm long and short strict panicles, which may have a culm length of over 150 cm. Equally short panicles of 6-15 cm long are found with upper cauline blades of over 40 cm long. In large colonies, tall stem-growth inevitably over shadows weak stem-growth and var. strictior becomes stunted. It seems clear that a random selection of short-stemmed plants will not necessarily have short panicles and/or short upper cauline blades. It is considered that Hackel’s taxonomic recognition of this variant of Festuca arundinacea is inappropriate and that those small plants with short, strict panicles and short upper cauline blades should only be considered as environmentally-induced variants. ACKNOWLEDGMENT [ thank Dick David for his help in the translation of Hackel’s description of subvar. strictior. REFERENCES Druce, G. C. (1931). Plant record. Rep. botl. Soc. Exch. Club Br. Isl. 9: 678. HACKEL, E. (1882). Monographia Festucarum Europaearum, p. 154. Berlin. SHORT NOTES 415 Hussarp, C. E. (1954). Grasses, p. 145. Harmondsworth. Kent, D. H. (1975). Historical Flora of Middlesex, p. 566. London. Sor, SURVEY OF ENGLAND & WALES (1983). Sheet 4, eastern England. TOWNSEND, C. C. (1953). Plant record. Watsonia 2: 356. Wesster, M.McC. (1978). Flora of Moray, Nairn and East Inverness-shire, p. 147. Aberdeen. P...O. TRist Glovers, 28 High Street, Balsham, Cambridge, CBI 6DJ CONTRIBUTIONS TO A CYTOLOGICAL CATALOGUE OF THE BRITISH AND IRISH FLORA, 1 A cytological catalogue of the British and Irish flora is being prepared (funded in part by a grant from the Welch Bequest), which will collate details of all published and unpublished chromosome counts made on indigenous material of native species (Gornall 1989). As only about 62% of the flora have properly documented counts, much original work still needs to be done; this is being conducted at three centres: the Botany Department, Leicester University; the Scottish Crop Research Institute, Invergowrie; and Trinity College, Dublin. The present paper is the first in a new series which will report the results of our studies as they progress. The chromosome counts of 61 species and their provenances are presented here. Figures in parentheses after the locality details indicate the number of plants counted. All counts were made from squashes of root-tips; supernumerary chromosomes are designated by the suffix ‘S’. Voucher specimens have been placed in LTR. Ammophila arenaria (L.) Link, 2n = 28: S. Lanes., v.c. 59, Ainsdale, 34/2.1 (3). Anthriscus sylvestris (L.) Hoffm., 2n = 16: W. Cornwall, v.c. 1, Buryas Bridge, 10/44.29 (1). Arctium minus Bernh., 2n = 36: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (3). Atropa bella-donna L., 2n = 72: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (3). Blackstonia perfoliata (L.) Hudson, 2n = 40: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (3). Callitriche brutia Petagna, 2n = 28: Caerns., v.c. 49, W. of Rhydlios, Methlem Farm, 23/174.301 (1). C. platycarpa Kutz., 2n = 20: Caerns., v.c. 49, N. of Llangwnnadl, Traeth Penllech, 23/20.34 (3). Campanula trachelium L., 2n = 34: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (3). Carduus acanthoides L., 2n = 16: Derbys., v.c. 57, Milldale, 43/14.55 (1). Centaurea nigra L., 2n = 44: W. Norfolk, v.c. 28, Fouldon Common, 52/761.997 (3). Chamaemelum nobile (L.) All., 2n = 18: Caerns., v.c. 49, SW. of Abersoch, S. of Mynydd Cilan, 23/292.240 (1). Clematis vitalba L., 2n = 16: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (1). Cynodon dactylon (L.) Pers., 2n = 36: S. Somerset, v.c. 5, Warren Point, Minehead, 21/986.463 (1); N. Somerset, v.c. 6, Weston-super-Mare, 31/317.603 (1); S. Essex, v.c. 18, Southend, Shoebury- ness, 51/448.852 (1). (No vouchers.) Digitalis purpurea L., 2n = 56: Caerns., v.c. 49, Llaniestyn, near Penbodlas, 23/281.337 (2). Erodium cicutarium (L.) L’Herit. subsp. cicutarium, 2n = 40: Derbys., v.c. 57, Dovedale, 43/14.52 CE Euonymus europaeus L., 2n = 32: Surrey, v.c. 17, Old Coulsdon, hillside above Old Lodge Lane, 51/31.59 (1); Caerns., v.c. 49, near Aberdaron, by the Afon Saint, 23/166.265 (1). Filipendula ulmaria (L.) Maxim., 2n = 16: Derbys., v.c. 57, Dovedale, 43/14.51 (1). Fragaria vesca L., 2n = 14: Dorset, v.c. 9, Wareham, Washer’s Pit, 30/86.94 (3). Frankenia laevis L., 2n = 30: W. Sussex, v.c. 13, East Wittering, 40/80.97 (1). 416 SHORT NOTES Galium aparine L., 2n = 66: Caerns., v.c. 49, E. of Sarn, 23/24.32 (1). G. mollugo L., 2n = 44: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (3). Geum urbanum L., 2n = 42: Dorset, v.c. 9, Wareham, Washer’s Pit, 30/86.94 (1). Hypericum hirsutum L., 2n = 18: Dorset, v.c. 9, Wareham, Washer’s Pit, 30/86.94 (2); Derbys., v.c. 57, Dovedale, 43/14.51 (1). H. humifusum L., 2n = 16: W. Cornwall, v.c. 1, Goonhilly 10/73.21 (1). Inula conyza DC., 2n = 32: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (3). Iris foetidissima L., 2n = 40: Dorset, v.c. 9, Wareham, Washer’s Pit, 30/86.94 (1). Lepidium heterophyllum Bentham, 2n = 16: Caerns., v.c. 49, near Aberdaron, overlooking Afon Saint, 23/16.26 (1). Leymus arenarius (L.) Hochst., 2n = 56 + 3S: S. Lancs., v.c. 59, Ainsdale, 34/2.1 (3). Lycopus europaeus L., 2n = 22: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (3). Lysimachia nemorum L., 2n = 18: Caerns., v.c. 49, Nanhoron valley, S. of Inkermann Bridge, 23/ 28.32 (3). Medicago arabica (L.) Hudson, 2n = 16: Caerns., v.c. 49, Aberdaron, 23/172.264 (1). Mercurialis perennis L., 2n = 64: Dorset, v.c. 9, Wareham, Washer’s Pit, 30/86.94 (1); Derbys., v.c. 57, Dovedale, 43/14.51 (1). Montia perfoliata (Willd.) Howell, 2n = 36: Leics., v.c. 55, Leicester, university campus, 43/59.04 (1). Myosotis sylvatica Hoffm., 2n = 18: Caerns., v.c. 49, Llaniestyn, near Penbodlas, 23/281.337 (1). Ononis spinosa L., 2n = 30: W. Norfolk, v.c. 28, Foulden Common, 52/761.997 (3). Origanum vulgare L., 2n = 30: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (2); Derbys., v.c. 57, Dovedale, 43/14.51 (1). Pastinaca sativa L., 2n = 22: N. Hants., v.c. 12, near Petersfield, W. of Shoulder of Mutton, above Steep, 41/73.26 (2). Pilularia globulifera L., 2n = 26: Caerns., v.c. 49, S. of Abersoch, Mynydd Cilan, 23/291.241 (1). Plantago coronopus L., 2n = 10: W. Cornwall, v.c. 1, near St Just, 10/39.31 (1). P. lanceolata L., 2n = 12: Derbys., v.c. 57, Dovedale, 43/14.52 (3). P. major L. subsp. major, 2n = 12: Derbys., v.c. 57, Milldale, 43/14.54 (3). P. major subsp. intermedia (DC.) Arcangeli, 2n = 12: Leics., v.c. 55, Cropston Reservoir, 43/54.10 (3). Potentilla sterilis (L.) Garcke, 2n = 28: Dorset, v.c. 9, Wareham, Washer’s Pit, 30/86.94 (1); Derbys., v.c. 57, Dovedale, 43/14.51 (1). Primula veris L., 2n = 22: Derbys., v.c. 57, Milldale, 43/14.55 (3). P. vulgaris Huds., 2n = 22: Caerns., v.c. 49, Llanbedrog, W. of Penarwel, 23/328.324 (1). Ranunculus repens L., 2n = 32: Caerns., v.c. 49, Llaniestyn, near Penbodlas, Llys Hyfryd, 23/28.33 CL), Rumex obtusifolius L., 2n = 40: Derbys., v.c. 57, Milldale, 43/14.54 (1). Sanguisorba minor Scop., 2n = 28: Derbys., v.c. 57, Dovedale, 43/14.52 (1). Sanguisorba officinalis L., 2n = 56: Leics., v.c. 55, Broughton Astley, 42/53.94 (3). Saxifraga hypnoides L., 2n = 52: Derbys., v.c. 57, Milldale, 43/14.55 (3). Sherardia arvensis L., 2n = 22: Derbys., v.c. 57, Dovedale, 43/14.52 (1). Sibthorpia europaea L., 2n = 18: S. Devon, v.c. 3, Dartmoor, north of Holne, Newbridge, 20/ 712.709 (1). Silaum silaus (L.) Schinz & Thell., 2n = 22 + 0-1S: W. Norfolk, v.c. 28, Foulden Common, 52/ 761.997 (3). Teucrium scorodonia L., 2n = 34: Derbys., v.c. 57, Dovedale, 43/14.51 (1). Tragopogon porrifolius L., 2n = 12: Co. Durham, v.c. 66, Crimdon to Blackhall, 45/47.38 (1). Umbilicus rupestris (Salisb.) Dandy, 2n = 48: Caerns., v.c. 49, Llaniestyn, near Penbodlas, 23/28.33 (ft). Urtica dioica L., 2n = 52: Derbys., v.c. 57, Dovedale, 43/14.52 (1). Verbascum thapsus L., 2n = 36: Dorset, v.c. 9, Wareham, Washer’s Pit, 30/86.94 (1). Veronica beccabunga L., 2n = 18: Derbys., v.c. 57, Milldale, River Dove, 43/14.55 (3). SHORT NOTES 417 Viburnum lantana L., 2n = 18: Surrey, v.c. 17, Old Coulsdon, hillside above Old Lodge Lane, 51/ 31.59 (1). Viola reichenbachiana Jord. ex Boreau, 2n = 20: Dorset, v.c. 9, Wareham, Washer’s Pit, 30/86.94 (iy REFERENCE GornaLlL, R. J. (1989). A proposed cytological catalogue of the British and Irish flora. B.S.B.I. News 51: 34. J. E. WENTWortTH, J. P. BAILEY & R. J. GORNALL Botany Department, The University, Leicester, LE] 7RH ] overs co? afte TUE —s abe ua; rods sbi See 10 re Hug. 1b riage» al 7 ; hs ny ee dea edit 9? ’ ‘ wa ‘ - 1B . > a J : | ! pe os HY. laonifecsun Ltn JW Corawalt "a var. oy sib he te radi F Shon ay) pe .<¢Q >a" > nh foal ve Reid ¢ ot? pees ee ne — id “fea ity . j —— | > ’ ne ) 7 - = ’ tha Vile ny . SRE: aa ATES LF Sass DP OOo reed ‘hie tebecenie Diels =< a bo. aS OUST SD Senne) Fa 4 attentions VEE x a. yy, Me : is a de Le fomtieido Ww of Sk , ’ i for Watsonia, 18, 419-436 (1991) 419 Plant Records Records for publication must be submitted to the appropriate Vice-county Recorder (see B.S.B.I. Year Book for 1991), and not the Editors. The record must normally be of species, hybrids or subspecies of native or naturalized alien plants belonging to one or more of the following categories: 1st or 2nd v.c. record; Ist post-1930 v.c. record; only extant v.c. locality, or 2nd such locality; a record of an extension of range by more than 100 km. Such records will also be accepted for the major islands in v.cc. 102-104 and 110. Only 1st records can normally be accepted for Rubus, Hieracium and hybrids. Records for subdivisions of vice-counties will not be treated separately; they must therefore be records for the vice-county as a whole. Records of Taraxacum are now being dealt with separately, by Dr A. J. Richards, and will be published at a later date. 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)). All records are field records unless otherwise stated. With the exception of collectors’ initials, herbarium abbreviations are those used in British and Irish herbaria by D. H. Kent & D. E. Allen (1984). Records from the following vice-counties are included in the text below: 1, 2, 4-6, 11, 12, 21, 24-29, 33-35, 38, 39, 41-43, 45-47, 49-54, 61, 62, 64, 67-70, 72, 73, 75-77, 79-81, 83, 87-90, 92-99, 103, 105, 109, 110, H30. The following signs are used: * before the record: to indicate a new vice-county record. + before the species number: to indicate that the plant is not a native species of the British Isles. + before 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. DIPHASIASTRUM ALPINUM (L.) Holub 81, Berwicks.: Stony slope, A68 road, Soutra, GR 36/47.56. D. G. Long, 1990. Only extant locality. 3/1. ISOETES LACUSTRIS L. 72, Dumfriess.: Shallow water, edge of Loch Skeen, GR 36/17.15. O. M. Stewart, 1989. Only extant locality. 4/5 X 6. EQUISETUM FLUVIATILE L. X E. PALUSTRE L. *73, Kirkcudbrights.: Marshy rough grassland, Southerness, GR 25/96.54. O. M. Stewart, 1990, E, det. C. N. Page. 7/1. HYMENOPHYLLUM TUNBRIGENSE (L.) Sm. 67, S. Northumb.: N.W.-facing sandstone crags, Roughside Moor, GR 35/73.83. G. A. & M. Swan, 1990, herb. G.A.S. 2nd record. +12/1. ONOCLEA SENSIBILIS L. 12, N. Hants.: Bank of R. Rother, Liss Forest, GR 41/77.28. J. Ockenden, 1990. 2nd record. *99, Dunbarton: Under Salix spp. on sandy river bank, R. Leven near Strathleven, GR 26/39.77. K. Futter, 1990, E. 15/5 qua. ASPLENIUM TRICHOMANES subsp. QUADRIVALENS D. E. MEYER *109, Caithness: Rocky outcrop, Yarrows, GR 39/30.42. J. K. Butler, 1980, E, det. C. R. Fraser-Jenkins. 16/1. CETERACH OFFICINARUM DC. *62, N. E. Yorks.: Mortar of brick wall, United Reform Church, Loftus, GR 45/71.18. A. M. Fryday, 1988, det. R. L. Gulliver. 21/2 cam. DRYOPTERIS AFFINIS subsp. CAMBRENSIS Fraser-Jenkins 41, Glam.: Ditch by wood, Cyfarthfa Castle, GR 32/04.07. G. Hutchinson, 1990, NMW. 2nd record. 21/2 aff. x 1. DRYOPTERIS AFFINIS (Lowe) Fraser-Jenkins subsp. AFFINIS X D. FILIX-MAS (L.) Schott *11, S. Hants.: Mixed plantation, Spearywell Wood, GR 41/31.27. R. M. Veall, 1989, herb. R.M.V., det. C. R. Fraser-Jenkins. 21/8. DRYOPTERIS AEMULA (Aiton) O. Kuntze 11, S. Hants.: Streambank in deciduous wood, Howe Copse, Roydon Woods, GR 40/32.99. A. E. Bolton, 1990, herb. R.P. Bowman. 2nd record. 24/4. GYMNOCARPIUM DRYOPTERIS (L.) Newman 28, W. Norfolk: Damp patch in coniferous wood, Weeting, GR 52/80.89. M. Keene, 1990, NWH. 420 PLANT RECORDS 25/1/cam. POLYPODIUM CAMBRICUM L. 11, S. Hants.: Mossy branches of oak, Lyndhurst Hill, Emery Down, GR 41/28.08. D. Winsland, 1989, herb. R.P. Bowman, det. R. H. Roberts. 2nd record. 25/1/int. POLYPODIUM INTERJECTUM Shivas *77, Lanarks.: By R. Clyde S. of Lanark, GR 26/ 8.4. A. McG. Stirling et al., 1989. 25/1/int. X vul. POLYPODIUM INTERJECTUM Shivas X P. VULGARE L. *77, Lanarks.: Rock face, Meikle Dripps, GR 26/58.55. P. Macpherson, 1985, herb. P.M., det. A. McG. Stirling. 40/1/1 X +var. ACONITUM NAPELLUS L. X A. VARIEGATUM L. *49, Caerns.: Ditch, Nanhoron, GR 23/2.3. J. Roberts, 1988, det. C. A. Stace. 2nd Welsh record. +43/bla. ANEMONE BLANDA Schott & Kotschy *69, Westmorland: Roadside verge between Kentmere and Staveley, GR 35/45.01. C. E. Wild, 1990, LANC. 46/19. RANUNCULUS FLUITANS Lam. “*4, N. Devon: R. Torridge, Weare Gifford, GR 21/ 47.21. C. D. Preston & N. F. Stewart, 1989, CGE, det. S. D. Webster. 1st confirmed record. 46/22c. RANUNCULUS PENICILLATUS subsp. PSEUDOFLUITANS (Syme) S. Webster *95, Moray: Auchernack Burn 2 km S. of Grantown, GR 38/02.24. J. Edelsten, 1990, E, det. S. D. Webster (as var. pseudofluitans). 46/23. RANUNCULUS BAUDOTII Godron 29, Cambs.: Fen ditch, Long Grove, Ouse Fen, GR 52/ 37.72. A. O. Chater, F. H. Perring & S. M. Walters, 1955, CGE, det. S. D. Webster. 1st record since 1859. 48/1. MyOSURUS MINIMUS L. 54, N. Lincs.: Bare ground near cattle trough, Lea Marshes, GR 43/81.86. A. J. Knibb & R. Nickerson, 1989, det. I. Weston. Ist record since c. 1900. +51/off. PAEONIA OFFICINALIS L. *69, Westmorland: Roadside, Fell Yeat, Casterton, GR 34/ 63.79. Woodland by R. Kent, Sedgwick, GR 34/50.86. Both C.E. Wild, 1990, LANC. 1st and 2nd records. 57/2. CERATOPHYLLUM SUBMERSUM L. *33, E. Gloucs.: Flooded, disused brick pit, Sandhurst, GR 32/81.23. S. H. Bishop & R. J. Cooper, 1987. Small field pond, Pamington, GR 32/93.31. J. W. Somerset, 1990. Ist and 2nd records. 58/3. PAPAVER LECOQi Lamotte 28, W. Norfolk: Spring barley field, Feltwell, GR 53/70.90. R. Tofts, 1989. 2nd record. 35, Mons.: Spoilheaps by R. Usk, Newport, GR 31/32.87. T. G. & U. T. Evans, 1988. 2nd record. 58/5. PAPAVER ARGEMONE L. 50, Denbs.: Weedy arable field near Fenns Moss, GR 33/50.38. J. A. Green, 1990. 2nd extant locality. +58/ori. PAPAVER ORIENTALE L. *69, Westmorland: Roadside north of Oxenholme, GR 34/ 53.91. C. E. Wild, 1988. Between Tebay and Orton, GR 35/62.06. C. E. Wild, 1989. 1st and 2nd records. 66/2. FUMARIA CAPREOLATA L. 11, S. Hants.: Roadside verge, Southern Lane, New Milton, GR 40/23.94. J. N. Le Rossignol, 1989, herb. R. P. Bowman, det. M. G. Daker. Only extant locality. 66/4. FUMARIA BASTARDII Boreau *73, Kircudbrights.: By field, Torrs Moss, GR 25/77.61. O. M. Stewart, 1988, E. Field edge, West Maryfield, New Abbey, GR 25/96.66. O. M. Stewart, 1990, E. Ist and 2nd records. +68/1. ERUCASTRUM GALLICUM (Willd.) O. E. Schultz 69, Westmorland: Roadside south of Black Esset Plantation, Killington, GR 34/58.87. C. E. Wild, 1989, LANC. 2nd record. +71/1. HIRSCHFELDIA INCANA (L.) Lagréze-Fossat 46, Cards.: Sandy roadside bank, Penyrer- gyd, Gwbert, GR 22/16.48. A. O. Chater, 1990. 2nd record. *77, Lanarks.: Waste ground, Springboig, Glasgow, GR 26/65.64. J. H. Dickson & K. Watson, 1989, GL. Rough grassland, Cambuslang, GR 26/63.59. K. Watson, 1989, GL. Ist and 2nd records. PLANT RECORDS 421 72/2. DIPLOTAXIS TENUIFOLIA (L.) DC. 38, Warks.: Side of railway, Birmingham, GR 42/ 08.87. J. W. Partridge, 1989, WAR, det. T. C. G. Rich. 2nd record. +72/3. DipPLOTAXIS ERUCOIDES (L.) DC. *28, W. Norfolk: Edge of conifer plantation, Hockwold-cum-Wilton, GR 52/69.86. T. C. G. Rich, 1990, LANC. +74/3. RAPHANUS SATIVUS L. 50, Denbs.: Shingly riverbank, Tregeiriog, GR 33/17.33. J. A. Green, 1989. 2nd record. +76/2. RAPISTRUM RUGOSUM (L.) All. 12, N. Hants.: Disturbed sandy wasteland, Aldershot, GR 41/85.50. T. Dove & A. R. G. Mundell, 1990, herb. A. Brewis. 2nd record. +81/1. CARDARIA DRABA (L.) Desv. *93, N. Aberdeen: Waste ground, Burnhaven, GR 48/ 13.44. D.Welch, 1990, ABD. +84/2. THLASPI ALLIACEUM L. *11, S. Hants.: Edge of arable field. Lower Clockhouse Farm, Winkton, GR 40/16.96. F. A. Woodhead, 1987, herb. F.A.W., det. R. P. Bowman. 89/1. SUBULARIA AQUATICA L. 72, Dumfriess.: Shallow water over peaty substrate, Loch Skeen, GR 36/17.16. R. W. M. Corner, 1989. Only extant locality. +90/2. BUNIAS ORIENTALIS L. 77, Lanarks.: Waste ground, Low Blantyre, GR 26/69.57. K. Watson, 1990, GL. 2nd post-1930 record. 94/4. DRABA MURALIS L. +*67,S. Northumb.: Recently cleared ground, Stantonfence, GR 45/ 13.89. G. A. Swan, 1990, herb. G.A.S. +96/1. ARMORACIA RUSTICANA P. Gaertner, B. Meyer & Scherb. *73, Kirkcudbrights.: Shore, Nun Mill, GR 25/65.48. O. M. Stewart, 1990. +98/3. BARBAREA INTERMEDIA Boreau 94, Banffs.: Roadside verge, Mulben, GR 38/35.50. J. Edelsten, 1988, E, det. T. C. G. Rich. 2nd record. 102/3. RORIPPA SYLVESTRIS (L.) Besser 94, Banffs.: River shingle, R. Spey, GR 38/34.64. J. Edelsten, 1990, E, det. T. C. G. Rich. 2nd record. +102/6. RoRIPPA AUSTRIACA (Crantz) Besser *77, Lanarks.: Waste ground, Ruchill, Glasgow, GR 26/58.68. J. H. Dickson, 1988, GL. Still present in abundance, 1990. 102/+6 X 3. RORIPPA AUSTRIACA (Crantz) Besser X R. SYLveEsTRIS (L.) Besser *77, Lanarks.: Cadder, Glasgow, GR 26/61.72. J. H. Dickson, 1989, GL, det. T. C. G. Rich. 7112/3. RESEDA ALBA L. 11, S. Hants.: Bare gravelly soil, West Cliff, Hythe, GR 41/41.08. A. P. Holliwell, 1990. Only extant locality. 113/6. VIOLA CANINA L. 81, Berwicks.: Sea braes, Burnmouth, GR 36/95.61. F. G. Hardy, 1990, det. A. J. Richards. 2nd localized record. 113/11. VioLA LUTEA Hudson *5,S. Somerset: Rough pasture, Rugg’s Hill, Brompton Regis, GR 21/98.31. G. Crouch & I. P. Green, 1990, herb. R. G. B. Roe, det. R. D. Meikle. 1st confirmed record and southerly extension of range, previous Somerset records having been doubted. Roadside knoll, Oldrey, GR 21/90.37. G. Crouch & I. P. Green, 1990. 2nd extant locality. 115/1. HYPERICUM ANDROSAEMUM L. +*77, Lanarks.: Pathside, Govan, GR 26/54.64. P. Macpherson, 1988. Waste ground, Hallside, GR 26/66.60. P. Macpherson, 1989. Both herb. P.M. Ist and 2nd records. 115/6a. HYPERICUM MACULATUM Crantz subsp. MACULATUM *69, Westmorland: Roadside, A685 road, Low Borrowbridge, GR 35/60.01. C. E. Wild, 1989, LANC, det. N. K. B. Robson. 115/6b X 5. HYPERICUM MACULATUM subsp. OBTUSIUSCULUM (Tourlet) Hayek < H. PERFORATUM fe: *4, N. Devon: Quarry spoil, Cove, GR 21/95.20. L. J. Margetts, 1990, det. N. K. B. Robson. *11, S. Hants.: Hedge, Ampfield, GR 41/40.23. R. M. Veall, 1990, herb. R. P. Bowman, det. N. K. B. Robson. 422 PLANT RECORDS 122/1. ELATINE HEXANDRA (Lapierre) DC. 47, Monts.: Lake margin, Llyn Ebyr, GR 22/9.8. I. C. Trueman & M. Wainwright, 1990. 2nd record. 123/12. SILENE NOCTIFLORA L. *35, Mons.: Garden weed, Chepstow, GR 31/52.93. T. G. Evans, 1982. Still present in 1990. 123/coe. SILENE COELI-ROSA (L.) Godron *35, Mons.: Gravelly island, Afon Llwyd east of Sebastopol, GR 31/30.97. T. D. Pollard & T. G. Evans, 1987, herb. T.G.E., det. E. J. Clement. 127/8. DIANTHUS DELTOIDES L. *75, Ayrs.: Castle Hill, Ardrossan, GR 26/2.4. Gardiner, no date, BM. 1st and only record. 131/2. CERASTIUM ARVENSE L. 70, Cumberland: Old sand quarry, East Brownrigg, GR 35/ 52.37. R. W. M.Corner, 1990, LANC. 1st post-1930 record. 133/3. STELLARIA PALLIDA (Dumort.) Piré 80, Roxburghs.: Bare soil, Peniel Heugh near Ancrum, GR 36/65.25. R. W. M. Corner, 1990, herb. R.W.M.C. 2nd record. 133/4. STELLARIA NEGLECTA Weihe *109, Caithness: Arable field, St. Mary’s Kirk, Forss, GR 39/0.7. M. McCallum Webster, 1968, E, det. A. O. Chater. 1st confirmed record. 141/1 mac. ARENARIA SERPYLLIFOLIA subsp. MACROCARPA (Lloyd) F. Perring & Sell *46, Cards.: Dune slack and grey dune, Ynys-las, GR 22/60.93. B.S.B.I. meeting, 1990, det. A. O. Chater. 153/bli. AMARANTHUS BLITOIDES S. Watson 25, E. Suffolk: Fallow field, Ramsholt, GR 62/ 31.42. E. M. Hyde, 1989, IPS, det. E. J. Clement. 2nd record. 154/9. CHENOPODIUM FICIFOLIUM Sm. 38, Warks.: Manure heap, Ettington Park, GR 42/ 24.47. H. A. Roberts, 1988, det. P. J. Copson. 2nd record. 154/14. CHENOPODIUM RUBRUM L. *43, Rads.: Roadside tip south of Upper Pitts Farm, Stanage, GR 32/31.70. R. G. Woods, 1990. Mud on lake margin, Llanbwchllyn, GR 32/12.46. D. R. Druit & I. D. Soane, 1990, det. R. G. Woods. 1st and 2nd records. 154/bot. CHENOPODIUM BoTrRYSs L. *6, N. Somerset: Roadside verge, Edington, GR 31/39.40. I. P. Green, 1990, herb. J.M.M., det. J. M. Mullin. 156/lon. ATRIPLEX LONGIPES Drejer 68, Cheviot: Under Phragmites at edge of salt-marsh, Alnmouth, GR 46/24.09. G. A. Swan, 1988, herb. G.A.S., det. J. R. Akeroyd. 2nd record. 156/lon. X 3. ATRIPLEX LONGIPES Drejer X A. PROSTRATA Boucher ex DC. *67,S. Northumb.: Under Phragmites at edge of pond near mouth of Blakemoor Burn, GR 45/28.94. G. A. Swan, 1988, herb. G. A.S., det. J. R. Akeroyd. *68, Cheviot: Under Scirpus maritimus at edge of salt-marsh near Beal Point, GR 46/08.42. G. A. Swan, 1987, det. P. M. Taschereau. 156/pra. ATRIPLEX PRAECOX Hiilphers *68, Cheviot: Sand above high-tide mark, mouth of Brunton Burn, GR 46/22.27. G. A. Swan, 1988. Sand above high-tide mark, Budle Bay near Links End, GR 46/14.35. G. A. & M. Swan, 1988. Both herb. G. A. S., det. J. R. Akeroyd. 1st and 2nd records. 160/2. SALICORNIA DOLICHOSTACHYA Moss *75, Ayrs.: Sandy/muddy shore, Hunterston, GR 26/19.52. A. McG. Stirling & A. Rutherford, 1990, E. 165/1. ALTHAEA OFFICINALIS L. +*69, Westmorland: Ditch, Rampside, Barrow-in-Furness, GR 34/23.66. P. Burton, 1989, LANC. 168/9. GERANIUM PYRENAICUM Burm. fil. *72, Dumfriess.: Roadside bank, Craigend Cottage, Georgetown, GR 35/01.74. M.E.R. Martin, 1990. 168/12. GERANIUM ROTUNDIFOLIUM L. +*50, Denbs.: Dry sandy ground, Marford Quarry, GR 33/35.56. P. Williams, 1990. 168/16b. GERANIUM ROBERTIANUM subsp. CELTICUM Ostenf. +*99, Dunbarton: Rosneath Caravan Park, GR 26/27.82. A. McSweeney, 1986, det. P. F. Yeo (as cv. Celtic White). PLANT RECORDS 423 4170/2. OXALIS CORNICULATA L. 73, Kirkcudbrights.: Edge of burn, Creetown, GR 25/47.58. O. M. Stewart, 1988. 2nd record. 4171/2. IMPATIENS CAPENSIS Meerb. *25, E. Suffolk: Bank of R. Waveney, Beccles, GR 62/ 41.90. T. Abrehart, 1989. +171/3. IMPATIENS PARVIFLORA DC. *42, Brecs.: Edge of plantation, Llangoed, GR 32/11.40. M. & C. Porter, 1990. Bank of R. Wye near Llangoed, GR 32/12.39. L. Bolingbroke, 1990. 1st and 2nd records. 94, Banffs.: Woodland by R. Spey 4 km N. E. of Rothes, GR 38/30.50. J. Edelsten, 1990, E. 2nd record. ’ +RHUS TYPHINA L. 41, Glam.: Cutting above cycle trackway, Lower Swansea Valley, GR 21/ 66.93. G. Hutchinson, 1989, NMW. 2nd record. Spreading by suckers. +182/2. PARTHENOCISSUS QUINQUEFOLIA (L.) Planchon *73, Kirkcudbrights.: Edge of wood, Balmae, GR 25/68.44. O. M. Stewart, 1990. 185/1 tin. GENISTA TINCTORIA L. subsp. TINCTORIA 45, Pembs.: Wet heath, Sageston Mountain Common, GR 22/06.03. S. B. Evans, 1990. 2nd extant locality. +CYTISUS STRIATUS (Hill) Rothm. *25, E. Suffolk: Purdis Heath Golf Course, GR 62/21.43. N. Kerr, 1990, herb. E. & M. Hyde, det. E. J. Clement. 190/6. MEDICAGO ARABICA (L.) Hudson +51, Flints.: St Asaph, GR 33/0.7. B. Gale, 1988. 2nd post-1930 record. +192/7. TRIFOLIUM INCARNATUM L. *52, Anglesey: Soil in car park, Traeth Lligwy, GR 23/ 49.87. B.S.B.I. meeting, 1990. 192/16. TRIFOLIUM SUFFOCATUM L. 6, N. Somerset: Short turf, Berrow Golf Course, GR 31/ 29.51. R. S. Cropper, 1990. 2nd record. 4198/1. ROBINIA PSEUDACACIA L. 70, Cumberland: Field edge, Lazonby, GR 35/55.39. R. W. M. Corner, 1989, LANC. 2nd record. 202/1. ORNITHOPUS PERPUSILLUS L. +*98, Main Argyll: Sides of forestry roads near Ardbreck- nish, GR 27/07.20. B. H. Thompson, 1989, herb. B.H.T., det. C. D. Preston. 207/2. LATHYRUS NISSOLIA L. +64, Mid-W. Yorks.: Waste ground, Hellifield, GR 34/85.56. J. Allinson, 1990. 2nd record. +69, Westmorland: Roadside verge between Brough and Brough Sowerby, GR 35/79.13. P. M. Atkins, 1990. 2nd record. +*99, Dunbarton: Waste ground, Dalmonach, GR 26/39.80. K. Futter, 1990. +209/2. SPIRAEA DOUGLASII Hooker subsp. DOUGLASII *77, Lanarks.: Old railway line, Meadowside, Glasgow, GR 26/55.66. P. Macpherson, 1989, herb. P.M., det. A. J. Silverside. +209/med. SPIRAEA MEDIA Franz Schmidt *69, Westmorland: Limestone quarry above Row, Whitbarrow, GR 34/45.88. C. E. Wild, 1989, LANC, det. A. J. Silverside. + ARUNCUS DIOICcUs (Walter) Fernald 69, Westmorland: Roadside between Newby Bridge and Bowland Bridge, GR 34/40.88. G. Halliday, 1988. 2nd record. 211/2. RUBUS SAXATILIS L. 47, Monts.: Ravine near Llanbrynmair, GR 22/9.9. G. Morton, 1989. 2nd record. *211/8. RUBUS SPECTABILIS Pursh *79, Selkirks.: Woodland, Glenkinnon Burn, Peel, GR 36/ 42.34. M. E. Braithwaite, 1990. +211/10. RuBus LOGANoBACccus L. H. Bailey *79, Selkirks.: Hedge south of Bridgelands, Selkirk, GR 36/47.30. R. W. M. Corner, 1989, herb. R.W.M.C., det. A. Newton. 211/11/2. Rusus scissus W. C. R. Watson *12, N. Hants.: Heathy ride, Sandford Woods, GR 41/55.59. D. E. Allen & A. O. Chater, 1982, BM, det. A. Newton. 211/11/27. RUBUS TUBERCULATUS Bab. *79, Selkirks.: Roadside, Selkirk, GR 36/47.29. R. W. M. Corner, 1989, herb. R.W.M.C., det. A. Newton. 424 PLANT RECORDS 211/11/32. RUBUS BRITANNICUS Rogers *26, W. Suffolk: Tuddenham Heath, GR 52/74.72. A. L. Bull, 1971, herb. A.L.B., det. A. Newton. 2211/11/44. RuUBUs NITIDIFORMIS Sudre *11, S. Hants.: Scrub margin, Southampton Common, GR 41/41.17. D. E. Allen, 1988, BM, det. A. Newton. 211/11/126. RUBUS ERRABUNDUS W. C. R. Watson [H30, Co. Cavan: Delete record in Edees & Newton (1988), Brambles of the British Isles, p. 274. Specimen in DBN is R. mucronulatus Boreau, det. D. E. Allen.] 211/11/129. RUBUS ULMIFOLIUs Schott *67, S. Northumb.: Grassy bank, Wallsend Dene, GR 45/31.66. D. N. Mitchell & A. Coles, 1990, herb. G. A. Swan, det. A. Newton. +211/11/139. Rupus procerus P. J. Mueller *77, Lanarks.: Lane-side, Hyndland, GR 26/ 55.67. A. McG. Stirling, 1989. 211/11/173. RUBUS BORAEANUS Genev. [11, S. Hants.: Delete record in Edees & Newton (1988), Brambles of the British Isles, p. 129. Provenance of specimen in SLBI, the basis of the record, is doubtful, fide D. E. Allen.] 211/11/240. RuBUs watsont W. H. Mills *25, E. Suffolk: Priestley Wood, Barking, GR 62/ 08.53. A. L. Bull, 1981, herb. A. L. B., det. A. Newton. 211/11/297. RuBUS RADULOIDES (Rogers) Sudre *12, N. Hants.: Open birchwood, Combe Wood, Lickenholt, GR 41/35.59. D. E. Allen & A. Newton, 1989, BM. 211/11/aqu. RuBUS AQUARUM Newton & Porter *42, Brecs.: Hedgerow west of Llanwrtyd Wells, GR 22/87.45. M. Porter, 1988, NMW, det. A. Newton. 211/11/bil. RuBUs BILOENSIS Newton & Porter *42, Brecs.: Lane-bank near Llanfillo, GR 32/ 11.33. M. Porter, 1988, NMW, det. A. Newton. +211/11/ele. RUBUS ELEGANTISPINOSUS (A. Schum.) H. E. Weber *26, W. Suffolk: Mildenhall Woods, GR 52/72.75. A. L. Bull, 1989, herb. A.L.B., det A. Newton. *79, Selkirks.: Old railway, Galashiels, GR 36/49.74. R. W. M. Corner, 1989, herb. R.W.M.C., det. G. Ballantyne & A. McG. Stirling. 211/11/ery. RUBUS ERYTHROPS Edees & Newton *11, S. Hants.: Margin of Red Wood, West Meon, GR 41/66.25. D. E. Allen, 1984, herb. D.E.A., det A. Newton. 211/11/gal. RUBUS GALLoFUsSCUS Newton & Porter *41, Glam.: Mynydd Bach near Lower Cwm-twrch, GR 22/76.09. M. Porter, 1988, NMW, det. A. Newton. *42, Brecs.: Gorsey bank, Penrhos near Ystradgynlais, GR 22/80.11. M. Porter, 1988, NMW, det A. Newton. 211/11/inf. RUBUS INFORMIFOLIUS Edees *12, N. Hants.: Margin of Chawton Park Wood, GR 41/67.36. D. E. Allen, 1975, herb. D.E.A., det. A. Newton. 211/11/mer. RUBUS MERLINI Newton & Porter *43, Rads.: Thickets, Allt Goch, GR 22/9.7. A. Ley, 1892, NMW, det. A. Newton. *44, Carms.: Roadside bank, Cilyewm, GR 22/75.39. T. A. W. Davis, 1973, NMW, det. M. Porter. 211/11/nor. RuBus NorvIcENSIs A. L. Bull & Edees *12, N. Hants.: Shady edge of car park, Winchester Station, GR 41/47.30. D. E. Allen, 1982, BM. 211/11/pyd. RUBUS PYDARENSIS Rilstone *4, N. Devon: Trackside, Challice’s Plantation, Eggesford, GR 21/68.09. W. H. Tucker, 1990, det. A. Newton. 211/11/rob. RusBus rosin (W. C. R. Watson) Newton *42, Brecs.: Wooded bank of R. Pyrddin, Pont-nedd-fechan, GR 22/89.09. M. Porter, 1990, herb. M.P., det. A. Newton. 211/11/sur. RUBUS SURREJANUS W. C. Barton & Riddelsd. *21, Middlesex: Shrubbery weed, Regent’s Park, GR 51/28.33. D. E. Allen, 1989, BM. PLANT RECORDS 425 211/11/tam. RUBUS TAMARENSIS Newton *11, S. Hants.: Heath margin, Chilworth Common, GR 41/41.17. D. E. Allen, 1974, herb. D. E. A., det. A. Newton. *12, N. Hants.: Hedge, Harewood Forest, GR 41/40.44. D. E. Allen, 1987, BM, det. A. Newton. 211/11/tav. RuBUS TAVENSIS Newton & Porter *42, Brecs.: Near Coed Wern Plemys, Ystradgynlais, GR 22/78.09. M. Porter, 1988, NMW, det. A. Newton. 211/11/tro. RuBUS TROIENSIS Newton *42, Brecs.: Hedge and wood, Llangynidr, GR 32/ 16.19. M. Porter, 1989, herb. M.P., det. A. Newton. 211/11/vig. RuBus vicursi Rilstone *4, N. Devon: Track through scrub, Kigbeare, Oke- hampton, GR 20/53.95. W. H. Tucker, 1990, det. A. Newton. 212/6. POTENTILLA ARGENTEA L. +*5, S. Somerset: Waste ground, Wembdon, GR 31/27.39. pee a Po Green. 1990. 72, Dumfriess.: Old sand pit, Dormont Grange, GR 35/11.74. M. E. R. Martin, 1986. Only extant locality. *212/12. POTENTILLA CRANTzII (Crantz) G. Beck ex Fritsch 72, Dumfriess.: Rocky ledge, Blackhope, GR 36/13.11. R. W. M. Corner & H. Lang, 1989. Only extant locality. 212/15. POTENTILLA REPTANS L. 95, Moray: River shingle, Boat o’Brig, GR 38/31.50. J. Edelsten, 1988, E, det. B. Harold. 2nd record. 220/3/9. ALCHEMILLA GLOMERULANS Buser *68, Cheviot: Cliff by stream in deep ravine, Bellyside Burn, GR 36/90.22. G. A. Swan, 1990, herb. G.A.S., det. S. M. Walters. 220/3/11. ALCHEMILLA WICHURAE (Buser) Stéfansson 72, Dumfriess.: Upland damp moor- land, Midlaw Burn area, GR 36/1.1. R. W. M. Corner, 1989. 2nd post-1930 record. +220/3/12. ALCHEMILLA MOLLIS (Buser) Rothm. *35, Mons.: Rough grassy roadside, Trede- gar, GR 32/13.09. T. G. Evans & J. Killick, 1988. *49, Caerns.: Waste ground near Llyn Chwythlyn, north-east of Llanrwst, GR 23/83.65. R. Lewis, 1990, NMW. *69, Westmorland: Elliscales Quarry, Dalton-in-Furness, GR 34/22.74. P. Burton, 1988. By pond, Pinfold Bridge, Raisbeck, GR 35/64.07. C. E. Wild, 1989, LANC. 1st and 2nd records. *73, Kirkcudbrights.: Roadside verge by Rosebank Bridge, GR 25/87.76. O. M. Stewart, 1990. +225/5. RosA RUGOSA Thunb. *46, Cards.: Top of shingle beach below Penyrangor, Aberystwyth, GR 22/58.80. A. O. Chater, 1990, NMW. *77, Lanarks.: Waste ground, Linthouse, GR 26/54.66. P. Macpherson, 1985, herb. P.M. Still present in 1990. 225/8 X 10. ROSA CANINA L. X R. OBTUSIFOLIA Desv. *41, Glam.: Near Llanedeyrn, Cardiff, GR 31/2.8. A. E. Wade, 1969, NMW, det. A. L. Primavesi. lst Welsh record. 225/8 X 14. ROSA CANINA L. X R. RUBIGINOSA L. *41, Glam.: Dutton’s Quarry, Rhiwbina Hill, Cardiff, GR 31/14.83. G. Hutchinson, 1989, NMW, det. A. L. Primavesi. 225/8 X 15. ROSA CANINA L. X R. MICRANTHA Borrer *11, S. Hants.: Grassland near tidal creek, Ports Creek, Hilsea, GR 41/65.04. R. P. Bowman, 1990, herb. R.P.B., det. A. L. Primavesi. 225/9 cae. ROSA CAESIA Sm. subsp. CAESIA *41, Glam.: Abernant, GR 32/01.03. J. W. Davies & A. E. Wade, 1973, NMW, det. A. L. Primavesi. *75, Ayrs.: Near Martnaham Loch, GR 26/ 40.17. A. McG. Stirling, 1988. 225/9 gla. ROSA CAESIA subsp. GLAUCA (Nyman) Graham & Primavesi 75, Ayrs.: Nobel Works, Ardeer, GR 26/27.40. A. McG. Stirling, 1988. 2nd record. 225/9 cae. X 8. ROSA CAESIA Sm. X R. CANINA L. *12, N. Hants.: Winnall Moors N.R., GR 41/48.30. J. Fryer, 1988, herb. A. L. Primavesi, det. A.L.P. 225/9 gla. X 8. ROSA CAESIA subsp. GLAUCA (Nyman) Graham & Primavesi X R. CANINA i. *41, Glam.: Abernant, GR 32/01.03. J. W. Davies, A. Newton & A. E. Wade, 1973, NMW, det. A. L. Primavesi. 1st Welsh record. 225/11 X 8. ROSA TOMENTOSA Sm. X R. CANINA L. *41, Glam.: Hedgerow north of Mountain Ash, GR 32/0.0. A. E. Wade, 1975, NMW, det. A. L. Primavesi. 426 PLANT RECORDS 225/12. ROSA SHERARDII Davies 11, S. Hants.: Roadside, Goatspen Plain, GR 41/23.01. Scrub, Burley Beacon, GR 41/20.02. Both R. P. Bowman, 1990, herb. R.P.B., det. A. L. Primavesi. Only extant localities. *75, Ayrs.: Ayr, GR 26/3.2. P. Ewing, 1889, GL, det. A. McG. Stirling. 1st confirmed record. 225/14. ROSA RUBIGINOSA L. *35, Mons.: Gritty bank near B4248 road, Blaenavon, GR 32/ 25.09. T. G. Evans & R. Fraser, 1988, herb. T.G.E., det. G. G. Graham. *79, Selkirks.: Old railway, Galashiels, GR 36/47.37. R. W. M. Corner, 1989, SUN, det. G. G. Graham. +226/ser. PRUNUS SEROTINA Ehrh. *46, Cards.: Sessile oakwood, Allt Fawr, Llanwnnen, GR 22/54.47. A. O. Chater, 1990, NMW. Abundant and locally dominant. +227/3. COTONEASTER HORIZONTALIS Decne *81, Berwicks.: Natural rock outcrop, Cockburn Mill, GR 36/77.58. M. E. Braithwaite, 1990. +227/5. COTONEASTER FRIGIDUS Wall. ex Lindley *35, Mons.: Rough grassland, Cwmtillery, GR 32/22.04. R. Fraser, 1988. +227/reh. COTONEASTER REHDERI Pojarkova *70, Cumberland: Clints Quarry, Egremont, GR 35/00.12. A. Dudman, 1989, LANC, det. J. Fryer. +227/ste. COTONEASTER STERNIANUS Boiss. *70, Cumberland: Old railway line, Harrington, GR 25/99.25. A. Dudman, 1989, LANC, det. J. Fryer. 232/7. SORBUS TORMINALIS (L.) Crantz 41, Glam.: Roadside near Earlswood Roundabout, Crymlyn Burrows, GR 21/72.93. C. R. Hipkin, 1990. 2nd post-1930 record. 237/1. CRASSULA TILLAEA Lester-Garland 12, N. Hants.: Old railway track, Woolmer Forest, GR 41/79.31. F. Rose, 1990. 2nd extant locality. +237/hel. CRASSULA HELMSII (T. Kirk) Cockayne *28, W. Norfolk: Pond on common, Tottenhill Row, GR 53/63.10. C. D. Preston, 1987, CGE. Pond, Tottenhill, GR 53/67.89. H. Williamson, 1990. 1st and 2nd records. 239/6. SAXIFRAGA HIRSUTA L. +*77, Lanarks.: Wood, Thorntonhall, GR 26/58.54. P. Mac- pherson, 1989, herb. P.M. +DARMERA PELTATA (Torrey) Voss *77, Lanarks.: Well naturalized in damp wood east of Busby, GR 26/58.55. P. Macpherson, 1989, herb. P.M., det. C. A. Stace. 247/2. DROSERA ANGLICA Hudson 72, Dumfriess.: Wet flushes near road below Auchenstroan Craig, GR 25/69.91. A. McG. Stirling, 1960. Wet pools at centre of bog, Lochar Moss, Longbridgemuir, GR 35/04.69. A. J. Kerr & L. V. Fleming, 1990. 1st and 2nd post-1930 records. 252/1. HIPPOPHAE RHAMNOIDES L. +*73, Kirkcudbrights.: Shingle bank, island south of St Mary’s Isle, GR 25/66.48. O. M. Stewart, 1990. Ist record of self-sown plants. +256/1. OENOTHERA BIENNIS L. 75, Ayrs.: Waste ground, Troon, GR 26/23.42. R. Mackech- nie, 1952. 2nd record. 4256/2. OENOTHERA ERYTHROSEPALA Borbas *43, Rads.: Ddol Road Industrial Estate, Llandrindod Wells, GR 32/06.62. R. G. Woods, 1990. +259/aqu. MyRIOPHYLLUM AQUATICUM (Velloso) Verdcourt *25, E. Suffolk: Pool, Landguard Common, GR 62/28.31. A. Copping, 1985. Still present in 1988. *39, Staffs.: Canal 1 km west of Aldridge, GR 43/04.00. J. P. Martin, 1990, herb. B.R. Fowler. *67,S. Northumb.: Mud at edge of small pond near Brenkley, GR 45/22.75. G. A. Swan, 1990, herb. G.A.S. 262/3. CALLITRICHE OBTUSANGULA Le Gall 73, Kirkcudbrights.: Glenamour Loch, GR 25/ 44.67. O. M. Stewart, 1989, E. 2nd record. 262/5. CALLITRICHE HERMAPHRODITICA L. *38, Warks.: Longmoor Pool, Sutton Park, GR 42/ 09.95. J. W. Partridge, 1990, WAR, det. J. C. Bowra & P. J. Copson. PLANT RECORDS 427 262/bru. CALLITRICHE BRUTIA Petagna 46, Cards.: Mud on side of dried-up pond, Glanrheidol, Capel Bangor, GR 22/66.79. A. O. Chater, 1990, NMW, det. C. D. Preston & N. F. Stewart. 2nd record, Ist since 1894. 4265/2. CORNUS SERICEA L. *73, Kirkcudbrights.: Wood near Gelston, GR 25/77.58. O. M. Stewart, 1988. 4271/1. ASTRANTIA MAJOR L. *26, W. Suffolk: Edge of flooded gravel pits, Lackford, GR 52/ 79.70. Suffolk Wildlife Trust recording team, 1987. 274/1. ANTHRISCUS CAUCALIS Bieb. *47, Monts.: Short grass and hedge bottom, Crew Green, GR 33/31.15. S. Stafford, 1977. Still present in 1990. +283/5. BUPLEURUM FALCATUM L. *35, Mons.: Garden weed, Chepstow, GR 31/52.93. T. G. Evans, 1980, herb. T.G.E. Still present in 1990. 2nd Welsh record. *64, Mid-W. Yorks.: Basic grassland on steep riverside bank, Bolton Abbey, GR 44/07.55. F. Draper & A. Gramshaw, 1990, det. J. Duncan. 291/1. CARUM VERTICILLATUM (L.) Koch *35, Mons.: Wet flush in woodland glade near Blaina, GR 32/2.0. J. Wohlgemuth, 1990, herb T. G. Evans. *96, Easterness: Amongst Molinia, east end of Loch Insh, GR 28/83.05. D. Wood, 1990. 291/2. CARUM CARVI L. 38, Warks.: Meadow, Bidford-on-Avon, GR 42/10.50. D. Boyce, 1990, WAR, det. J. C. Bowra. 2nd record, Ist since 1860. 300/1. OENANTHE SILAIFOLIA Bieb. *54, N. Lincs.: Water meadows, Lea Marshes near Gainsborough, GR 43/81.86. R. Nickerson, 1990, det. T. Dixon & I. Weston. +319/4. EUPHORBIA CORALLIOIDES L. 35, Mons.: Garden weed, Chepstow, GR 31/52.93. T. G. Evans, 1980. Still present in 1990. 2nd record. 319/416 X +15. EUPHORBIA CYPARISSIAS L. X E. ESULA L. *26, W. Suffolk: Forest ride, King’s Forest, GR 52/83.73. E. M. Hyde, 1990, IPS, det. A. Radcliffe-Smith. | +319/17 rob. EUPHORBIA AMYGDALOIDES subsp. ROBBIAE (Turrill) Stace *39, Staffs.: Side of pasture west of Upper Penn, Wolverhampton, GR 32/88.95. B. R. Fowler, 1990, herb. B.R.F., det. A. Radcliffe-Smith. +319/wal. EUPHORBIA WALDSTEINII (Sojak) R.-Sm. *77, Lanarks.: Grass bank above railway, Govan, GR 26/54.64. P. Macpherson, 1986, herb. P.M., det. A. Radcliffe-Smith. Still present in 1990. 320/1/3. POLYGONUM RURIVAGUM Jordan ex Boreau 46, Cards.: Cabbage field, Mwnt, GR 22/ 19.52. A. O. Chater, 1990, NMW. 2nd record. 320/1/are. POLYGONUM ARENASTRUM Boreau 50, Denbs.: Rubbish tip, Denbigh, GR 33/05.65. J. A. Green, 1989. 2nd record. 320/3. POLYGONUM MARITIMUM L. 1, W. Cornwall: Shingly sand amongst boulders, Gunwalloe Church Cove, GR 10/66.20. P. Bowman, 1990, det. R. J. Murphy. Only extant locality. 11, S. Hants.: Sand spit, Hengistbury, GR 40/18.91. T. Warwick, 1990, det. A. J. Byfield, D. A. Pearman & R. M. Walls. Ist record since 1902. 320/13. POLYGONUM MITE Schrank 50, Denbs.: Edge of farm pond, Little Green Farm, Penley, GR 33/41.40. J. A. Green, 1990. 2nd record. 320/14. POLYGONUM MINUS Hudson 34, W. Gloucs.: Small dried-out pond, Coalway Lane End, Coleford, GR 32/59.10. K. Herring, 1990. Only extant locality. +320/18. FALLOPIA AUBERTI (Louis Henry) J. Holub *42, Brecs.: Hedge, Llanfilo, GR 32/ 11.33. Hedge, Glangrwyney, GR 32/23.16. Both M. Porter, 1990. 1st and 2nd records. 51, Flints.: Gronant, GR 33/0.8. P. I. Morris, 1985. 2nd record. 428 PLANT RECORDS 7320/20. REYNOUTRIA SACHALINENSIS (Friedrich Schmidt Petrop.) Nakai 49, Caerns.: Waste ground, Glyn Isa Hotel, Rowen, GR 23/76.72. R. Lewis, 1990. 2nd record. *94, Banffs.: Neglected garden, Forglen, GR 38/69.52. J. Edelsten, 1988, E, det. A. P. Conolly. +320/22. POLYGONUM CAMPANULATUM Hooker f. 70, Cumberland: Knowefield Wood, Stan- wix, GR 35/40.57. R. E. Groom, 1989, LANC. 2nd record. 325/11 lit. RUMEX cCRISPUS subsp. LITToREUS (Hardy) Akeroyd *75, Ayrs.: Shingle shore, Knock, Largs, GR 26/19.63. Shingle shore, Ardneil Bay, GR 26/18.48. Both A.McG. Stirling, 1990. lst and 2na records. *99, Dunbarton: Shingle beach, Knockderry, GR 26/21.83. A. Ruther- ford, 1990. +330/tho. ULMUS THOMASII Sarg. *50, Denbs.: Disused gravel quarry, Marford near Wrex- ham, GR 33/35.56. J. A. Green, 1989. 1st Welsh record. +336/2. ALNUS INCANA (L.) Moench *35, Mons.: Roadside hedge, Llanbadoc, GR 31/37.98. T. G. & U. T. Evans, 1988. 343/2 X 4. SALIX ALBA L. X S. FRAGILIS L. *52, Anglesey: Margin of Llyn yr Wyth-Eidion, GR 23/47.81. C. D. Preston & N. F. Stewart, 1988, CGE, det. R. D. Meikle. 343/11b. SALIX CAPREA subsp. SERICEA (N. J. Andersson) B. Flod. *79, Selkirks.: Wooded cleuch, Hermanlaw Burn, Little Yarrow, GR 36/22.16. D. J. McCosh, 1988, herb. D.J.McC., det. R. D. Meikle (as S. caprea var. sphacelata (Sm.) Wahlenb.). Ist record outwith Scottish Highlands. 343/11 X 9. SALIX CAPREA L. X S. VIMINALIS L. *5, S. Somerset: Hedges, Clinton Trinity, GR 31/29.39. P. R. Green, 1990, det. R. D. Meikle. *52, Anglesey: Edge of Llyn Maelog, GR 23/ 32.72. C. D. Preston & N. F. Stewart, 1988, CGE, det. R. D. Meikle. 343/12 * 15. SALIX CINEREA L. X S. PHYLICIFOLIA L. *96, Easterness: Bushy ground, Kincraig, GR 28/83.05. B. H. Thompson, 1990, herb. B.H.T., det. R. D. Meikle. 343/16a. SALIX REPENS L. subsp. REPENS *43, Rads.: Wet pasture, Rhosgoch Common, GR 32/19.48. R. G. Woods, 1990. Ist localized record. 4351/1. GAULTHERIA SHALLON Pursh 70, Cumberland: Woodland, Eskdale Green, GR 35/ 14.00. M. M. Gill, 1990. 2nd record. 354/1. ARCTOSTAPHYLOS UvA-URSI (L.) Sprengel 73, Kirkcudbrights.: Rock ledge in gorge, Louran Burn, Cairnsmore of Fleet, GR 25/48.68. R. W. M. Corner, 1990. 2nd record. 358/4. VACCINIUM OXYCOccos L. 51, Flints.: Upland heather moor, Llyn Cyfynwy, GR 33/2.5. G. Wynne, 1989. 2nd record. 364/2. EMPETRUM HERMAPHRODITUM Hagerup *73, Kirkcudbrights.: Ledges and boulder areas, Cairnsmore of Fleet, GR 25/49.67. R. W. M. Corner, 1990, herb. R.W.M.C. 385/3/2 GENTIANELLA AMARELLA subsp. SEPTENTRIONALIS (Druce) Pritchard *64, Mid-W. Yorks.: Steep limestone grassland, Kettlewell, GR 34/96.72. P. P. Abbott, 1990, det. N. M. Pritchard. Extends range by 160 km to south. +392/ibi. SYMPHYTUM IBIRICUM Stevens *69, Westmorland: Roadside verge south of Bendrigg Lodge, Killington, GR 34/57.88. C. E. Wild, 1990, LANC. 400/8 umb. MyosoTis ARVENSIS subsp. UMBRATA (Rouy) Schwarz *99, Dunbarton: Wood margin near Meikleross Bay, GR 26/26.80. A. McG. Stirling & A. Rutherford, 1986, E. 401/3. LITHOSPERMUM ARVENSE L. 61, S. E. Yorks.: Arable field south-east of Thornton, GR 44/77.44. D. R. Grant, 1990. Only extant locality. 402/1. MERTENSIA MARITIMA (L.) S. F. Gray 73, Kirkcudbrights.: Stones near high-water mark, Howell Bay, GR 25/69.43. O. M. & N. F. Stewart, 1990. 2nd extant locality. PLANT RECORDS 429 406/+2 < 1. CALYSTEGIA PULCHRA Brummitt & Heywood x C. sepium (L.) R. Br. subsp. SEPIUM *54, N. Lincs.: Hedge, Riseholme Lane, Riseholme, GR 43/98.75. I. Weston, 1989, K, det. R. K. Brummitt. 409/1. LYCIUM BARBARUM L. *73, Kirkcudbrights.: Wall, Kirkandrews old churchyard, GR 25/ 60.48. D. Cockerill, 1978. Old wall near Senwick, GR 25/65.46. O. M. Stewart, 1990. Ist and 2nd records. +420/2. LINARIA PURPUREA (L.) Miller 72, Dumfriess.: Street weed, Tynron near Thornhill, GR 25/80.93. M. E. R. Martin, 1990. 2nd record. 420/+2 < 3. LINARIA PURPUREA (L.) Miller X L. REPENS (L.) Miller *46, Cards.: Scrub on railway ballast, Aberystwyth, GR 22/58.81. A. O. Chater, 1990, NMW. With both parents. 420/3 < 4. LINARIA REPENS (L.) Miller x L. vuULGARIS Miller *77, Lanarks.: Along fence, Kelvindale, GR 26/55.68. K. M. Calver, 1964, herb. P. Macpherson. Still present in 1989. 427/1. SIBTHORPIA EUROPAEA L. *35, Mons.: Mossy bank near stream, Cwmfelinfach, GR 31/ 18.92. R. Fraser, 1990, herb. T. G. Evans. 4428/1. ERINUS ALPINUS L. *39, Staffs.: Mossy wall, Wolseley Hall, Colwich, GR 43/02.20. B. R. Fowler, 1990. +430/9. VERONICA LONGIFOLIA L. *69, Westmorland: Roadside verge between Casterton and Kirkby Lonsdale, GR 34/64.79. K. Butler, 1989, LANC. *73, Kirkcudbrights.: Roadside bank south of Cassencarie, GR 25/47.57. O. M. Stewart, 1988. 430/14. VERONICA PEREGRINA L. 81, Berwicks.: Walled vegetable garden, Newton Don, GR 36/70.37. M. E. Braithwaite, 1990, herb. M.E.B. Ist record since 1893 record in same locality. 430/20 hed. VERONICA HEDERIFOLIA L. subsp. HEDERIFOLIA *81, Berwicks.: Waste ground, Duns, GR 36/79.53. M. E. Braithwaite, 1990, herb. M.E.B. 434/3. MELAMPYRUM PRATENSE L. 29, Cambs.: Ditch bank by hedge, Balsham, GR 52/58.51. P. J. O. Trist, 1990, CGE. 1st record since 1930. 61,S.E. Yorks.: Woodland west of Thornton, GR 44/72.45. T. E. Dixon, 1989. Only extant locality. 435. EUPHRASIA. All records below are based on specimens determined by A. J. Silverside. 435/1/2. EUPHRASIA SCOTTICA Wettst. *83, Midlothian: Base-rich upland flush, Trously, GR 36/38.45. D. R. McKean, 1988, E. 435/1/1 X 2. EUPHRASIA MICRANTHA Reichenb. X E. sScorrica Wettst. *42, Brecs.: Montane flush, Cwm Dergwm, GR 32/05.21. M. Porter, 1987, herb. M.P. 1st Welsh record. 435/1/15 X 1 EUPHRASIA CONFUSA Pugsley X E. MICRANTHA Reichenb. *73, Kirkcudbrights.: Forestry track below Craignane, GR 26/57.04. O. M. Stewart, 1988, E. 435/1/15 xX 13. EUPHRASIA CONFUSA Pugsley X E. NEMOROSA (Pers.) Wallr. *35, Mons.: Pathside, Chepstow Park Wood, GR 31/49.97. T. G. & U. T. Evans, 1985, herb. T.G.E. 435/1/16. EUPHRASIA PSEUDOKERNER!I Pugsley 46, Cards.: Rough grassland on cliff top, R. A. E. site, Aberporth, GR 22/24.52. A. O. Chater, 1979, NMW. 2nd record. 435/1/17 X 13. EUPHRASIA ARCTICA subsp. BOREALIS (Townsend) Yeo X E. NEMOROSA (Pers.) Wallr. *35, Mons.: Heathy grassland west of Pontypool, GR 32/26.00. T. G. & U. T. Evans, 1986, herb. T.G.E. Ist Welsh record. 437/1. PARENTUCELLIA VISCOSA (L.) Caruel +95, Moray: Damp, acidic grassland, R. A. F. Lossiemouth, GR 38/19.69. D. Law, 1989, E, det. D. J. Hambler. Only extant locality. 440/10. OROBANCHE HEDERAE Duby +12, N. Hants.: On Hedera hibernica, road between Old Alresford and Bighton, GR 41/59.34. E. Nicholl, 1989, herb. A. Brewis (forma monochroma G. Beck). 2nd record. +*42, Brecs.: Deserted garden, Brecon, GR 32/04.28. C. Cooper, 1990. 1st localized record. 430 PLANT RECORDS 445/2. MENTHA PULEGIUM L. 12, N. Hants.: Edge of pond, Police College, Bramshill, GR 41/ 75.58. B. Meyrick, 1977. 1st record since 1870. 725, E. Suffolk: Porters Wood, Woodbridge, GR 62/26.48. J. Ryland, 1990, det. M. N. Sanford. 445/3 X +5. MENTHA ARVENSIS L. X M. SPICATA L. *42, Brecs.: Edge of pool, Glasbury, GR 32/18.40. M. Porter, 1990, NMW. 445/75 xX 7. MENTHA X VILLOSA Hudson nm. ALOPECUROIDES (Hull) Briq. 7715, AyEse: Foreshore, Hunterston, West Kilbride, GR 26/10.52. A. McG. Stirling & A. Rutherford, 1990. 461/1 arg. LAMIASTRUM GALEOBDOLON subsp. ARGENTATUM (Smejkal) Stace *77, Lanarks.: Grassy bank, Maryhill, Glasgow, GR 26/56.69. P. Macpherson, 1988, herb. P.M., det. A. McG. Stirling. Roadside verge, Moodiesburn, GR 26/70.71. H. J. Noltie & K. Watson, 1988. 1st and 2nd records. 3 465/4/2. GALEOPSIS BIFIDA Boenn. 109, Caithness: Garden weed, Thurso, GR 39/10.68. J. K. Butler, 1977, herb. J.K.B., det. C. C. Townsend. Only extant locality. 472/1 int. PLANTAGO MAJOR subsp. INTERMEDIA (DC.) Arcangeli *39, Staffs.: Dry margin of Knypersley Reservoir, GR 33/89.55. I. W. Brown, 1989, det. J. R. Akeroyd. 73, Kirkcud- brights.: Damp ground by shore west of Kirkandrews, GR 25/59.48. O. M. Stewart, 1990. 2nd record. 472/2. PLANTAGO MEDIA L. 42, Brecs.: Grass verge, Buckland, GR 32/13.21. M. Porter, 1988. 2nd record. 94, Banffs.: Grassland in cemetery, Keith, GR 38/43.51. J. Edelsten, 1990, E. 1st post-1930 record. 484/1. GALIUM CRUCIATA (L.) Scop. *93, N. Aberdeen: Laneside bank, Huntly, GR 38/53.39. D. Welch, 1990. 1st confirmed record. *487/1. SAMBUCUS EBULUS L. 70, Cumberland: Grass verge and hedge bottom, Oulton, GR 35/24.50. R. E. Groom, 1990, LANC. 1st post-1930 record. +491/1. LONICERA XYLOSTEUM L. 50, Denbs.: Deciduous wood, Loggerheads, Mold, GR 33/ 19.62. B. Ing, 1988. 2nd post-1930 record. *95, Moray: Wood on riverbank, Dulnain Bridge, GR 28/99.24. J. Edelsten, 1989, E. *99, Dunbarton: Wood by shore, Cove, Loch Long, GR 26/ 22.82. A. McG. Stirling & A. Rutherford, 1990, E. +491/inv. LONICERA INVOLUCRATA Banks *70, Cumberland: Large patch between road and shore near Cote lighthouse, Silloth, GR 35/11.54. R. E. Groom, 1990, LANC. +491/nit. LONICERA NITIDA Wilson *77, Lanarks.: Along fence, Hutchesontown, GR 26/59.64. Waste ground, Govan, GR 26/55.65. Both P. Macpherson, 1988, herb. P.M., det. D. McClintock. lst and 2nd records. 4506/4. SENECIO SQUALIDUS L. *109, Caithness: Trackside, Pennyland Farm, Thurso, GR 39/ 11.68. J. M. Gunn, 1990, det. J. K. Butler. +BRACHYGLOTTIS Cultivar SUNSHINE C. Jeffrey (Senecio greyi hort.) *77, Lanarks.: Grassy area near river, Meadowside, Glasgow, GR 26/55.66. P. Macpherson, 1988, herb. P.M., det. D. McClintock. +509/3. PETASITES JAPONICUS (Siebold & Zucc.) Maxim. *35, Mons.: Bank of Berthin Brook, Usk, GR 32/370.019. C. Titcombe, 1989, det. T. G. Evans. Bank of Berthin Brook, Usk, GR 32/ 365.019. T. G. & U. T. Evans, 1990, NMW. Ist and 2nd records. *73, Kirkcudbrights.: Banks of Old Water near Rosebank, GR 25/87.76. O. M. Stewart, 1990. 514/2. FILAGO LUTESCENS Jordan 29, Cambs.: Sand pit, Little Abington, GR 52/5.5. B. Jackson & J. Hirsh, 1990, CGE, det. P. D. Sell. 1st post-1930 record. 517/1. ANTENNARIA DIOICA (L.) Gaertner 72, Dumfriess.: Upland moorland near Midlaw Linn, GR 36/1.1. R. W. M. Corner, 1989. Only extant locality. PLANT RECORDS 431 +518/3. SOLIDAGO GIGANTEA Aiton *73, Kirkcudbrights.: Edge of wood west of Summerville, GR 25/95.76. O. M. Stewart, 1980. 2nd record. +536/exa. ECHINOPS EXALTATUS Schrader 25, E. Suffolk: Roadside verge, Finningham, GR 62/06.68. 2nd record. *26, W. Suffolk: Roadside verge south of Daisy Green, Great Ashfield, GR 62/00.67. Both R. Addington, 1989, herb. E. & M. Hyde, det. E. J. Clement. 538/3. ARCTIUM PUBENS Bab. *67, S. Northumb.: Edge of field near Dinnington, GR 45/ 22093: *68, Cheviot: Roadside verge, Cawledge Burn, GR 46/19.10. Roadside verge near East Allerdean, GR 36/98.46. 1st and 2nd confirmed records. All G. A. Swan, 1990, herb. G.A.S., det. F. H. Perring. 542/1. ONOPORDON ACANTHIUM L. +50, Denbs.: Railway embankment, Marford, GR 33/ 36.56. C. R. J. Williams, 1990. 2nd post-1930 record. +544/2. CENTAUREA MONTANA L. *99, Dunbarton: Rough ground near Station, Rhu, GR 26/ 27.84. A. Rutherford, 1975. *109, Caithness: Roadside verge 9 km north-west of Wick, GR 39/ 30.57. M. McCallum Webster, 1978. +552/2. TRAGOPOGON PORRIFOLIUS L. 50, Denbs.: Field gateway, Colwyn Bay, GR 23/85.77. D. Lloyd, 1989. 2nd record. 558. HIERACIUM. Contrary to normal practice, some Ist post-1930 records of Hieracium sect. Alpina are published below. These have been provided by D. J. Tennant, who has revised the British species in this section in collaboration with P. D. Sell. A list of specimens of Hieracium sect. Alpina determined by Mr Tennant has been deposited with the Biological Records Centre. 558/1/5. HTERACIUM HOLOSERICEUM Backh. 87, W. Perth: Rocky ledge near Creagan Liatha, Stob Binnein, GR 27/42.21. D. J. Tennant, 1981, herb. D.J.T. 1st record since 1891. 89, E. Perth: Rocky ground between Cairnwell and Carn nan Sac, Glen Shee, GR 37/12.77. D. J. Tennant, 1974, herb. D.J.T. 1st post-1930 record. 558/1/8. HIERACIUM CALENDULIFLORUM Backh. 90, Angus: Granitic outcrop above Loch Brandy, GR 37/33.75. D. J. Tennant, 1975, herb. D.J.T. 1st post-1930 record. 558/1/9. HIERACIUM MACROCARPUM Pugsley 92, S. Aberdeen: Rocky ground, Soldier’s Corrie, Cairn Toul, GR 27/96.96. D. J. Tennant, 1975, CGE, det. P. D. Sell. 1st record since 1887. 558/1/10. HIERACIUM GRANITICOLA W. R. Linton 94, Banffs.: Rocky slope above Loch Avon, GR 38/00.03. D. J. Tennant, 1981, herb. D.J.T. 1st record since 1898. 96, Easterness: Turf over scree, E. slope of Cairngorm, GR 38/01.04. D. J. Tennant, 1981, herb. D.J.T. 1st record since 1898. 558/1/16. HIERACIUM HANBURY! Pugsley *87, W. Perth: Rock ledges, Stob Garbh, Cruach Ardrain, GR 27/41.21. D. J. Tennant, 1971, CGE. 558/1/32. HIERACIUM DASYTHRIX (E. F. Linton) Pugsley *77, Lanarks.: Falls of Clyde, GR 26/ 88.41. D. J. McCosh, 1988, herb. D.J.McC., det. P. D. Sell. 558/1/41. HIERACIUM FLOCCULOSUM Backh. *77, Lanarks.: Vertical rocks, Falls of Clyde, GR 26/88.41. D. J. McCosh, 1988, herb. D.J.McC., det. P. D. Sell. 558/1/42. HIERACIUM SHOOLBREDII E. S. Marshall *71;,. Lanarks’: ‘Rocks ‘by..R: Clyde; Bonnington Linn, GR 26/8.4. A.McG. Stirling, 1970. 558/1/78. HIERACIUM SANGUINEUM (A. Ley) W. R. Linton *77, Lanarks.: Slab rocks, Falls of Clyde, GR 26/88.41. D. J. McCosh, 1988, herb. D. J. McC., det. P. D. Sell. 558/1/91. HIERACIUM CYMBIFOLIUM Purchas *77, Lanarks.: Slab rocks, Corra Linn, GR 26/ 88.41. D. J. McCosh, 1988, CGE, det. P. D. Sell. 558/1/219. HIERACIUM SABAUDUM L. *81, Berwicks.: Riverside rocks, Preston Bridge, GR 36/ 78.56. M. E. Braithwaite, 1990, det. D. J. McCosh. *94, Banffs.: Roadside verge 1 km south- west of Aberchirder, GR 38/61.51. J. Edelsten, 1987, E, det. J. Bevan. 432 PLANT RECORDS 558/1/ins. HIERACIUM INSIGNE Backh. 92, S. Aberdeen: Rocky slope, Coire an Lochain UVaine, Cairn Toul, GR 27/95.98. D. J. Tennant, 1974, CGE, det. D. J. T. & P. D. Sell. 1st record since 1857. 558/1/mar. HIERACIUM MARGINATUM Sell & C. West *97, Westerness: Mica-schist cliffs, Coire Ardair, Creag Meagaidh, GR 27/43.88. D. J. Tennant, 1979, herb. D.J.T., det. D.J.T. & P. D. Sell. 558/1/mem. HIERACIUM MEMORABILE Sell & C. West 92, S. Aberdeen: Cliff ledge, head of Glen Callater, GR 37/19.81. D. J. Tennant, 1978, herb. D.J.T., det. D. J. T. & P. D. Sell. 1st post- 1930 record. 558/1/ten. HIERACIUM TENUIFRONS Sell & C. West 88, Mid Perth: Crumbling scree, Creag Roro, Ben Lawers, GR 27/63.45. D. J. Tennant, 1979, herb. D.J.T. 1st post-1930 record. 92,S. Aberdeen: Rock gulley, Coire Etchachan, GR 37/01.99. D. J. Tennant, 1976, herb. D.J.T., det. D. J. T. & P. D. Sell. 1st post-1930 record. 558/1/sub. HIERACIUM SUBCRASSUM (Almgq. ex Dahlst.) Johans. *76, Renfrews.: Roadside, Langside Drive, Glasgow, GR 26/57.60. P. Macpherson, 1988, herb. D. J. McCosh, det. D. J. McC. +558/2/8. HIERACIUM BRUNNEOCROCEUM Pugsley 94, Banffs.: Grass verge, Banff, GR 38/ 69.63. J. Edelsten, 1990, E. 2nd record. 4559/3. CREPIS SETOSA Haller fil. *5, S. Somerset: Sea bank, Otterhampton, GR 31/26.44. A.C. Leslie & P. R. Green, 1990, herb. A.C.L. *11,S. Hants.: Lightly shaded streamside path, Bartley Water, Rum Bridge, Totton, GR 41/35.12. Grassy area near Kingfisher Lake, Testwood, GR 41/35.14. Both R. M. Veall, 1990, herb. R.P. Bowman. Ist and 2nd records. *77, Lanarks.: Rough lawn, Shieldhall, Glasgow, GR 26/53.65. P. Macpherson, 1984. Grassy bank, Linthouse, Glasgow, GR 26/54.66. P. Macpherson, 1985. Both herb. P.M., det. C. D. Preston. Ist and 2nd records. 559/8. CREPIS PALUDOSA (L.) Moench *35, Mons.: Wet margin of Afon Cibi, The Park, Abergavenny, GR 32/28.17. R. Fraser, 1990, herb. T. G. Evans. +571/1. LAGAROSIPHON MAJOR (Ridley) Moss 29, Cambs.: Roadside pond, Swavesey, GR 52/ 36.68. J. H. Belcher & E. M. F. Swale, 1989. 2nd record. 557/5 X 1. POTAMOGETON LUCENS L. X P. NATANS L. *27, E. Norfolk: Dyke, Limpenhoe Marshes, GR 63/39.02. C. Doakes & J. Spray, 1989, NWH, det. C. D. Preston & N. F. Stewart. 577/6 X 1. POTAMOGETON GRAMINEUS L. X P. NATANS L. *87, W. Perth: R. Leny south of Loch Lubnaig, GR 27/58.09. C. D. Preston & N. F. Stewart, 1986 and 1990, CGE. *97, Westerness: Edge of Loch Shiel east of Acharacle Pier, GR 17/68.67. M. G. B. Hughes, D. Morgana & C. D. Preston, 1990, CGE. 577/16. POTAMOGETON TRICHOIDES Cham. & Schlecht. [12, N. Hants.: Delete record published in Watsonia 11: 402 (1977); specimen in herb. A. Brewis is P. pusillus L., det. C. D. Preston.] 577/21. POTAMOGETON PECTINATUS L. 43, Rads.: Eutrophic lake, Llanbwchllyn, GR 32/12.46. D. R. Druit & I. D. Soane, 1990, det. R. G. Woods. 2nd record. +586/1. HEMEROCALLIS FULVA (L.) L. *49, Caerns.: Ditch near Ty Mawr, Porth Neigwl, GR 23/25.28. A. P. Connolly, 1990. 1586/2. HEMEROCALLIS LILIOASPHODELUS L. *70, Cumberland: Grassy roadside south of Orton Moss, GR 35/34.54. R. E. Groom, 1989, LANC. 597/1. GAGEA LUTEA (L.) Ker-Gawler 75, Ayrs.: Wooded bank of R. Girvan near Merkland, Kirkmichael, GR 26/34.07. M. Faulds, 1989, conf. A.McG. Stirling, 1990. 2nd record. 598/3. ORNITHOGALUM PYRENAICUM L. +*51, Flints.: Point of Ayr, GR 33/1.8. P. I. Morris, 1988. 605/4. JUNCUS COMPRESSUS Jacq. *77, Lanarks.: Malls Mire, Rutherglen, GR 26/60.62. J. N. Davies, 1985, GL, det. H. J. Noltie. abthles Prandicd aca sade 28 PLANT RECORDS 433 605/amb. JUNCUS AMBIGUUS Guss. 46, Cards.: Muddy depressions in car park by beach, Ynys- las, GR 22/60.92. B.S.B.I. meeting, 1990, det. T. C. G. Rich. 2nd record. 607/6. ALLIUM OLERACEUM L. 26, W. Suffolk: Verge of old railway track, Hadleigh, GR 62/ 03.41. M. Parker & F. W. Simpson, 1989. Only extant locality. +607/7. ALLIUM CARINATUM L. *49, Caerns.: Grassy roadside verge 3 km south of Conway, GR 23/77.74. R. Lewis, 1988, NMW. 607/8. ALLIUM SCHOENOPRASUM L. +*39, Staffs.: Base-rich spoil of old industrial site, Wood Common, Pelsall, GR 43/01.04. J. P. Martin, 1990. +607/10. ALLIUM TRIQUETRUM L. *25, E. Suffolk: Under trees by footpath, Waldringfield, GR 62/28.44. P. Furze, 1989, IPS, det. E. J. Clement. +607/11. ALLIUM PARADOXUM (Bieb.) G. Don *69, Westmorland: Wooded river bank, R. Rothay, Ambleside, GR 35/37.04. A. Franks, 1989, LANC. 616/3. IRIS FOETIDISSIMA L. +*69, Westmorland: Roadside verge, Prince’s Way, Heversham, GR 34/49.83. C. E. Wild, 1990, LANC. 7618/1. CROCUS NUDIFLORUS Sm. *73, Kirkcudbrights: Gravelly island in R. Cree near Creebridge, GR 25/41.65. O. M. Stewart, 1988. +620/pot. Crocosmia poTtsi (Baker) N. E. Brown *73, Kirkcudbrights.: Embankment by R. Dee below Glenlochar Dam, GR 25/73.64. O. M. Stewart, 1982, E. Bank of R. Cree below Minnigaff, GR 25/41.65. O. M. Stewart, 1982. 1st and 2nd records. 627/1. SPIRANTHES SPIRALIS (L.) Chevall. *42, Brecs.: Old lawn, Llangattock, GR 32/21.18. S. Sankey-Barker, 1989. 627/3. SPIRANTHES ROMANZOFFIANA Cham. 103, Mid Ebudes: Partially drained bog, Mornish, GR 17/3.5. F. Horsman, 1990. Ist Mull record. 635/1. COELOGLOSSUM VIRIDE (L.) Hartman 25, E. Suffolk: Meadow on boulder-clay, Winks Meadow, Metfield, GR 62/30.79. E. Krutysza, 1990, det. M. N. Sanford. Only extant locality. 636/1b. GYMNADENIA CONOPSEA subsp. DENSIFLORA (Wahlenb.) G. Camus, Bergon & A. Camus *105, W. Ross: Marsh near Mellon Charles, GR 18/8.9. A. G. Kenneth, M. R. Lowe & D. J. Tennant, 1984, E, det. D. J. T. & R. H. Roberts. 642/7. ORCHIS MASCULA (L.) L. 72, Dumfriess.: Ledges above Tail Burn near Loch Skeen, GR 36/18.15. R. W. M. Corner, 1989. Only extant locality. 643/1 X 3. DACTYLORHIZA FUCHSII (Druce) So6 X D. INCARNATA (L.) S06 *41, Glam.: Kenfig Burrows, GR 31/79.81. D. C. Lang, 1990. 643/1 X 3b X 7. DACTYLORHIZA FUCHSII (Druce) Sod X D. INCARNATA subsp. PULCHELLA (Druce) So6 X D. TRAUNSTEINERI (Sauter) S06 *62, N. E. Yorks.: Marsh near Thornton Dale, GR 44/ 8.8. F. Horsman, 1989. 643/1 x 4. DACTYLORHIZA FUCHSII (Druce) So6 X D. PRAETERMISSA (Druce) S06 *46, Cards.: Moist pasture above Plas Cwmcynfelin, Llangorwen, GR 22/60.83. A. P. Fowles, 1989, det. D. C. Lang. 643/2b xX 4. DACTYLORHIZA MACULATA subsp. ERICETORUM (E. F. Linton) P. F. Hunt & Summerhayes X D. PRAETERMISSA (Druce) So6 *42, Brecs.: Marshy meadow near Llangorse Lake, GR 32/12.26. M. Porter, 1990. 643/2b X 5 maj. DACTYLORHIZA MACULATA subsp. ERICETORUM (E. F. Linton) P. F. Hunt & Summerhayes X D. PURPURELLA subsp. MAJALIFORMIS E. Nelson *110, Outer Hebrides: Coastal marsh, Borve, S. Harris, GR 18/02.94. A. G. Kenneth & D. J. Tennant, 1985, herb. D.J.T., det. D. TEGPsOORGHS Roberts. 434 PLANT RECORDS 643/3c < 5 maj. DACTYLORHIZA INCARNATA Subsp. COCCINEA (Pugsley) Sod X D. PURPURELLA subsp. MAJALIFORMIS E. Nelson *110, Outer Hebrides: Coastal marsh, Borve, S. Harris, GR 18/ 02.94. A. G. Kenneth & D. J. Tennant, 1985, herb. D.J.T., det. D. J. T. & R. H. Roberts. 643/4 x 5. DACTYLORHIZA PRAETERMISSA (Druce) So6 X D. PURPURELLA (T. & T. A. Stephenson) So6 *64, Mid-W. Yorks.: Calcareous area in Lawkland Moss, GR 34/76.66. F. Horsman, 1990. 643/5. DACTYLORHIZA PURPURELLA (T. & T. A. Stephenson) Sod *41, Glam.: Dune slacks, Kenfig Burrows, GR 21/79.81. D. C. Lang, 1990. 643/5 maj. DACTYLORHIZA PURPURELLA subsp. MAJALIFORMIS E. Nelson *105, W. Ross: Marsh near Mellon Charles, GR 18/8.9. A. G. Kenneth, M. R. Lowe & D. J. Tennant, 1984, herb. D.J.T., det. D. J. T. & R. H. Roberts. *110, Outer Hebrides: Coastal marsh, Northton, S. Harris, GR 08/99.90. Coastal marsh, Borve, S. Harris, GR 18/02.94. Both A. G. Kenneth, M. R. Lowe & D. J. Tennant, 1985, herb. D.J.T., det. D. J. T. & R. H. Roberts. 1st and 2nd records (plant previously reported as D. majalis subsp. occidentalis (Pugsley) P. D. Sell). 643/7. DACTYLORHIZA TRAUNSTEINERI (Sauter) So6 *45, Pembs.: Flush in calcareous fen, Castlemartin Corse, GR 11/89.99. J. Donovan, S. B. Evans & D. C. Lang, 1990. 1st Welsh record. 643/5 x 636/1. DACTYLORHIZA PURPURELLA (T. & T. A. Stephenson) Sod X GyYMNADENIA CONOPSEA (L.) R. Br. *105, W. Ross: Wet ditch near Mellon Charles, GR 18/8.9. A. G. Kenneth, M. R. Lowe & D. J. Tennant, 1984, E, det. D. J. T. & R. H. Roberts. +648/1. LySICHITON AMERICANUS Hultén & St John *35, Mons.: Ditch in wood, Llanarth Fawr, GR 32/36.08. C. Titcombe, 1990. 654/2. ERIOPHORUM GRACILE Roth 41, Glam.: Fen, Pant-y-Saes Fen near Neath, GR 21/71.94. C..R. Hipkin, 1990. 2nd record. 654/3. ERIOPHORUM LATIFOLIUM Hoppe 50, Denbs.: Marshy field, Graianrhyd, GR 33/20.54. J. A. Green, 1989. 1st record since 1830. 655/12. ELEOGITON FLUITANS (L.) Link 61, S. E. Yorks.: Dike c. 1-5 miles west of Thornton, GR 44/72.45. T. E. Dixon, 1990. Only extant locality. 657/1. BLYSMUS COMPRESSUS (L.) Panzer & Link *42, Brecs.: Small marsh by tufa outcrop, Hay-on-Wye, GR 32/2.3. M. Porter, 1989, NMW. Ist record, and only extant Welsh locality. 658/1. CYPERUS LONGUS L. +35, Mons.: Pond margin, Kilgwrrwg, GR 31/47.97. T. G. Evans, 1990, herb. T.G.E. Only extant locality. 663/16 X 17. CAREX ROSTRATA Stokes X C. VESICARIA L. *47, Monts.: With C. rostrata at marshy edge of lake, Maesmawr Hall, GR 33/1.0. P. M. Benoit, 1989, NMW, det. A. O. Chater & | A. C. Jermy. 663/21 * 20. CAREX ACUTIFORMIS Ehrh. X C. RIPARIA Curtis *25, E. Suffolk: Framlingham Mere, GR 62/28.63. A. C. Jermy, 1990. 663/47 X 21. CAREX ACUTA L. X C. ACUTIFORMIS Ehrh. *25, E. Suffolk: Framlingham Mere, GR 62/28.63. A. C. Jermy, 1990. 663/23. CAREX STRIGOSA Hudson 69, Westmorland: Clayey drainage channel, High Wood, Askham-in-Furness, GR 34/20.76. P. Burton, 1989, LANC. Ist post-1930 record. 663/24. CAREX PALLESCENS L. *81, Berwicks.: Wet meadow, Leescleugh Burn, GR 36/74.52. M. E. Braithwaite, 1990, herb. M.E.B. Ist record since 1880. Flushed grassland, Wellcleugh Burn, GR 36/73.52. M. E. Braithwaite, 1990. 2nd extant locality. 663/28. CAREX LIMOSA L. 50, Denbs.: Soligenous flush, Llansannan, GR 23/93.61. P. Day, 1989, det. A. O. Chater. 2nd record. 663/52. CAREX BIGELOW Torrey ex Schweinitz *47, Monts.: Moel Sych, GR 33/0.3. A. P. & R. A. Dawes, 1990, NMW, det. P. M. Benoit. Confirms hitherto disregarded record from same locality. 72, Dumfriess.: Moffat Hills, GR 36/1.1. R. W. M. Corner, 1989. 2nd extant locality. PLANT RECORDS 435 663/54 X 71. CAREX PANICULATA L. X C. REMOTA L. *35, Mons.: With parents in woodland flush, Cwm Merddog, GR 32/18.06. R. Fraser, 1990, herb. T. G. Evans, det. A. O. Chater. *47, Monts.: With parents in marshy oak-birch woodland near Penegoes, GR 23/7.0. A. Morton, S. Grasse & P. M. Benoit, 1990, NMW, det. A. O. Chater & A. C. Jermy. *96, Easterness: Swamp, Loch Battan, GR 28/53.38. D. Gilbert, 1988, herb. N.C.C. Inverness, det. A. C. Jermy. 663/57. CAREX OTRUBAE Podp. *77, Lanarks.: Damp grassland, Garthamlock, Glasgow, GR 26/66.66. K. Watson, 1986, GL. Ist localised record. +663/59. CAREX VULPINOIDEA Michx 77, Lanarks.: Waste ground, Hallside near Cambuslang, GR 26/66.60. J. H. Dickson, 1989, GL. 2nd record. 663/61. CAREX ARENARIA L. +*47, Monts.: Railway ballast, Dovey Junction Station, GR 22/ 6.9. P. M. Benoit, 1989, NMW. 663/69. CAREX ELONGATA L. 12, N. Hants.: Swampy area under Salix by R. Blackwater, Ash, GR 41/88.51. C. R. Hall, 1990, herb. A. Brewis, det. F. Rose. Only extant locality. 670/2 X 1. FESTUCA ARUNDINACEA Schreber X F. PRATENSIS Hudson *38, Warks.: Meadow, Moreton Morrell, GR 42/30.55. J. C. Bowra, 1990, WAR, det. P. J. O. Trist. 670/2 X 3. FESTUCA ARUNDINACEA Schreber X F. GIGANTEA (L.) Vill. *42, Brecs.: Shaded margin of pool, Glasbury, GR 32/18.40. M. Porter, 1990, NMW. +670/5. FESTUCA HETEROPHYLLA Lam. 35, Mons.: Woodland edge, The Hill, Abergavenny, GR 32/29.15. R. Fraser, 1987, NMW, det. R. F. & T. G. Evans. 2nd vice-county and Welsh records. *73, Kirkcudbrights.: Grassy edge of Marbreck Burn, Burnfoot, GR 25/59.92. O. M. Stewart, 1989, E, det. P. J. O. Trist. Edge of roadside wood, Garroch, GR 25/59.80. O. M. Stewart, 1990, E. Ist and 2nd records. 670/6b. FESTUCA NIGRESCENS Lam. subsp. NIGRESCENS 2, E. Cornwall: Hedgebank, West Looe, GR 20/23.52. R. W. Gould, 1989, LTR, det. A.-K. Al-Bermani. Ist record since 1926. 673/5. PUCCINELLIA RUPESTRIS (With.) Fernald & Weatherby 53, S. Lincs.: Track behind sea wall between R. Ouse and R. Nene, GR 53/51.26. S. J. Leach, 1989. 2nd record. 674/2. DESMAZERIA MARINA (L.) Druce *47, Monts.: Dovey Junction Station, GR 22/6.9. P. M. Benoit, 1989, NMW. 667/1 aqu. CATABROSA AQUATICA L. subsp. AQUATICA *75, Ayrs.: Near Ardrossan, GR 26/2.4. W. Barclay, 1988, E. Near Burnfoot, Bennane, GR 25/11.88. D. Counsell, 1990, herb. A. McG. Stirling, det. A. McG. S. Ist and 2nd records. 667/1 min. CATABROSA AQUATICA subsp. MINOR (Bab.) F. Perring & Sell *75, Ayrs.: Seamill, West Kilbride, GR 26/2.4. R. H. Meldrum, 1889, E, det. A.McG. Stirling. +680/3. BRIZA MAXIMA L. *75, Ayrs.: Foreshore, Ballantrae, GR 25/08.82. J. G. Michie, 1960, E. Quarry, Noddsdale, Largs, GR 26/22.64. A. McG. Stirling, 1989, E. 1st and 2nd records. 681/2. MELICA NUTANS L. 93, N. Aberdeen: Gorge, Gordonsburn, GR 38/47.38. D. Welch, 1989, ABD. Ist record since c. 1765. +683/4. BROMUS INERMIS Leyss. *67, S. Northumb.: Reclaimed pit heap near Rising Sun Farm, GR 45/29.68. D. Mitchell, 1990, herb. G.A. Swan, det. P. J. O. Trist. +683/7. BROMUS DIANDRUS Roth *77, Lanarks.: Pavement, Cadder, GR 26/61.72. J. H. Dickson, 1989, GL. More abundant in 1990. Foot of wall, Meadowside, Glasgow, GR 26/55.66. P.Macpherson, 1989, herb. P.M., det. P. Smith. Still present in 1990. 1st and 2nd records. £93. N. Aberdeen: Set-aside field, Pitcaple, GR 38/71.24. D. Welch, 1990, ABD. ¥683/9. BROMUS TECTORUM L. 11, S. Hants.: By new road, West End, Southampton, GR 41/ 48.14. P. D. Stanley, 1990, herb. R.P. Bowman. 2nd extant locality. *42, Brecs.: Garden weed, Llangynidr, GR 32/16.19. M. Porter, 1988, det. T. A. Cope. 436 PLANT RECORDS 7683/19. BROMUS CARINATUS Hooker & Arnott *28, W. Norfolk: Verge of track, Feltwell, GR 52/67.89. R. Tofts, 1989. Verge of green lane, Methwold, GR 52/73.91. R. Tofts & K. A. & G. Beckett, 1989. Ist and 2nd records, both det. P. J. O. Trist. 51, Flints.: Foot of church wall, Rhuddlan. GR 33/02.78. T. C. G. Rich, 1987, det. T. A. Cope. 2nd record. 685/4 x 3. ELYMUS PYCNANTHUS (Godron) Melderis x E. REPENS (L.) Gould *46, Cards.: Edge of golf course, Ynys-las, GR 22/60.93. A. O. Chater, 1990, K, det. T. A. Cope. +697/3. AIRA CARYOPHYLLEA subsp. MULTICULMIS (Dumort.) Bonnier & Layens *77, Lanarks.: Bare ground, Shieldhall, GR 26/53.66. Sandy ground, Westburn, Cambuslang, GR 26/65.60. Both P. Macpherson, 1989, herb. P.M., det. P. J. O. Trist. 1st and 2nd records. 700/3. CALAMAGROSTIS STRICTA (Timm.) Koeler 75, Ayrs.: Wetlands by R. Doon between Waterside and Dalmellington, GR 26/45.07. D. Counsell, 1990, E, det. A. McG. Stirling. 2nd record. a 701/4. AGROSTIS GIGANTEA Roth 81, Berwicks.: Arable land near Dunslaw Farm, GR 36/ 79.55. M. E. Braithwaite, 1990, det. R. W. M. Corner. 2nd record. 701/sca. AGROSTIS SCABRA Willd. *46, Cards.: Graveyard, Llangwyryfon, GR 22/59.70. P. Macpherson, 1987, NMW, det. C. A. Stace. 2nd Welsh record. 708/4. ALOPECURUS AEQUALIS Sobol. 5, S. Somerset: Drying sand at edge of Hawkridge Reservoir, GR 31/20.35. and 20.36. P. R. Green, 1990. 2nd record. 708/4 xX 3. ALOPECURUS AEQUALIS Sobol. X A. GENICULATUS L. *24, Bucks.: Lakeside, Tiddenfoot, Leighton Linslade, GR 42/91.23. C. M. Dony, 1986, LTN, det. C. A. Stace & P. J. O. Trist. 708/5 X 3. ALOPECURUS BULBOSUS Gouan X A. GENICULATUS L. *5,S. Somerset: Fixed dunes, Wall Common, GR 31/26.45. P. Harmes, 1990, det. A. C. Leslie. *54,N. Lincs.: Old saltmarsh, now grazed by ponies, Winteringham Haven, GR 44/93.23. W. Earnshaw, 1989, det. P. J. O. Trist. 708/6. ALOPECURUS ALPINUS Sm. 72, Dumfriess.: Flushed ledges, White Coomb, GR 36/1.1. R. W. M. Corner, 1989. Only extant locality. 711/1. H1EROCHLOE ODORATA (L.) Beauv. 109, Caithness: Sand dune, Dunnet Sands, GR 29/ 21.70. J. K. Butler, 1978, E, det. S. M. Walters. 2nd record. 714/2. PARAPHOLIS INCURVA (L.) C. E. Hubbard 5, S. Somerset: By pool on sandy beach, Greenaleigh, GR 21/95.48. C. J. Giddens, 1990. 2nd record. 54, N. Lincs.: Sand on top of sea defences, Trusthorpe, GR 53/51.83. J. Southey, 1989. 2nd record. Watsonia, 18, 437-445 (1991) 437 Book Reviews John Lightfoot: his work and travels, with a biographical introduction and a catalogue of the Lightfoot Herbarium. J. K. Bowden. Pp. vi + 255, illust. The Bentham-Moxon Trust, Kew, and Hunt Institute for Botanical Documentation, Pittsburgh. 1989. Price £16 (ISBN 0—-913196—-S1-7). The contribution made by the English clergy to the development of descriptive British natural history, especially during the eighteenth and nineteenth centuries, was nothing short of phenome- nal. The Reverend Gilbert White (1720-1793) is perhaps the most famous, the Reverend William Kirkby (1759-1850), father of British entomology, not far behind, and here, of course, is Whites’s friend, the Reverend John Lightfoot, M.A., F.R.S., F.L.S. (1735-1788), who is the subject of a new biography by Jean Bowden. Lightfoot’s reputation is founded upon his botany and conchology: his Flora Scotica, 1777 and the Catalogue of the Portland Museum, 1786, being the somewhat slender published evidence of his activities. It is through the statements of his many correspondents, however, that a broader picture unfolds of his wide-ranging knowledge, his unselfish help to others, and the pursuit of scholarly excellence. Lightfoot’s close friend Thomas Pennant (1725-1798) summed things up: ““He was an excellent scholar in many branches of literature; but after the study of his profession, he addicted himself chiefly to that of botany and conchyliologie [sic]. He excelled in both.” Not much has been written hitherto about the life of John Lightfoot. Thomas Pennant, zoologist, traveller, and correspondent of many of the eminent figures of natural philosophy in the eighteenth century, knew Lightfoot well. In the second edition of Flora Scotica, 1789, published just after Lightfoot’s premature death, Pennant included an additional preface entitled ‘Some account of the author of this work.’ In 1819 James Edward Smith’s short biography appeared in Rees’ Cyclopaedia, and, apart from a short obituary notice in The Gentleman’s Magazine in 1788, that is all. John Lightfoot was born on 9 December 1735 at Newent in the Forest of Dean, about 13 km from Gloucester, the son of a gentleman farmer. Little is known of his upbringing until 1753, when at the age of seventeen he entered Pembroke College, Oxford. He graduated in 1756, and proceeded to an M.A. in 1766. After taking holy orders, he was appointed curate at Colnbrook, Middlesex, sometime between 1756 and 1762. In 1765 Lord Chancellor Northington gave him the living of Shelden, near Alton in Hampshire, a stone’s throw from Gilbert White’s parish of Selborne, which he held until 1777. He married Matilda Raynes, daughter of a wealthy mill-owner of Uxbridge, Middlesex in 1780, when he was nearly 45 and she just 20. They had three daughters and two sons, although one daughter died before the age of two. Like so many churchmen, Lightfoot used his spare time for the observation of nature, study and research. It was almost certainly as a result of this that in 1767 he was chosen, at the age of 31, to become Chaplain to the Dowager Duchess of Portland (the second Duke having died in 1762). Lady Margaret Cavendish Bentinck, only daughter and heiress of the second Earl of Oxford, was the leading patroness of the natural sciences in England during her lifetime. Sir Joseph Banks (1743- 1820) and Daniel Solander (1733-1782), fresh from their triumphant circumnavigation in 1768-1771 with Cook, were frequent visitors to the Dowager Duchess’s great house at Bulstrode in Buckinghamshire. The focus of attention there was her natural history collection on which she lavished most of her fortune and energy. On the Dowager Duchess’s death in 1785, Lightfoot was mainly responsible for drawing up the sale catalogue, the well-known Catalogue of the Portland Museum, 1786. Lightfoot’s travels to Scotland (1772) and Wales (1773) were undertaken at the instigation of Thomas Pennant and Sir Joseph Banks respectively. Pennant had early recognised the excellence of Lightfoot’s botany and invited him to be his companion on a five-month tour in Scotland, including the Hebrides, for the purpose of observing and working up the botany of a region little studied in that respect. Pennant strongly encouraged Lightfoot to prepare the work for publication and actually provided financial support for Flora scotica which came out in 1777. It was the first Flora of the north of Britain in which Linnaean binomials were used. However, during Lightfoot’s lifetime 438 BOOK REVIEWS Flora scotica was heavily criticised by those perhaps envious of Pennant’s financial support and it was not the success it deserved to be. Lightfoot’s tour in Wales with Banks did not result in a publication. Jean Bowden's new biography, written whilst she was employed at Kew, is based on Lightfoot’s letters to his friends and associates, especially the Dowager Duchess of Portland, Sir Joseph Banks, Sir John Cullum, Thomas Pennant, William Curtis and Gilbert White. She has scoured libraries and record offices for material, photocopies of which have been deposited in the Kew Archives. By using extensive and numerous quotes from this correspondence, Ms Bowden has tackled the organisation of the biography in an unconventional way. A long introduction outlining Lightfoot’s life precedes sections devoted to the principal events — the tours in Scotland, Wales, Devon and Cornwall, Portland Catalogue and British Flora-— using quotations to chart a step-by-step account of itineraries and activities. A final section consists of a Catalogue of the Lightfoot Herbarium at Kew, and five Appendices. | The Catalogue of the Lightfoot Herbarium comprises, at 90 pages, over a third of the volume. With the exception of modern identifications, verified by expert helpers, summary counts of Lightfoot’s folders and sheets and any attributions or comments, all the information in the entries is taken directly from the writing on the folders or sheets. The Catalogue is extraordinarily detailed. Its ease of use would have been enhanced by inclusion in the index of names used in it. This point of criticism applies also to names which appear in the Appendices. The index is rather basic and reduces usefulness of the biography for quick-reference purposes. Scholarship in biographical study can result in exceptional factual accuracy but also a loss of the readability. Here the over-use of quotations from newly-revealed sources becomes almost an irritation. The reader longs for some analysis and speculation to chew upon. At least footnotes have been omitted though, and all references are gathered together into three pages at the end. Typography and lay-out are workmanlike, but the nine illustrations have printed poorly in the text. The map of Scotland showing Lightfoot’s itinerary requires the use of a magnifying glass. The overwhelming impression of this biography is that much new light is shed on John Lightfoot’s natural history activities through recently uncovered manuscript sources. For botanists, the Catalogue of his herbarium, linked with a detailed investigation of collecting localities, will provide much essential information about the British flora. For the student of biography, Ms Bowden’s study fills another gap in the who’s who of eighteenth century British natural history. R. E. R. BANKs The botanist in Berwickshire. M. E. Braithwaite & D. G. Long. Pp. 111, with map. The Berwickshire Naturalists Club. 1990. Price £6 (ISBN 0—9516434—0-1) This attractively produced work is an excellent example of the type of local botanical guide now becoming popular, providing the reader with something more than a basic check list though falling short of a comprehensive flora. The soft cover is attractively designed, incorporating a very professional colour illustration of Rock Rose, and the size — a slim 20 x 15 cm — makes for easy portability. The inclusion of a bryological section is somewhat unconventional and there may be differing views as to the desirability of this format, but in an economically priced work there is much to commend the combination. An introductory section sets out the format and scope of the contents and includes an historical account of botanical recording in the county. The order of species and their scientific names follows the 3rd edition of Clapham, Tutin & Warburg’s Excursion Flora and the common names are according to English Names of Wild Flowers, 2nd edition. Local and possibly unfamiliar names are rarely used. The section on flowering plants and ferns by Michael Braithwaite (B.S.B.I. Recorder for v.c 81) is preceded by an overview of the flora based on the principal habitat types found in the county, indicating their botanical features and the changes which have taken place over the years. A statistical summary of the native and introduced flora indicates its composition in terms of species, subspecies, microspecies and hybrids. The recorded taxa, past and present, total 1151, of which 946 are known to occur at the present day. The check list which follows provides basic information for each species, ranging from a simple BOOK REVIEWS 439 indication of relative abundance in the case of the more common plants, to more explicit notes including recording dates and sites for the more interesting or localised species. Conventional symbols indicate non-native species and those considered no longer present. Rather unconventio- nally an indication is also given of species for which there is no record, but which might be expected in view of their national distribution. A useful gazetteer assists the user to identify many of the localities mentioned in the text, and there is a short bibliography. Readers may find the lack of a comprehensive index inconvenient. Only the genera are included, presumably in the interests of space-saving. For the convenience of those more familiar with common names there is a table indexing the families of flowering plants and ferns, but the value of this is somewhat reduced by its not being listed alphabetically. Of the bryological section, which is by David Long of the Royal Botanic Garden, Edinburgh, and will no doubt be suitably reviewed elsewhere, it is only necessary to say that this is very comprehensive and follows the same format of overview followed by a check list. The authors of The botanist in Berwickshire are to be commended for the production of this useful volume which should prove a popular guide to the flora of the area. A. McG. STIRLING Sussex plant atlas: selected supplement. M. Briggs. Pp. 32, with introductory notes and 37 distribution maps. Booth Museum of Natural History, Brighton. 1990. Price £3.50 (ISBN 0-948723- 14-9). This book is intended to be used in conjunction with the Sussex plant atlas published by P.C. Hall in 1980 (see review by Rose in Watsonia 13: 353-354, 1981). It has the same format and the same attractive cover. It is to be hoped that the colour reproduction of the latter is not so poor on all copies as it is on mine. The book consists of a selection of records made by members of the Sussex Botanical Recording Society from 1979 to 1988. Only those records for which the more recent information adds significantly to the Plant Atlas have been included. The brief text is interesting and informative. It includes corrections of errors published in the Atlas, effects of the 1987 storm and sites of deliberate introduction of native species. There are some enviable new county records and rediscoveries. Those plants which Francis Rose considered under-recorded in his review of the Atlas have clearly been chased up and a more comprehensive distribution pattern results. I would have thought that some of these species are still under-recorded, but perhaps Sussex is fortunate and its ponds and waterways are not disappearing under that unwelcome trio: Crassula helmsii, Lemna minuscula and Elodea nuttallii. Clearly no-one except the author was interested in looking for further specimens of Prunus X fruticans. A list of aliens and adventives additional to those in the AZlas is given. It is surprising that this does not include the well established colony of Tropaeolum speciosum at Wivelsfield Green. With the assistance of Jenny Moore of the Natural History Museum, an account has been given of the Sussex Charophytes, with tetrad maps for four of the species. Owners of the Sussex plant atlas will need a copy of this Supplement to bring their botanical knowledge of the county up to date. J. E. SMITH The Davis & Hedge Festschrift. Edited by Kit Tan, assisted by R. R. Mill & T. S. Elias. Edinburgh University Press. 1989. Pp. xxvi + 351; 1 colour plate and many b/w illustations. Price £47.50 (ISBN 0—85224-638-2). The subtitle of this book, “‘Plant Taxonomy, Phytogeography and Related Subjects’’, describes not only the scope of the 25 commissioned articles but also the life-long research goals of the two distinguished Edinburgh botanists in whose honour it was published. It starts with a biographical 440 BOOK REVIEWS sketch of Peter H. Davis (‘P. D.’ to his friends and colleagues), a list of his publications, an illustrated description of Biarum davisii and an even briefer account of the career of Ian C. Hedge with an extensive bibliography. Both men have a lifetime’s involvement with taxonomic botany at Edinburgh, one as a University lecturer and professor as well as editor of the Flora of Turkey, the other as a taxonomist and curator of the herbarium at the Royal Botanic Gardens and contributor to most of the post-war Floras of south-west Asia. It is appropriate that south-west Asian botany should be the theme of most of the authors selected by Kit Tan, all of whom have had a close professional relationship with Davis and Hedge. One contribution, Jennifer Lamond’s ‘Plantsmen and Pottery’, reflects the private interests of Peter Davis as a collector of ceramics, while Karl-Heinz Rechinger’s account of ‘Fifty years of botanical research in the Flora Iranica area (1937—-1987)’ is a tour de force which should be read by anyone contemplating botanical field-work in the region. Rupert Barneby’s account of ‘A far-rolling stone’ deals with the bizarre story of a ‘dimpled egg’ excavated in a paddock in Leicestershire which proved to be the seed of a Brazilian rain-forest liana; it reminds us that the flora of the British Isles is full of surprises. There is little of purely local interest in this book; it is hard to see who might buy a personal copy, since its price of nearly £50 represents a major outlay for what is a rather eclectic range of topics. Academic libraries, on the other hand, will find its articles to be much sought after and will need to buy it. It is a worthy momento to mark the 70th and 60th birthdays of two of Edinburgh’s outstanding botanists, who are widely respected for their taxonomic expertise and judgement; it also displays the high standards of editorship for which they are known. J. R. EDMONDSON Plants for people. A. Lewington. Pp. vii + 232; lavishly illustrated. Natural History Museum Publications, London. 1990. Price £19.95 (ISBN 0—-565—01094-8). Plants for people is exactly as its title suggests. Each one of the seven chapters lucidly describes the botanical basis of our daily lives: plants that cover us; plants that feed us; plants that protect us; plants that cure us; plants that transport us and plants that entertain us. A purist would categorize the book as Economic or Ethno-Botany, but these descriptions would fail to do justice to the highly readable, informative and entertaining nature of its contents. Plants for people does not become bogged down in any one aspect of plants or their uses, but skilfully takes the reader through the scientific aspects of plants and their extracts, to why they are used as they are and at what cost both in financial and environmental terms. The book is as up to date as it could be, telling us that a chemical (castanospermine) from the seed of an Australian evergreen tree called the Moreton Bay Chestnut (Castanospermum australe) has recently been isolated and found to have a dramatic effect on the AIDS virus HIV. The historical development of plant use is also well presented. The Egyptians used plant oils and fragrant flowers in their ablutions. The Romans were aware of the disinfectant properties of Lavender. Indeed, its scientific name, Lavandula, is derived from the Latin ‘lavare’ meaning to wash. “Ring a ring of roses, a pocket full of posies”’ testifies to the once held belief that carrying a posy of fragrant flowers would ward off disease. The development and context of plant use is by no means confined to the western or Classical world. Some of the uses made of plants and their by-products by indigenous peoples worldwide have also been the subject of research. Colouring the skin, particularly the face, with patterns of pigments conveys important messages about the wearer be they from Burma, New Guinea, Japan, Cameroon or the Indian sub-continent. The main text guides the reader through the information at an easy but attention-holding pace. Every second page presents a profile of a particular plant which gives a little more in-depth information about it, the areas in which it is found and the uses to which it has been put by people. As a book it is well constructed and any reader will appreciate the unpretentious language, the excellent and numerous colour plates and illustrations and the excellent but over-done design. It is also gratifying to find a comprehensive index. In her Foreword to Plants for people, Anita Roddick expresses the hope that when people read BOOK REVIEWS 44] this book they will gain a renewed appreciation of what plants do for people the world over. In that it succeeds. It is not just another contribution to environmental friendship. It is a book whose literary yet scientific approach gradually reveals the fundamental part played by plants in our various lifestyles and makes the reader consider and realise the implications of this relationship. It neither hesitates in nor apologises for telling the facts as they are. It gives everyone, be they scholar or amateur, an opportunity to reflect upon the impact they themselves make on the environment by doing something as simple as brushing their teeth. The Royal Botanic Gardens, Kew, the Natural History Museum and, above all, Anna Lewington have provided their public with a worthwhile and informative read. E. I. KWASNIK A guide to some difficult plants. Edited by E. Norman. Pp. 131 with numerous line drawings and keys. Wild Flower Society, Loughborough. 1990. Price £6. Available from 68 Outwoods Road, Loughborough, Leics. This booklet usefully gathers together several series of articles from the Wild Flower Society Magazine from 1973-88, published by the generosity of the mother of Christine Hibbert, a member who died too young. The authors are all experts in their fields and although the chatty nature of the articles occasionally gives a feeling of talking down, they are packed with useful facts to aid all field botanists. John Mason’s guide to bird-seed aliens is full of drawings you will find elsewhere in British books, with keys and salient features for a wide range of species. [t makes one wish that rubbish-tips were as rich as they used to be. Willows are comprehensively covered by the Howitts, with hints from a lifetime’s experience augmented by the first of Olga Stewart’s excellent drawings. Alpine willows are not considered. Alan Silverside takes us skilfully through yellow Compositae, even tackling Hieracium and Taraxacum (inevitably the least useful part of the book). Next, Juncus and Luzula by Dr Tom Cope are dealt with in some depth including J. subulatus, J. planifolius, the J. bufonius splits and some hybrids. But I feel you would look in vain for J. capitatus on Anglesey these days. The seed characters of L. campestris and L. multiflora are not mentioned. Norman Robson gives a fascinating discourse on Hypericum. I had not realised that androsaemum signified ‘man’s blood’ after the red sap of Tutsan. The genus is placed in its world context which helps to explain several puzzles. Major-General Turpin deals with heathers, calling on a wide range of sources listed at the end. Considerable detail about all species and hybrids is given and we are told how to spot and conclusively identify Erica X watsonii (tetralix X ciliaris) which the inevitable visitors to Dorset would value. Comments about the 47 variants of Calluna described make one want to know more! Finally, Tim Rich gives a foretaste of his Crucifer Handbook in a chapter on “nasty” yellow crucifers. Few people would not find useful his forthright comments on which books not to use and the practical hints on how and when to determine his favourite plants. ‘All You Want to Know About Yellow Crucifers’ might be an alternative title. So, if you are active in the field or want your appetite stimulating, here is a very good and cheap booklet to help you. Buy it before it is out of print. G. M. Kay Atlas of the British flora. Edited by F. H. Perring & S. M. Walters. Pp. xxiv + 444. Botanical Society of the British Isles, London. 1990. Price £20 (ISBN 0—90115—819-4). The Atlas has had a long history; it was first published by Thomas Nelson and Sons in 1962 and reprinted only a year later, such was the importance and demand. The work was originally reviewed by D. H. Kent in Watsonia 5: 396-397 (1963). A second edition was produced by Epworth Press in 1976 with two more reprints before a third edition in 1982. The changes between the editions are rather less dramatic than might be supposed, for the second edition only updates the rare species. 442 BOOK REVIEWS The completion of the Red Data Book of British Vascular Plants after seven years of work on some 350 of our rarest species allowed these maps, where necessary, to be updated and elsewhere, only obvious errors corrected. Any maps which were altered were given a + sign to the right of the species number in the inset. Similarly, maps altered in the third edition were given an * symbol. Sadly, even the Pteridophytes were not updated by using the more recent maps in the Aflas of ferns of the British Isles (1978) edited by Jermy, Arnold, Farrell & Perring, and the under-recording of Primula vulgaris in Ireland due to an administrative error, subsequently rectified in the Critical supplement, has again been overlooked. This new reprint of the third edition of the Aflas is in a much smaller (and hence more convenient and portable) size, c. 35.0 x 17.5 cm. The work can at last be taken off the table and into the car — even carried in the ruck-sack. The maps are clear and usable. The original transparent overlays (six, each with two maps) of the first and third editions have been reproduced as three ordinary pages. Sadly, in an attempt to keep the cost of the reprint reasonable the reader is left to produce his or her own transparent overlays. However, there is one very important addition: an Index and Bibliography. This has been prepared by C. D. Preston in his usual meticulous and careful way. This welcome addition enables the user to check if a more recent (1962-1989) and up-to-date map has been published elsewhere. We are especially grateful to him for his efforts; for these nine pages alone the work is worth buying. S. L. Jury Guide des fougeres et plantes alliées. R. Prelli. Pp. viii + 232, illustrated. Editions Lechevalier, Paris. 1990. Price not stated (ISBN 2—720-50528-S). This second edition of Prelli’s Guide follows the same pattern as the first, a brief general account of the essential features of the Pteridophyta, followed by a catalogue raisonnée of the species to be found in France (including Corsica). The anatomy and morphology of the pteridophytes are dealt with at an elementary level, and there is little at which to cavil. Not all fern spores however possess a perispore (as is implied). Beginners will search the spores of Pteridium (for example) in vain. Mention might have been made of the manner in which the gametophytes of many ferns pass through male and female phases as they develop. This feature, tending to discourage intragametophytic selfing, may be an important aspect of their population biology. The condensed account of the fossil history of the pteridophytes is not wholly in accord with modern views, although similar will be found in some current textbooks. There is little evidence of any close relationship between Psilotum and Rhynia. Many would now doubt the relevance of the kind of heterospory seen in Selaginella to the evolution of seeds. The reader might assume from the text that all the fern-like fronds found in the Carboniferous were referable to ferns. Many belonged to the all-important pteridosperms (which receive no mention). The systematic section, which is brought into line with current knowledge, forms a useful Flora. Keys are provided for the larger genera. Asplenium is made to include Phyllitis and Ceterach. The analysis of the probable relationships of Dryopteris affinis, D. oreades and D. caucasica (reflecting the views of Fraser-Jenkins and others) is clearly set out. The descriptions of species are frequently supported by excellent drawings, and those of many ferns by photo-silhouettes of the fronds. Unfortunately no magnifications are given. In fact, dimensions are entirely absent. “Spores smaller than in the two other sub-species’’ (following Dryopteris affinis subsp. affinis) is of little use for purposes of identification if no measurements are given for any form. Chromosome numbers are also conspicuously missing, although ploidy is often referred to. According to the Introduction the Guide is intended for all those seeking information about the biology, ecology, evolution and classification of the Pteridophyta. Having mastered the first part of the book, it is presumably hoped that the readers will then proceed into the field to see the living representatives that France has to offer. It is a laudable aim; let us hope it succeeds. P. R. BELL BOOK REVIEWS 443 Wild flower habitats of Hertfordshire; past, present and future. B. Sawford. Pp. xvi + 299 with 16 colour plates. Castlemead Publications, Ware, Herts. Price £18.95 (ISBN 0-948555—09-2). This book by the Senior Keeper of the North Hertfordshire Museum’s Natural History Department sets an excellent pattern that one would like to see repeated for every county in Britain. He discusses first the usual preliminaries — geology, climate and so forth — and then describes in some detail the vegetation of the county’s various habitats. Finally, in a long series of appendices, which occupy almost one-third of the book, he lists the characteristic flora of these habitats. In fact he has produced a most admirable vade-mecum for botanists, both resident and visiting, in a county which nobody could describe as the most botanically exciting in Britain but which nevertheless has its specialities. North Hertfordshire, for instance, is the headquarters of Bunium bulbocastanum in England. The habitat core of the book goes into considerable detail, describing for instance, eight woodland types in the Peterken classification plus, ‘“‘recent woods and plantations”, which most botanists pass swiftly by. And even Mr Sawford cannot offer a great deal more than Sycamore, Ivy, Bracken and Urtica dioica. However, assiduous botanists prepared to watch one plantation for several decades may also hope to log the arrival of Bluebell and Dog’s Mercury. There are also analyses of eight types of grassland and ten types of wetland, with seven more sections on various man-made habitats, from arable fields to walls and churchyards. The whole enterprise, coupled with compiling the lists of characteristic plants of the various habitats, clearly entailed an enormous amount of, no doubt extremely pleasurable, field work. The book is excellent and accurate, but could have been made even more useful by a fuller index. For instance, seeking for information on the typical habitat of Bromus benekenii, which I have actually seen in a Hertfordshire beechwood, I drew a blank, and quailed at checking through all the lists, or even all the likely lists. Likewise I should have welcomed more details of actual sites in the habitat section, though of course there are often good reasons for not mentioning particular sites. A somewhat depressing Appendix is that of the 107 plants now extinct in Hertfordshire, some of them, including Spiranthes spiralis and Cirsium dissectum, having been lost as recently as the early 1980s. One advantage of publishing such lists is that it stimulates people to go and see whether the plants really are extinct — as I shall do with a local population of Senecio fluviatilis, which I saw in 1958, and is here stated to have become extinct c. 1960. In future, when botanising in other counties, I shall regret not having the equivalent of Mr Sawford’s book. So, more please. R. S. R. FItrer Wild flowers of north east Essex. T. Tarpey & J. Heath. Pp. 302, with 7 line illustrations and 659 maps. Colchester Natural History Society. 1990. Price £8 (ISBN 0-9516312-0-9). As one who took part in the final stages of recording for Stanley Jermyn’s Flora of Essex, published in 1974, I welcome this opportunity to find out what has been happening in my native county since then. Jermyn’s flora was the culmination of many years of work and even by the time it was published it was already becoming outdated in this rapidly changing county. Unsurprisingly, this new work is often depressing reading; the phrase, “‘not found during our survey” appears all too frequently. Only north-eastern Essex is covered, but it is covered well. Basically it is a report of a ten-year survey by the Colchester Natural History Society, recording on a 1 km square basis. The coverage has been remarkably thorough, with reasonable species totals from all but a handful of their 1322 squares. Maps of common species look complete, or nearly so; Urtica dioica has been verified from 1252 squares and Crataegus monogyna from 1238. The data storage (over 180,000 records) and map production have been handled by desk-top computer and should serve as an inspiration to those of us struggling to devise similar systems. The bulk of the book consists of the accounts, sometimes brief, sometimes expanded, of all of the species recorded from the area, past or present. Most currently known species are mapped, the iss BOOK REVIEWS maps, up to six to a page, usually being opposite the text entry. Nomenclature follows Clapham, Tutin & Warburg’s Excursion Flora of the British Isles, 3rd edition, and synonyms are given only in the index. The introductory sections include similar 1 km square maps of surface geology, including the complex drift deposits. The accounts of the habitats are detailed and thorough and contain much on how changing management regimes have affected the flora, rarely for the better. Even popular perceptions of ‘green’ issues have caused problems; referring to one of the few remaining heathlands, the authors write, “‘there is much more to do here if the conservation message is to be interpreted as managing a habitat rather than just growing trees.”’ On a lighter note, a mention of how lion dung may have a use in conserving woodland orchids is typical of the snippets of information to be found throughout the book. There is much else to be found here; the remarkable linear map of Cochearia danica as it spreads along the central reservation of the A12 and the description of how moth collectors are threatening Peucedanum officinale are just two examples. Critical taxa have been carefully recorded and I was pleased to see recognition that Poa subcaerulea is not so rare, even in this southern county. Coverage of infraspecific taxa is, at best, sketchy, but I was interested to see the map of Plantago major subsp. intermedia, while their mapping of the two leaf shapes of Lactuca serriola is precisely the sort of exercise I feel should be included in such projects. The account of Taraxacum is substantial and up to date, though I would have liked to have seen more habitat notes. My only real criticism of the text is that there is not always a clear indication of non-native status. The text seems to indicate significantly greater losses of the native flora than is suggested by the list near the end of the book. This is a book that should certainly be owned by every Essex botanist, but it has to interest anyone else concerned with recording schemes or conservation matters. The authors and recorders are to be congratulated. A. J. SILVERSIDE The biogeography of the British Isles — an introduction. P. Vincent. Pp. 315. Routledge, London. 1990. (ISBN 0-415—03470-1 hardback, 0-415-03471—x paperback). Designed as an introductory text, this is an ambitious attempt to interest undergraduates in the interdisciplinary subject of biogeography — the study and interpretation of plant and animal distributions in geographic space. At face value such a definition appears unexceptional but, in geographical as in botanical circles, a chorological approach finds both support and opposition. By placing emphasis on patterns rather than on the processes which generate them, spatial explanations fail to disentangle the separate effects of endogenous and exogenous factors and serve rather to focus attention on the similarities in the ranges of environmental conditions at the cores and limits of distributions. In practice, Vincent achieves a reasonable balance between these competing approaches but, in consequence, the treatment of concepts and methods of enquiry is often partial and superficial. For example, although paleoecological evidence has proved useful for evaluating a number of the concepts used in the book — the causes of disjunct distributions, rates of dispersal, migration and extinction — this evidence itself needs to be applied with care. Likewise the concept of community finds no place in the discussion, even though its contribution to the history of the subject in Britain has proved to be a particularly important one. It is appropriate, for example, to examine the contribution of community ecology and vegetation description and mapping when addressing the history of nature conservation in Britain. These concepts too have played an important role in examining the impact of urban and industrial growth. Both are topics which are addressed in the latter part of the book. A partial approach means therefore, that undergraduate readers, whether in botany, ecology, environmental science or geography, are likely to need further guidance and assistance before they can adequately address the range of topics introduced here. The strength of the book lies in its inclusion of both animal and plant distributions, of marine and terrestrial organisms and environments and in the attention given to the problem of explaining why the distribution of certain plants and animals does not extend to Ireland — the latter a particularly BOOK REVIEWS 445 useful vehicle for exploring many of the central concerns of the book. The book is very fully illustrated, has an extensive glossary and a full index. Had more attention and detail been directed to the development of a rigorous and consistent method of enquiry, then these attractive features would have ensured a wider audience than a reading of the text itself suggests. C. M. HARRISON ve ile op sic Bhp ocapehdiorely Doe mais raletdelehen mere @ 27824 pl Ae sesl re GA fn 6: Salina ~ bre een wee The (art THEA in thy, ie. 9a Mark seat Soieas Pt TS. Ab eat: sib ay hole ps2 MVC Us regimes bare attested e Hota, wel wt | 2 * sired Kave phasis’ | wale oe PocMie ‘ 7 : 1h hunls, Doe eulho write “theta is tach iniey 8 dy here if the Gomera WISE aS ng a hemihar hadier thai jet provekiz | 72 a” Nigh wwe Wt CATT . ww ainsi tr chy is typ te yay i * j 7.7 meghcua Li ‘ “ \ a. is : ; p4 , a - _¥ ‘ J . ¢ mele | Fa wieg* \ M)fLT ri if if eke 2 il co , 4 7 om 7 , fo an (earrintie wheyahampeetmsi ret XS $6 a leis an aed } ; cat | Seen carctulls ha : i . eM rey ved Th + thin «i - as * ied tO Gee anianr, ect V : , uta out a id | oa mn a } : if TArL The : : ¢ list new tof a] Watsonia, 18, 447-452 (1991) 447 Obituaries JOHN FREDERICK GUSTAV CHAPPLE (1911—1990) John Chapple, who died on 4 December 1990, had been no more than a name for over 40 years to all but a handful of the present generation of British botanists. A demanding and immensely successful second career in educational publishing, much of it spent overseas, had so completely taken him from his earlier interests that his obituary in The Times made no more than a passing and almost grudging mention of his one-time involvement in “botanical research at Oxford”. Yet there was probably no one who played a more crucial role in the 1930s in holding together the Botanical Exchange Club after the death of Dr G. C. Druce and ensuring its continuance and gradual transformation into the B.S.B.I. of the post-war years. It was to Druce that Chapple owed his entry on to the botanical stage. Born on 23 December 1911 and brought up in poverty-stricken circumstances, he was taken on by Druce as his assistant after leaving school, relieving that by then elderly and increasingly frail veteran not only of various routine curatorial chores, but even of the burden of his vasculum on their collecting trips into the field. On his death in 1932 Druce bequeathed the larger part of his very substantial fortune to Oxford University on condition that it took over his house in the city with its very extensive herbarium and library, maintained it as an institute of taxonomic botany and appointed Chapple as the Curator, “‘to see that the collections are kept on good order, and to carry on as far as possible work at British Field Botany”. The will further directed that the Curator “should undertake to give adequate assistance to the Botanical Society and Exchange Club, if so desired by that body’. And, for good measure, there was a personal bequest to Chapple of £1000, quite a handsome sum for those days. Unfortunately the will contained legal loopholes and became the subject of a lawsuit between the University and the Executors which dragged on and on, leaving Chapple in a position of continuing uncertainty and, for a time, acute financial difficulty (for he could not be paid out of the estate until probate was granted and that was delayed for two whole years). Only a critical £50 from the Botanical Exchange Club, which Chapple had joined in 1931, enabled him to keep afloat. Even after the dispute was settled and the University took over as his employer, the situation remained by no means satisfactory. Taxonomy was at a very low ebb in the academic world at that period and the new institute was effectively cold-shouldered. Required nonetheless to be permanently on hand, seldom able to escape into the field, Chapple found himself frustrated and dispirited. The one clear gainer was the B.E.C, which, thanks to Druce’s forethought, acquired the services free of charge of a very efficient Executive Secretary. This proved a godsend during the prolonged and strained conversion of Druce’s personal fiefdom into a learned society of ordinary character. Chapple was able to deal with much of the routine correspondence and, with the excellent knowledge he had acquired of British plants and with the Druce Herbarium on hand for reference, field many of the requests for assistance with identification which the Club felt obliged to continue to offer to meet as a major way of attracting and holding members. By 1936 it had become increasingly apparent that no single person could cope with the range of duties that went with the Honorary Secretaryship, and on the death in that year of W. H. Pearsall, Chapple’s role belatedly received official recognition and he was elected into a more narrowly- conceived version of that linchpin office. It was consequently a severe blow to the Club when, only three years later, war broke out and he was called up almost immediately. It was a different B.E.C. and a different state of affairs at the University to which he returned in 1945. Although he resumed the Honorary Secretaryship, the Club’s centre of gravity had decisively shifted to London, where its two new leading (and forceful) figures, A. J. Wilmott and J. E. Lousley, both lived and worked. At Oxford, in the meantime, taxonomy was at last beginning to revive and the Druce Herbarium to be seen as having a function within the Botany School. As a 448 OBITUARIES result Chapple was placed under pressure to equip himself with an academic qualification. The prospect of the years of study that this would have entailed was hardly an appealing one; and it is little surprise that he decided to take up an offer of employment made to him some time before by his wartime commanding officer, who had formed a very high opinion of his abilities. This was K. B. Potter, a director of the old-established publishing house of Longman’s Green, then just embarking on a major expansion Overseas in response to the post-war boom in the English-language textbook market. This new career proved at once entirely to his taste, and before ten years were out he had been promoted to the board. Later, after the Longman Group merged with Penguin Books, his entrepreneurial flair received a culminating accolade in the shape of the chairmanship of the new joint marketing company that resulted. Despite this spectacular climb he nonetheless remained modest and self-effacing throughout, mixing little in trade circles and declining office in the Publishers’ Association. Although he-had felt obliged to resign the Honorary Secretaryship on his change of career in 1947, he lingered on in the world of British botany just sufficiently long to do the B.S.B.I. (as the B.E.C. had by then become) one final, invaluable service. This was as one of the three hard-working members who constituted the historic Advertising Committee, which in the course of the 1950s was largely instrumental in increasing the size of the Society by more than half. It was a highly fitting way in which to have bowed out. As a botanist — as opposed to a botanical administrator — he will be remembered in particular for much useful work on the floras of Berkshire and Oxfordshire, especially in the more critical groups. Anyone who consults the Fielding-Druce Herbarium today cannot but be struck by the extent and range of his contributions, both to it and to field botany locally, during that all-too-short span of years when they jointly benefited from his enthusiasm, care and expertise. D. E. ALLEN ARTHUR ROY CLAPHAM (1904—1990) Born on 24 May 1904, Roy Clapham was educated at the City of Norwich School. He was elected to an Open Scholarship at Downing College, Cambridge in 1922, and gained a double first class degree in Natural Sciences, specialising in Botany. His first research was also at Cambridge, under the foremost plant physiologist of the day, F. F. Blackman. In 1928 he moved to Rothamsted Agricultural Research Station as a crop physiologist, where his interests in sampling techniques and biological statistics were fostered by the renowned statistician R. A. Fisher. In 1930 he moved to Oxford University and held the post of Demonstrator in the Department of Botany until 1944. This was a period when his botanical interests were widened by contact with many leading academic botanists. One major influence was Sir A. G. Tansley, the great plant ecologist, who had also founded and edited one of the leading botanical journals, The New Phytologist. In 1931 he passed the editorship on to Clapham, H. Godwin and W. O. James, who jointly edited the journal for the next 30 years. Roy Clapham’s published work carried out while at Oxford indicates the breadth of his interests and includes papers on quantitative ecology, cytotaxonomy and post-glacial vegetation history. It was also at Oxford that his stress on the importance of synthesis of information began to develop and he played a leading role in the launch of the Biological Flora of the British Isles in 1940 and he wrote, with P. W. Richards, several accounts of Juncus species. An early problem was the lack of an up-to-date list of British vascular plants. Again Roy Clapham demonstrated his scholarship and ability to achieve a consensus with the publication of a Check-list of British Vascular Plants in the Journal of Ecology in 1946. By this time Roy Clapham had moved to the University of Sheffield, where he was Professor of Botany from 1944 until his retirement in 1969. The university benefited from his leadership in many ways. He built up a major Department of Botany and in 1961 founded the Research Unit of Grassland Ecology, supported by the Nature Conservancy. This very successful group still continues in Sheffield as the Unit for Comparative Plant Ecology, supported by the Natural Environment Research Council. He also served as Dean, Pro Vice-Chancellor and, finally, acting Vice- OBITUARIES 449 Ficure 1. Arthur Roy Clapham, 1904-1990. Chancellor for a period in 1965. During all this teaching was never neglected and he continued to lead first-year field excursions into Derbyshire, including Kinder Scout, until the year of his retirement. The post-war years were a period of increasing productivity, culminating in 1952 with the publication, together with the late T. G. Tutin and E. F. Warburg of the Flora of the British Isles, our first completely new Flora this century and the work by which all three, affectionately reduced to the acronym ‘CTW’, are best known to generations of botanists. The Flora and its abbreviated companion, the Excursion Flora, are both now in their third editions; Roy was involved in the preparation of them all, spanning a period of 35 years of publication. In 1950 Roy Clapham, speaking at a conference organised by the B.S.B.I. (which he joined in 1945) on “‘The Study of the Distribution of British Plants”, persuasively laid out the ground rules for the production of distribution maps of British vascular plants based on the 10-kilometre squares of the National Grid, even forseeing the production of transparent overlays; a conference proposal endorsing his suggestions was carried “‘with acclamation’’. So was set in train perhaps the most successful project ever carried out by the B.S.B.I. and certainly the one which involved most members. Roy Clapham was heavily involved in the project’s organisation, which culminated in the publication of the Atlas of the British flora, edited by F. H. Perring and S. M. Walters, in 1962. Subsequently the distributions of many other groups of British plants and animals have been similarly mapped. The 1950s were a time of great importance for the development of nature conservation in Britain. Roy Clapham was deeply committed, being a member of the then Nature Conservancy from 1956 to 1972 and chairing several influential committees during that period. When the Nature Conservancy 450 OBITUARIES came under the direction of the Natural Environment Research Council he was also appointed a council member. From 1964 to 1975 he was also Chairman of the British National Programme of the International Biological Programme. This international project aimed to study the biological basis of productivity and human welfare, by carrying out comparable research in the main ecological areas of the world. Roy Clapham was a natural choice for British Chairman both because of the breadth of his scientific knowledge and his skills in reaching wise decisions from committee discussion. His diplomatic skills were also recognised by the Royal Society; in 1964 he and Professor Blackett, the physicist, visited China as the Society’s representatives and guests of the Academia Sinica in one of the few scientific contacts to be made during the ravages of the Cultural Revolution. Even during these years of national responsibility Roy Clapham did not neglect his home area, helping to found the Derbyshire Wildlife Trust and acting as its first chairman. He also edited the Flora of Derbyshire, published in 1969, and personally wrote ecological notes for every species. It was entirely characteristic that when he came to the draft of Hieracium and found it wanting, he took it upon himself to collect material from a number of Derbyshire localities, identify it himself, and then send it to P. D. Sell at Cambridge for comment. His breadth of ecological knowledge was also put to good use in his authorship of the Oxford book of trees, published in 1975. Roy Clapham’s achievements were recognised by the award of a Fellowship of the Royal Society in 1959, the Presidency of the Linnean Society of London from 1967 to 1970, followed by the award of its Gold Medal in 1972, a C.B.E. in 1969, and an LI.D. from Aberdeen University in 1970. He was also a Trustee of the then British Museum (Natural History) from 1965 to 1975. Throughout his career Roy Clapham remained modest and unassuming, achieving his influence by quiet persuasion and logic. He was a complete scholar, widely interested in science and natural history, but also a first rate linguist with a love of etymology. It was significant that the University of Sheffield awarded him an Honorary Litt.D. in 1970. He relaxed at home, where he and his wife Brenda entertained their many friends, colleagues and family with good food, wine and conver- sation. T. T. ELKINGTON BARBARA MARY STEYNING EVERARD (1910—1990) Barbara Everard (née Beard), the botanical artist, died on 17 June 1990. She was born at Telscombe, Sussex, on 27 July 1910 in the Manor House now owned by the National Trust. Her father was the first cousin of the well-known gardener and author E. A. Bowles (1865-1954) and this may well have inspired her life-long interest in plants and gardening. When I first met her some 17 years ago, she gave me a guided tour of her garden, pointing out enthusiastically the many choice ‘E. A. Bowles’ plants she had collected from all over the country. She joined the B.S.B.I. in 1956 and exhibited her paintings and drawings of plants regularly at the Society’s Annual Exhibition meeting each November. Many members will have purchased her poster of ‘Endangered and vanishing plants’ which she promoted at these events. She was introduced to the Society by Ted Lousley at the time of her being commissioned by the Medici Society to produce 900 illustrations of British wild flowers for a proposed Flora of the British Isles. She had completed this task by 1963, thanks to the many amateur and professional botanists who had sent her flowering plants by post from all over the country. Her son Anthony remembers as many as 15 parcels a day arriving at their Farnham home, the most memorable being a many times folded specimen of Giant Hogweed. Sadly, the Medici Society never published the book. Barbara will probably be best remembered by B.S.B.I. members for her illustrations in Oleg Polunin’s Trees and bushes of Europe (Oxford University Press, 1976). Although reproduced at a much reduced scale from the originals, these drawings and paintings exemplify her work. She produced bold and botanically accurate illustrations that reproduced well and this, together with her ability to work quickly, made her popular with both authors and publishers. Barbara also provided line drawings for Oleg’s other Field Guides to European flowers. Barbara’s main passion in life was for orchids — growing them, painting them and conserving OBITUARIES 451 them. Her introduction to orchids began 50 years ago in Malaya where she joined her late husband Ray, who had taken up a posting there as a rubber planter. From 1938 until World War II intervened in 1942, and again from 1946 until 1950, Barbara lived only a stone’s throw from the tropical rain forest and its orchid riches. By 1950 she had gathered together enough paintings of tropical flowers for her first exhibition, which was held at the Singapore Flower Show. From then on her love of orchids grew apace and in parallel with her reputation as a botanical artist. She won eight Gold Medals for her botanical art from the Royal Horticultural Society, including the prestigious Grenfell Gold Medal in 1950 for a display of paintings of Malayan wild flowers. In 1976, combining her interest in tropical flowers, orchids and a growing awareness of conservation matters, she returned to Malaya on a Winston Churchill Travelling Fellowship to illustrate the threatened plants of Malaya. The original paintings from this fellowship have featured in several books, articles and calendars on threatened plants and, were generously donated by Barbara to the Royal Botanic Gardens Kew, in 1987. There they join Barbara’s original paintings from Brian Morley’s monumental Wild Flowers of the World (Michael Joseph, 1970) which will be a lasting testimonial to her skill and dedication. For many years, Barbara was a leading light in orchid growing circles in the British Isles, originally an active member of the Thames Valley Orchid Society and latterly in the Devon Orchid Society and the Orchid Society of Great Britain. She was a skilled grower and won many prizes for her orchids, the last an Award of Merit from the Royal Horticultural Society for a fine specimen of the Burmese Slipper-orchid, Paphiopedilum charlesworthii. Over the years, she became more and more concerned by the disappearance of the tropical forests and their orchids. Her response to their plight was characteristically positive. Barbara decided that, while she could do little to protect the orchids and their habitats directly, it was possible to protect those same species that were already in cultivation. She had often seen wonderful collections of orchids, lovingly assembled and cared for over decades by their owners, decline or die in a matter of days or weeks when their owner became ill or died. Plants of inestimable value have perished for the want of the owner leaving instructions for their care in the event of an unforseen accident or illness. Barbara, after consultation with friends representing the Orchid Society of Great Britain, other orchid societies, the Royal Horticultural Society and the Royal Botanic Gardens, Kew, found the answer! In 1986 the Barbara Everard Trust for Orchid Conservation was established as a registered charity provided with its initial capital by the generosity of Barbara herself. The Trustees, representatives of the Orchid Society of Great Britain, Kew and the Royal Horticultural Society decided that the Trust should be named after Barbara Everard in gratitude for her generosity and the leading role she had taken in its establishment. The primary aim of the Trust is to conserve orchids in cultivation. One of its main roles is seen to be the establishment of a mechanism by which collections can be rescued in the event of the owner’s death or serious illness. Several thousand plants have already been rescued and will continue to delight future generations. P. J. Crips RICHARD PEARSE LIBBEY (1911—87) During the last five years, Norfolk has lost four of its eminent botanists. Richard Libbey, the authors of the most recent Flora of the county, Charles Petch (1909-87) and Eric Swann (1904-89), and the widely loved all-round naturalist, Ted Ellis (1909-86), the subject of a recent biography (Stone 1988), represented a remarkable generation of field botanists. The late 1980s has been indeed, to use that well-worn but appropriate phrase, the end of an era (vide Daniels 1990). Richard Libbey was born on 9 March 1911, the son of the Rev. H. C. Libbey of Huddersfield, Yorkshire. He was educated at St John’s School, Leatherhead in Surrey, and the Technical College, Huddersfield. He went up to the University of Reading in 1930, where he read Agricultural Botany and graduated, after an absence through illness, with First Class Honours in 1936. In 1936-7 he carried out research on Thames Valley grasslands under the supervision of Professor W. B. Brierley, although he did not pursue this work through to a higher degree. Instead, he followed up 452 OBITUARIES his interest in grasses by joining the staff of the Board of Greenkeeping Research (later the Sports Turf Institute) at Bingley near Leeds in his native Yorkshire. During World War II, after volunteering for active service in 1939, he was appointed to the War Agricultural Committee for Norfolk. By then he had married Alice Clark, another graduate of Reading University. They had three children. After the war, the family returned to Yorkshire, but subsequently moved back to Norfolk, to King’s Lynn, where Richard worked for a company that manufactured fertilizers. He was an enthusiastic member of the B.S.B.I., which he joined in 1943, and he was an important contributor to the Atlas mapping scheme, as well as attending many meetings all over the country especially after his retirement in 1971. Much of his Norfolk field-work was with Eric Swann, and records that they collected together were included in Swann’s Supplement to the Flora of Norfolk (1975). I remember Richard Libbey as a friendly and helpful person, willing to give encouragement to a young, inexperienced botanist, not least in indicating most tactfully a gross error in my first publication. Hé was a member of the Council of the British Bryological Society, and devoted much of the last four years of his life to bryology. His bryophyte herbarium has been lodged at the Castle Museum, Norwich, although his herbarium of Higher Plants, comprising some 12,000 sheets, is now at the University of Leicester herbarium (LTR). As well as having a wide knowledge of grasses, he was the B.S.B.I. referee for the genus Oxalis. REFERENCES DanieELs, E. T. (1990). The end of an era. Trans. Norfolk Norwich Nat. Soc. 28: 336-340. STonE, E. (1988). Ted Ellis, the people’s naturalist. Norwich. J. R. AKEROYD INDEX WATSONIA VOLUME 18 (1990-91) Prepared by C. R. Boon Abbott, P. P. — Rev. of Flora of the East Riding of Yorkshire 323-324 Abies 325 Acacia 328 Acer campestre 364, 388; pseudoplatanus L.) from infructescences, Detection of protandry and protogyny in Sycamore (, 17-20, (illus.) 19, 388 Aceras anthropophorum 45 Achillea millefolium 258, 261; ptarmica 66 Acinos arvensis 76, 79 Aconitum napellus 326, subsp. lusitanicum 326, subsp. napellus 326; napellus X variegatum (v.c. 49) 420 Aconogonon 357 Acorus calamus (v.c. 8) 227 Actaea 76; spicata 74, 79 Adenostyles 274 Adiantum capillus-veneris 231 Adonis 252 Aeschimann, D. & Burdet, H. M. — Flore de la Suisse et des territoires limitrophes (Bk Rev.) 229 Aesculus hippocastanum 181 Aethusa cynapium 329 Aglaophyton 100 Agropyron 249; caninum 186 Agrostemma githago 205, 329 Agrostis curtisii 349; gigantea (v.c. 81) 436; scabra (v.c. 46) 436; vinealis (v.c. 43) 228 Alolon 252 Aira caryophyllea subsp. multiculmis 187, (v.c. 67, 68) 228, (v.c. 77) 436 Akeroyd, J. R. — Anthyllis vulneraria L. subsp. polyphylla (DC.) Nyman, an alien kidney- vetch in Britain 401—403 Akeroyd, J. R. — Obit. of Richard Pearse Libbey (1911-1987) 451-452 Al-Bermani, A.-K. K. A. & Stace, C. A.— A new subspecies of Festuca rubra L. 315-316 Alchemilla alpina 182; filicaulis subsp. vestita (v.c. 46) 218; glomerulans (v.c. 68) 425; mollis (v.c. 35, 49, 69, 73) 425; vestita 87; wichurae (v.c. 72) 425 Aleksandrova, V. D., with D. Love (trans.) — Vegetation of the Soviet polar deserts (Bk. Rev.) 229-230 Alexander, J. C. M., with S. M. Walters et al. — The European garden flora. A manual for the identification of plants cultivated in Europe, both out-of-doors and under glass. Vol. II, Dicotyledons (Part I) (Bk Rev.) 235-236 Alisma 97 Allen, D. E. — Micromorphy in New Forest Rubi 81 Allen, D. E. — The Rubus flora of the Isle of Wight 21-31 Allen, D. E. — Obit. of John Frederick Gustav Chapple (1911-1990) 447-448 Alliaria petiolata 84 Allium 381, 383, 384; carinatum L., A roseum L. and Nectaroscordum — siculum (Ucria) Lindley on St Vincent’s Rocks, Avon Gorge, Bristol, Allium sphaerocephalon L. and introduced, 381-385, (v.c. 77) 226, (v.c. 49) 433; nigrum (v.c. 5) 226; oleraceum 79, 381, (v.c. 26) 433; paradoxum (v.c. 69) 433; roseum L. and Nectaroscordum_ siculum (Ucria) Lindley on St Vincent’s Rocks, Avon Gorge, Bristol, Allium sphaerocephalon L. and introduced A. carinatum L., 381-385; schoenoprasum (v.c. 77, 80) 226, (v.c. 39) 433; scorodoprasum 79; siculum 381-383; sphaerocephalon L. and introduced A. car- inatum L., A. roseum L. and Nectaroscor- dum siculum (Ucria) Lindley on St Vincent’s Rocks, Avon Gorge, Bristol 381-385; triquetrum (v.c. 25) 433; ursinum 117, 185; vineale 381, 384 Alnus incana 179, 184, (v.c. 35) 428 Aloe 328 Alopecurus aequalis (v.c. 5) 436; aequalis xX geniculatus (v.c. 24) 436; alpinus (v.c. 72) 436; bulbosus Gouan in Dorset 206-207; bulbosus X geniculatus 206, (v.c. 5, 54) 436; geniculatus X pratensis (v.c. 14, 29) 228; xX plettkei 206; pratensis 187 Althaea officinalis (v.c. 69) 422 Amaranthus blitoides (v.c. 25) 422; retroflexus (v.c. 4) 216 Ammania boraei 254; borysthenica 254 x Ammocalamagrostis baltica 369 Ammophila arenaria 370, 374-378, 415, var. arundinacea 369; arenaria X Calamagrostis 453 454 INDEX TO WATSONIA VOLUME 18 epigejos 369 Amphibromus neesii 413 Amsinckia calycina 329; micrantha 329 Anacamptis pyramidalis 79 Anagallis arvensis 184; minima (v.c. H36) 221; tenella 184 Anchusa barrelieri 329 Anemonanthea 252 Anemone 252; blanda (v.c. 69) 420; nemorosa 66, 117 Angelica 245 Anon — Biological collections UK. A report on the findings of the Museums Association Work- ing Party on natural science collections based on a study of biological collections in the United Kingdom (Bk Rev.) 108-109 Antennaria dioica 75, 80, (v.c. 72) 430 x Anthemimatricaria maleolens 212 Anthemis arvensis (v.c. 33) 224 Anthoxanthum odoratum 87 Anthriscus caucalis (v.c. 47) 427; sylvestris 84, 183, 415 Anthyllis vulneraria 39, 401, 402, subsp. carpatica 401, 402, subsp. carpatica var. pseudovulner- aria 401, 402, subsp. corbieri 401, subsp. lapponica 401, 402, subsp. polyphylla (DC.) Nyman, an alien kidney-vetch in Britain 401- 403, (v.c. 29) 217, subsp. vulneraria 401, 402 Apera spica-venti (v.c. 7, 8) 228 Aphanes 249 Apium repens 102 Aquilegia 76; vulgaris 79 Arabidopsis thaliana 181 Arabis glabra 205 Arcteranthis 252 Arctium minus 415, subsp. nemorosum 185; pubens (v.c. 67, 68) 431 Arctostaphylos uva-ursi 184, (v.c. 73) 428 Arenaria balearica (v.c. 73) 216; leptoclados 181; serpyllifolia subsp. macrocarpa (v.c. 46) 422 Arisaema triphylla 340 Armeria maritima 116 Armoracia 248; rusticana (v.c. 73) 421 Arnett, R. (ed), with E. Crackles — Flora of the East Riding of Yorkshire (Bk Rev.) 323-324 Arrhenatherum elatius 364, subsp. bulbosum (v.c. 67, 68) 228 Artemisia 261; vulgaris 325, 360 Arum 111, 333, 337, 341; hygrophilum 337; itali- cum 111; maculatum L. — a historical review and new observations. The pollination of, 333-342, 186, 388, (illus.) 334; nigrum 111, 335; pictum 111 Aruncus dioicus (v.c. 69) 423 Asarina 107 Asclepias syriaca 337 Asparagus officinalis subsp. prostratus 97, (v.c. 2) 226 Aspidophyllum 252 Asplenium 442; adiantum-nigrum 180; billotii 319; marinum 104; ruta-muraria 180; scolo- pendrium 388; trichomanes 13, 231, subsp. quadrivalens 180, (v.c. 109) 419 x Asplenophyllitis microdon (T. Moore) Alston in Edinburgh, Herbarium sheets of the rare fern, 319 Aster laevis X novi-belgii (v.c. 39) 224 Astragalus danicus 80 Astrantia major (v.c. 26) 427 Atriplex glabriuscula 181, 206, 320; glabriuscula x longipes 320; X gustafssoniana 320; hali- mus (v.c. 5) 216; longipes 320, (v.c. 35) 216, (v.c. 68) 422; longipes X prostrata 320, (v.c. 67, 68) 422; praecox (v.c. 108) 216, (v.c. 68) 422; X taschereaui Stace, hyb. nov. 320 Atropa belladonna 71, 79, 415 Avenula pubescens 66, 87 Azolla 231; filiculoides (v.c. 26, H36) 215 Bailey, J. P., with J. E. Wentworth & R. J. Gornall — Contributions to a cytological cata- logue of the British and Irish flora 415—417 Baker, A. J. M. — Plants of metalliferous mine workings in the British Isles (Talk) 116-117 Baldellia ranunculoides 97, (v.c. 21) 225 Banks, R. E. R. — Rev. of John Lightfoot: his work and travels, with a biographical intro- duction and a catalogue of the Lightfoot herbarium 437-438 Barbarea stricta (v.c. 25) 216; vulgaris 180, 181; intermedia (v.c. 94) 421 Barbula unguiculata 364 Barneoudia 252 Bartsia alpina 87 Batrachium bachii 144, 412 Beckett, A., with L. C. Frost, L. Houston, L. & C. M. Lovatt — Allium sphaerocephalon L. and introduced A. carinatum L., A roseum L. and Nectaroscordum siculum (Ucria) Lindley on St Vincent’s Rocks, Avon Gorge, Bristol 381-385 Beckett, G. — Obit. of Eric Lister Swann (1904— 1989) 112-113 Beckwithia 252 Bell, P. R. — Rev. of Guide des fougéres et plantes alliées 442 Bellis perennis 325 Benham, P. E. M., with A. J. Gray (eds) - Spartina anglica — a research review (Bk Rev.) 324-325 Berberis 235, 236, 258, 294; aggregata (v.c. 12) 215; darwinii 329 Berg, R., with L. Ryvarde (ed.) & K. Faegri — Norges ville blomster — fra bier og blomster til fro og frukt [Norway’s wild flowers — from bees and flowers to seeds and fruits| (Bk Rev.) 234 | Berula 249; erecta 73, 75, 79 INDEX TO WATSONIA VOLUME 18 455 Beta vulgaris subsp. maritima 181 Betula 231, 298; pendula 329; pubescens 13, 329 Bevan, D. & Rich, T. C. G. — Bevan’s bittercress: the small, white-flowered variant of Carda- mine X fringsii Wirtgen also present in Bri- tain 403-405 Bevan, J. — Hieracium britannicum F. J. Hanb. in Wales 199-200 Biarum davisii 440 Bidens cernua 71, 73, 80 Binggeli, P. — Detection of protandry and proto- gyny in Sycamore (Acer pseudoplatanus L.) from infructescences 17—20 Birks, H. J. B. — Rev. of Norges ville blomster — fra bier og blomster til fro og frukt |Norway’s wild flowers — from bees and flowers to seeds and fruits] 234 Blackmore, S., with W. Punt & G. C. S. Clarke (eds) — The northwest European pollen flora. Vol. 5 (Bk Rev.) 105 Blackstonia perfoliata 39, 66, 415, (v.c. 67) 221 Blamey, M. & Grey-Wilson, C. — The illustrated flora of Britain and northern Europe (Bk Rev.) 97 Blepharis 328; ciliaris 328; dhofarensis 328; linar- laefolia 328 Blysmus compressus (v.c. 42) 434; rufus 186, (v.c. 44) 227 Book Reviews 97-109, 229-237, 323-329, 437- 445 Booth, Evelyn Mary (1897-1988) (Obit. ) 238-239 Borago officinalis 184 Boswellia sacra 328 Botrychium lunaria 117, 180 Bowden, J. K. — John Lightfoot: his work and travels, with a biographical introduction and a catalogue of the Lightfoot herbarium (Bk Rev.) 437-438 Brachyglottis c.v. Sunshine (v.c. 77) 430 Brachypodium sylvaticum 388 Brachythecium rutabulum 388 Braithwaite, M. E. & Long, D. C. — The botanist in Berwickshire (Bk Rev.) 438-439 Brassica juncea (v.c. 38) 215; napus 181; oleracea 272; rapa 181, (v.c. 94) 215; tournefortii (v.c. 38) 215 Brewis, A. — Yellow Ivy Broomrape 81-82 Bridson, D., with L. Forman (eds) — The herbar- tum handbook (Bk Rev.) 324 Briggs, M. — Annual General Meeting (1989) (Rpt) 115-116, (1990) (Rpt) 331-332 Briggs, M. — Obit. of Oliver Buckle (1903-1989) ial Briggs, M. — Rev. of Flowering plants of Sey- chelles 105-106 Briggs, M. — Rev. of The illustrated flora of Britain and northern Europe 97 Briggs, M. — Sussex plant atlas: selected supple- ment (Bk Rev.) 439 Briza maxima (v.c. 75) 435; media 87 Bromus arvensis (v.c. 33) 228; benekenii 443; carinatus (v.c. 5, 12) 228, (v.c. 28, 51) 436; diandrus (v.c. 77, 93) 435; erectus 66, 364; hordeaceus subsp. ferronii 272, subsp. hor- deaceus 186; inermis (v.c. 5) 228, (v.c. 67) 435; ramosus 186; stamineus 413; sterilis (v.c. 93) 228; tectorum (v.c. 11, 42) 435; valdivia- nus 413 Bryum 364; pseudotriquetrum 201 Buckle, Oliver (1903-1989) (Obit.) 111 Buddleja davidii 329 Bunias orientalis (v.c. 77) 421 Bunium bulbocastanum 443 Bupleurum falcatum (v.c. 35, 64) 427; rotundifo- lium 75, 76, 80 Burdet, H. M., with D. Aeschimann — Flore de la Suisse et des territoires limitrophes (Bk Rev.) 229 Burges, N. A. — Rev. of The contented botanist. Letters of W. H. Harvey about Australia and the Pacific 98-99 Burkhardt, F. & Smith, S. (eds) — The correspon- dence of Charles Darwin. Vol. 4, 1847-1850 (Bk Rev.) 98 Burton, R. M. — Rev. of Biological collections UK. A report on the findings of the Museums Association Working Party on natural science collections based on a study of biological collections in the United Kingdom 108-109 Butomus umbellatus 80 Buxus sempervirens 182 Caesalpina sappan 326 Calamagrostis epigejos 187, 369, 370, 374-376; stricta (v.c. 75) 436 Calamintha ascendens 75, 80 x Calammophila baltica (Fligge) Brand, in the British Isles, The distribution of hybrid Marram Grass, 369-379, (map) 371, (map) 373, var. subarenaria 369 Calla palustris (v.c. 39) 227 Callitriche autumnalis 73; brutia 415, (v.c. 46) 427; hamulata 183; hermaphroditica 80, (v.c. 53) 220, (v.c. 38) 426; intermedia subsp. hamulata 183; obtusangula (v.c. 73) 426; platycarpa 415; stagnalis 183; truncata (v.c. 53) 220 Calluna 214, 217, 441; vulgaris 89, 407 Caloplaca 113 Caltha 248; palustris 34 Calystegia 249; pulchra 184; pulchra X sepium (v.c. 54) 429; sepium 184; silvatica 329; solda- nella 270 Campanula glomerata 39; lactiflora (v.c. 79) 223; latifolia 74, 185; patula (v.c. 39) 223; trache- lium 415, (v.c. 73) 223 Camus, J. M. — Rev. of Ferns: their habitats in the British and Irish landscape. (A natural history 456 INDEX TO WATSONIA VOLUME 18 of Britain’s ferns.) 103-104 Cannabis sativa 260 Capethia 252 Capsella bursa-pastoris 181 Cardamine 201, 403, 404; amara L.: its occur- rence in montane habitats in Br tain 200-201; flexuosa 403, 404; flexuosa X pratensis 404, (v.c. 38) 216; x fringsii Wirtgen also present in Britain, Bevan’s bittercress: the small, white-flowered variant of, 403405; x haussknechtiana 404; hirsuta 181; pratensis 201, 403, 404. Cardaria draba (v.c. 93) 421 Carduus acanthoides 415; nutans 271, (v.c. 103) 224 Carex 233, 246, 407; acuta 221; acuta X acutifor- mis (v.c. 25) 434; acuta X elata (v.c. 6) 227; acutiformis X riparia (v.c. 25) 434; appro- pinquata Schumacher (C. paradoxa Willd.) in Great Britain and Ireland, The dis- tribution of, 201-204, (v.c. 45) 227; arenaria (v.c. 47) 435; bigelowii 186, 233, (v.c. 47, 72) 434; capillaris 87; curta 186; curta X panicu- lata (v.c. 45) 227; demissa 233; diandra 202, 203, 227; dioica 407; dioica X echinata 406; distans 233; disticha 203; divisa 207; echinata 407; elongata (v.c. 47) 227, (v.c. 12) 435; extensa 186; flacca 87; X gaudiniana Guth- nick — a rarity? 406-407; hostiana 233; lepido- carpa 154, 186, 234; limosa (v.c. 50, 81) 227, (v.c. 50) 434; microglochin 224; muricata 233, subsp. muricata 234; ornithopoda 87; otrubae (v.c. 77) 435; pallescens (v.c. 81) 434; paniculata 186, 201-203, 323; paniculata x remota (v.c. 35, 47, 96) 435; paradoxa Willd.) in Great Britain and Ireland, The distribution of Carex appropinquata Schu- macher (, 201-204; pendula (v.c. 86) 227; pilulifera 233; pseudoparadoxa 202; remota 186; rostrata 201, 434; rostrata X vesicaria (v.c. 46) 227, (v.c. 47) 434; serotina 186; spicata 66, 233; stellulata 407; strigosa (v.c. 69) 434; sylvatica 186, 388; tomentosa (v.c. 7) 227; vesicaria 186; viridula subsp. viridula 186, (v.c. 39) 227; vulpinoidea (v.c. 77) 435 Carum carvi (v.c. 38) 427; verticillatum (v.c. 35, 96) 427 Castanospermum australe 440 Catabrosa aquatica (v.c. 76, 80) 228, subsp. aquatica (v.c. 75) 435, subsp. minor 186, (v.c. 75) 435 Caucalis platycarpos 75, 80 Cedrus atlantica var. glauca 92; libani subsp. atlantica f. glaucissima P. D. Sell. nom. nov. 92, subsp. deodara (D. Don) P. D. Sell, comb. nov. 92, var. glauca 92 Centaurea 229, 253, 258, 267; cyanus 325; mon- tana 205, (v.c. 99, 109) 431; nigra 87, 267, 364, 415; scabiosa 248, 362, 364 Centaurium capitatum 272; littorale 205; pulchel- lum (v.c. 68) 221 Cephalanthera longifolia 73, 80, 186 Cerastium arvense (v.c. 70) 422 Ceratochloa staminea (E. Desv.) Stace, comb. nov. 413 Ceratophyllum demersum (v.c. 83, H36) 215; submersum (v.c. 33) 420 Ceterach 442; officinarum (v.c. 62) 419 Chaloner, W. G., with E. M. Friis & P. R. Crane (eds) — The origins of Angiosperms and their biological significance (Bk Rev.) 99 Chamaemelum nobile 80, 415 Chamerion angustifolium 183 Chapman, G. T. L., & M. N. Tweddle (eds), W. Turner — A New Herball (Bk Rev.) 107-108 Chapple, John Frederick Gustav (1911-1990) (Obit.) 447-448 Chater, A. O. — Rev. of Colour identification guide to the grasses, sedges, rushes and ferns of the British Isles and north-western Europe 233-234 Chatters, C. — The status of Pulicaria vulgaris Gaertner in Britain in 1990 405-406 Chenopodium 248; album 181; botrys (v.c. 6) 422; ficifolium (v.c. 38) 422; rubrum 329, (v.c. 43) 422 Chernov, Y.I., with D. L6ve (trans.) — The living tundra (Bk Rev.) 229-230 Chrysanthemum leucanthemum 325; maximum 89; segetum 325; X superbum 89 Chrysosplenium oppositifolium 183, 201 Circaea alpina 80; X intermedia 80; lutetiana 183, 388, (v.c. 94) 220 Cirsium 249; arvense 265; dissectum 243, 443; eriophorum 79, 265; helenioides 185; heter- ophyllum 185, 265; vulgare 360, 364 Clapham, Arthur Roy (1904-1990) (Obit.) 448- 450, (plate) 449 Clark, J. W. & Jermy, A. C. — Additions to the flora of Mull and adjacent small islands 179- 187 Clarke, G. C. S., with W. Punt & S. Blackmore (eds) — The northwest European pollen flora. Vol. 5 (Bk Rev.) 105 Clematis 248, 252; vitalba 360, 364, 384, 415 Clematopsis 252 Clinopodium vulgare 364 Cochlearia (Cruciferae), A new combination in, 82; danica 181, 444; officinalis 201, subsp. officinalis 181, B alpina 82; pyrenaica 82, 116, subsp. alpina (Babington) Dalby, comb. nov. 82 Coeloglossum viride 39, 72, 79, 232, (v.c. 79) 226, (v.c. 25) 433 Colchicum autumnale 79 Commiphora 328 Conium maculatum 183 Conolly, A. P. — Polygonum lichiangense W. INDEX TO WATSONIA VOLUME 18 457 Smith: rejected as a naturalized British spe- cies 351-358 Convallaria majalis 71, 79 Convolvulus 249 Conyza albida 329; bilbaoana 329; canadensis 329; parva 329; sumatrensis 329 Cooksonia 100 Corallorhiza trifida 307 Corner, R. W. M. — Cardamine amara L.: its occurrence in montane habitats in Britain 200-201 Cornus alba 33; sanguinea 33, 360; sericea L. in Ireland: an incipient weed of wetlands 33—36, (v.c. 73) 427, var. flaviramea 35; stolonifera 33 Coronopus squamatus 181 Corydalis claviculata 181, (v.c. 77) 215 Corylus 388; avellana 231, 388 Cotoneaster, Two new species of, 311-313; Section Cotoneaster Subsection Adpressi Series Adpressi 312, 313; ascendens 312; atropurpureus Flinck & Hylm6, sp. nov. 311, 312; bullatus (v.c. 77) 218; dielsianus (v.c. 12, 77) 218; divaricatus (v.c. 29, 77) 219; franche- tli (v.c. 77) 219; frigidus (v.c. 35) 426; hjelmqvistii Flinck & Hylm6, sp. nov. 312, 313; horizontalis 311-313, (v.c. 77, 79) 218, (v.c. 81) 426, var. perpusilla 312, var. perpu- sillus 311, var. prostratus 311, var. wilsonii 312, cv. Coralle 312, cv. Robusta 312; mic- rophyllus 182, (v.c. 39) 218; perpusillus 311; Prostratus 311; racemiflorus 312; rehderi (v.c. 70) 426; sanguineus (v.c. 77) 219; simonsii (v.c. 77) 218; sternianus (v.c. 70) 426; X watereri (v.c. 12, 77) 219 Cotula coronopifolia (v.c. 39) 224 Crackles, E., with R. Arnett (ed.) — Flora of the East Riding of Yorkshire (Bk Rev.) 323-324 Crackles, F. E. — Hypericum xX _ desetangsii Lamotte nm. desetangsii in Yorkshire, with special reference to its spread along railways 63-67 Crambe 248; maritima (v.c. 45) 216 Crane, P. R., with E. M. Friis & W. G. Chaloner (eds) — The origins of Angiosperms and their biological significance (Bk Rev.) 99 Crassula helmsii 439, (v.c. 46, 47, 53, 54) 219, (v.c. 28) 426; tillaea (v.c. 2) 219, (v.c. 12) 426 Crataegus 49-51, 56, 57, 60, 61; laevigata (Poiret) DC. in the Upper Thames Valley, England, Introgressive hybridization between Cratae- gus monogyna Jacq. and, 49-62, subsp. /aevi- gata 49; monogyna Jacq. and C. laevigata (Poiret) DC. in the Upper Thames Valley, England, Introgressive hybridization between, 49-62, 360, 364, 388, 443, subsp. nordica 49 Cratoneuron commutatum 201; filicinum 201 Crepis 85, 259, 268, 269; biennis (v.c. 2) 225; capillaris 185; foetida 329; mollis 85; palu- dosa 74, 79, (v.c. 35) 432; praemorsa (L.) Tausch, new to western Europe 85-87, (illus.) 86, subsp. praemorsa 85; setosa (v.c. 5, 11, 77) 432; virens 271 Cribb, P. J. — Obit. of Barbara Mary Steyning Everard (1910-1990) 450-451 Crithmum maritimum 206 Crocosmia pottsii (v.c. 73) 433 Crocus nudiflorus (v.c. 73) 433 Cronk, Q. C. B. — Rev. of Plant names of medieval England 325-326 Cronk, Q. C. B. — Rev. of Popular medicine in thirteenth century England 325-326 Cryptogramma crispa 180 Cryptomeria japonica subsp. sinensis (Miq.) P. D. Sell, stat. nov. 92, var. sinensis 92 Cuscuta europaea (v.c. 12) 222 Cynodon dactylon 415 Cynoglossum officinale 79 Cynoglottis barrelieri 329 Cynorkis kirkii 105 Cynosurus cristatus 87 Cyperus longus (v.c. 25) 227, (v.c. 35) 434 Cystopteris dickieana 104; fragilis 180, (v.c. 7) 214 Cytisus 274; scoparius 263, subsp. maritimus 263, subsp. scoparius 182, 263; striatus (v.c. 25) 423 Dactylis glomerata 87 Dactylorchis traunsteineri 170; traunsteinerioides 170 Dactylorhiza, Two new varieties of British, 307— 309, 243, 396; x dinglensis 396, nothosubsp. dinglensis 396, nothosubsp. robertsii F. Horsman, nothosubsp. nov. 395-397, (illus. ) 398; francis-drucei 243; fuchsii (Druce) Sod, Differential pollination efficiency within a hybrid swarm between Dactylorhiza purpur- ella (T. & T. A. Stephenson) So6 and, 391- 393039; 66, 1544 1553186,0307-4308% 395, subsp. fuchsii 396, var. rhodochila D. M. T. Ettlinger, var. nov. 308; fuchsii X incarnata (v.c. 41) 433; fuchsii X incarnata subsp. incarnata (v.c. 64) 226; fuchsii X incarnata subsp. pulchella (v.c. 64) 226; fuchsii xX incarnata subsp. pulchella X_ traunsteineri (v.c. 62) 433; fuchsii X praetermissa (v.c. 46) 433; incarnata 79, 155, 170, 307, subsp. cocci- nea 186, subsp. coccinea X purpurella subsp. majaliformis (v.c. 110) 434, subsp. cruenta 160-162, 164, 168-170, 243, subsp. incarnata 154, 186, subsp. pulchella 154, 160-162, 164, 168-170, 186, subsp. pulchella XxX majalis subsp. cambrensis (v.c. 46) 226, subsp. pul- chella X praetermissa (v.c. 46) 226; insularis var. bartonii 307; lapponica 103, 163, 243; maculata 154, 155, 396, subsp ericetorum 307, 395, 396, (illus.) 398, subsp. ericetorum 458 Xx praetermissa (v.c. 42) 433, subsp. erice- torum X purpurella subsp. majaliformis (v.c. 110) 433, var. concolor 307, 308; maculata x praetermissa 396; majalis 154, 164, 170, 391, 395, 396, subsp. alpestris 396, subsp. cam- brensis 395-397, (illus.) 398, subsp. cambren- sis X maculata subsp. ericetorum 395-397, subsp. occidentalis 154, 156, 160-162, 164, 168, 169, 396, 434, subsp. occidentalis var. cambrensis 396, subsp. occidentalis X macu- lata subsp. ericetorum 396, subsp. prae- termissa 154,.155, 160-162, 164, 166-169, subsp. purpurella 154, 160-162, 164, 166— 169, 307, 308, 391, subsp. purpurella var. atrata A. J. Richards, var. nov. 308, subsp. scotica 396, subsp. traunsteinerioides 154, 170, subsp. traunsteinerioides var. eborensis 162, 170, subsp. traunsteinerioides var. fran- cis-drucei 170, subsp. traunsteinerioides var. traunsteinerioides 170; praetermissa 395; praetermissa X purpurella (v.c. 64) 434; pur- purella (T. & T. A. Stephenson) So6 and D. fuchsii (Druce) Sod, Differential pollination efficiency within a hybrid swarm between, 391-393, 327, 395, 396, (v.c. 43) 226, (v.c. 41) 434, subsp. majaliformis (v.c. 105, 110) 434; purpurella x Gymnadenia conopsea (v.c. 105) 434; russowii 163; sambucina 307, var. rubra 307; X townsendiana 396; traun- steineri (Sauter) Soo in the British Isles and a comparison with others from Continental Europe. An assessment of populations of, 153-172, 243, 395, (map) 155, (v.c. 45) 434, subsp. curvifolia 164, subsp. francis-drucei 170, subsp. hibernica 170, subsp. traunstei- nerioides 170; traunsteinerioides 164, 170 Dalby, D. H. — A new combination in Cochlearia (Cruciferae) 82 Daphne laureola79, (v.c. 43) 219; mezereum (v.c. 43, 44) 219 Darmera peltata (v.c. 77) 426 Daucus 274; carota subsp. gummifer 270, 272 David, R. W. — Carex X gaudiniana Guthnick —a rarity? 406—407 David, R. W. — The distribution of Carex appro- pinquata Schumacher (C. paradoxa Willd.) in Great Britain and Ireland 201—204 Dawson, H. J. — Chromosome numbers in two Limonium species 82-84 Deschampsia cespitosa 186, 388 Desmazeria marina (v.c. 47) 435; rigida (v.c. 47) 228 Desmond, R. — Rev. of A New Herball 107-108 Dianthus deltoides 80, (v.c. 75) 422 Diaz, A., with A. J. Lack — The pollination of Arum maculatum L. — a historical review and new observations 333-342 Dicentra formosa 181 Dietrich, W. — The status of Oenothera cambrica INDEX TO WATSONIA VOLUME 18 Rostanski and O. (Onagraceae) 407-408 Digitalis purpurea 415 Diphasiastrum alpinum 180, (v.c. 81) 419; com- planatum (v.c. 96) 214, subsp. issleri 243 Diplachne muelleri 413 Diplotaxis erucoides (v.c. 28) 421; tenuifolia (v.c. 38) 421 Dipsacus fullonum 185, subsp. sylvestris 360; pilosus (v.c. 41) 224 Dony, J. G. — Obit. of John Campbell Gardiner (1905-1989) 239-240 Doronicum columnae X pardalianches X planta- gineum 321; X excelsum (N. E. Br.) Stace, comb. et stat. nov. 321; plantagineum var. excelsum 321 Draba muralis (v.c. 67) 421 Drosera anglica 73, 80, (v.c. 72) 426; intermedia 73, 79; rotundifolia 73, 79, 164 Dryopteris 231; aemula (v.c. 11) 419; affinis 104, 231, 442, (v.c. 53) 214, subsp. affinis 442, (v.c. 75) 214, subsp. affinis x filix-mas (v.c. 11) 419, subsp. borreri (v.c. 75) 214, subsp. cambrensis (v.c. 41, 75) 214, (v.c. 41) 419; assimilis 13; carthusiana 231, (v.c. 93) 214; caucasica 442; dilatata 13, 231; expansa 13; filix-mas 11, 180, 388; oreades 442 Ducker, S. C. (ed.) — The contented botanist. Letters of W. H. Harvey about Australia and the Pacific (Bk Rev.) 98-99 Dudman, A. — Obit. of Christopher C. Haworth (1934-1989) 240-242 Dunn, A. J. — Further observations on Stachys germanica L. 359-367 Dunster, C. W., with G. I. C. Ingram - A nonresupinate abnormality in Orchis pur- purea Hudson 317-319 novae-scotiae Gates Echinops exaltatus (v.c. 25, 26) 431 Edmondson, J. R. — Rev. of Flore de la Suisse et des territoires limitrophes 229 Edmondson, J. R. — Rev. of Honey identification 106 Edmondson, J. R. — Rev. of The Davis & Hedge Festschrift 439-440 Edmondson, J. R., with H. Hofmann & G. Halliday — The Cumbrian herbarium of W. H. Youdale 204-206 Elatine hexandra (v.c. 47) 422 Eleocharis 248; uniglumis 186 Eleogiton fluitans (v.c. 61) 434 Elias, T. S., with K. Tan (ed.) & R. R. Mill— The Davis & Hedge Festschrift (Bk Rev.) 439-440 Elkington, T. T. — Obit. of Arthur Roy Clapham (1904-1990) 448-450 Elkington, T. T. — Rev. of Comparative plant ecology. A functional approach to common British species 230-231 Elkington, T. T. — Rose End Meadows Nature INDEX TO WATSONIA VOLUME 18 459 Reserve, Derbyshire (Fld Mtg Rpt) 117 Ellis, R. G. & Perry, A. R. — Obit. of Arthur Edward Wade (1895-1989) 113-114 Elodea 176; canadensis Michx by Elodea nuttallii (Planch.) H. St John in the British Isles, Displacement of, 173-177; nuttallii (Planch. ) H. St John in the British Isles, Displacement of Elodea canadensis Michx by, 173-177, 439, (v.c. 26, 46, 67, H36) 225 Elymus 249; caninus 186; farctus 270; pycnanthus Xx repens (v.c. 46) 436 Empetrum hermaphroditum 184, (v.c. 73) 428; nigrum 71, 75, 79 Epilobium alsinifolium 201; anagallidifolium 201; angustifolium 183; ciliatum 179, 183, 329; hirsutum 183; obscurum 183; obscurum xX roseum (v.c. 67) 220; parviflorum 183; tetragonum subsp. tetragonum (v.c. 67) 219 Epipactis leptochila (v.c. 8) 226; palustris 87, 154, 163, 164; phyllanthes (v.c. 6) 226; purpurata 261; youngiana 103 Epipogium aphyllum 103 Equisetum fluviatile 180; fluviatile x palustre (v.c. 73) 419; hyemale 180, (v.c. 39) 214; palustre x telmateia (v.c. 52, 72) 214; pratense 327 Erica biformis 93; ciliaris 14, subsp. mackaiana 93; cinerea 14; lusitanica 329; mackaiana Bab., The typification and nomenclature of, 92-93, 14; mackaii 93; tetralix 164, subsp. mackaiana 93, subsp. mackaii 93; tetralix Xx ciliaris 441; vagans 14, 329; X watsonii 441 Erigeron mucronatus 111 Erinus alpinus (v.c. 39) 429 Eriophorum gracile (v.c. 41) 434; latifolium 164, (v.c. 44, 50) 227, (v.c. 50) 434 Erodium cicutarium subsp. cicutarium 415 Erucastrum gallicum (v.c. 69) 420 Eryngium maritimum 183, 270 Erysimum cheiranthoides 76, 79 Ettlinger, D. M. T. —- Two new varieties of British Dactylorhiza 307-309 Eucalyptus 329 Eulophia guineensis 328 Euonymus europaeus 74, 79, 360, 415 Eupatorium cannabinum 185 Euphorbia amygdaloides subsp. robbiae (v.c. 39) 427; corallioides (v.c. 35) 427; cyparissias X esula (v.c. 26) 427; dulcis 183; esula x wald- steinii (v.c. 39) 220; lathyrus 205; peplus 97; platyphyllos (v.c. 38) 220; serrulata (v.c. 5) 220; waldsteinii (v.c. 77) 427 Euphrasia 97, 249, 348, 429; Section Euphrasia Subsection Euphrasia Series Euphrasia 350; Section Semicalcaratae 349; Section Semical- caratae Subsection Ciliatae Series Grandif- lorae 350; Section Semicalcaratae Subsection Ciliatae Series Hirtellae 350; Series Euphra- sia 349; anglica 344, 348, 349, var. graciles- cens 348; arctica subsp. borealis (v.c. 12) 222, subsp. borealis * nemorosa (v.c. 12) 223, (v.c. 35) 429, subsp. tenuis 347; brevipila 347; campestris 349; confusa (v.c. 25) 222; confusa x micrantha (v.c. 73) 429; confusa X nemor- osa (v.c. 35) 429; confusa X scottica (v.c. 77) 222; curta 343-345, var. glabrescens 345; fennica 344, 346, 347; foulaensis (v.c. 98) 222; frigida 346; hirtella 344, 350; micrantha 349; micrantha X nemorosa (v.c. 12) 222; micrantha X scottica (v.c. 42) 429; montana 347, 348; nemorosa 343-345, 349, var. cal- carea 349, var. curta 345; officinalis L. and its nomenclatural implications, The identity of, 343-350, subsp. anglica (Pugsl.) Silverside, comb. et stat. nov. 348, 349, subsp. monticola Silverside, nom. nov. 347, subsp. officinalis 346, subsp. rostkoviana 347-349, var. mon- tana 347, 348; officinalis X nemorosa 349; officinarum 347, onegensis 344, 346; ostenfel- dii 345; pratensis 347, 348; pseudokerneri (v.c. 46) 429; rivularis 349; rostkoviana 343- 350, subsp. fennica 346, 347, subsp. montana 346, 348, subsp. rostkoviana 344, 346, var. fennica 347, subvar. montana 348; scottica 350, (v.c. 35) 222, (v.c. 83) 429; stricta 343, 345, 347, 349; tricuspidata 343; uliginosa 348; vigursil 349 Eurhynchium praelongum 364; swartzii 388 Everard, Barbara Mary Steyning (1910-1990) (Obit.) 450-451 Fallopia aubertii (v.c. 42, 51) 427 Farrell, L. — Rev. of A guide to the wild orchids of Great Britain and Ireland 103 x Fatshedera 273 Fatsia 273 Festuca 249, 250; arundinacea 186, 414, Schreb. var. strictior (Hack.) K. Richt. 414415, subvar. strictior 414; arundinacea X gigantea (v.c. 42) 435; arundinacea X pratensis (v.c. 38) 435; diffusa 315; filiformis (v.c. 75) 228; gigantea 388; guesifalica (v.c. 54) 228; hetero- malla 315; heterophylla (v.c. 35, 73) 435; Juncifolia 315; nigrescens 315, (v.c. 77) 227, subsp. nigrescens (v.c. 2) 435; ovina 87; richardsonii 315; rubra L., A new subspecies of, 315-316, 87, subsp arctica 315, subsp. multiflora 315, subsp. scotica S. Cunn. ex Al- Bermani, subsp. nov. 315, var. commutata 315; tenuifolia 186; vivipara 234 Ficus 236; carica 84, 117, 179, 184 Filago lutescens (v.c. 29) 430 Filipendula 249;ulmaria 11, 66, 415; vulgaris 66, IDR (act 5) 217, Fissidens taxifolius 364 Fitter, R. S. R. — Rev. of Wild flower habitats of Hertfordshire; past, present and future 443 FitzGerald, R. — Obit. of Evelyn Mary Booth (1897-1988) 238-239 460 INDEX TO WATSONIA VOLUME 18 Flinck, K. E. & Hylm6, B. — Two new species of Cotoneaster 311-313 Foley, M. J. Y.— An assessment of populations of Dactylorhiza traunsteineri (Sauter) Sod in the British Isles and a comparison with others from Continental Europe 153-172 Foley, M. J. Y. — The current distribution and abundance of Orchis ustulata L. in southern England 37-48 Forman, L. & Bridson, D. (eds) — The herbarium handbook (Bk Rev.) 324 Foster, A. S., with E. M. Gifford — Morphology and evolution of vascular plants (Bk Rev.) 100 Fragaria vesca 414 Frangula alnus 79 Frankenia laevis 415 Fraxinus excelsior 387, 388 Friis, E. M., Chaloner, W. G. & Crane, P. R. (eds) — The origins of Angiosperms and their biological significance (Bk Rev.) 99 Frost, L. C., Houston, L., Lovatt, C. M. & Beckett, A. — Allium sphaerocephalon L. and introduced A. carinatum L., A. roseum L. and Nectaroscordum siculum (Ucria) Lindley on St Vincent’s Rocks, Avon Gorge, Bristol 381-385 Fuchsia 248; excorticata 248; procumbens 248 Fumaria bastardii 181, (v.c. 73) 420, var. major 92; capreolata 80, (v.c. 11) 420; muralis subsp. boraei 181, (v.c. 38) 215; subsp. boraei var. major (Boreau) P. D. Sell, comb. nov. 92; officinalis 181; magellanica 183 Futter, K. & Raynes, P. — The flora of Derby (Bk Rev.) 99-100 Fegri, K., with L. Ryvarde (ed.) & R. Berg — Norges ville blomster — fra bier og blomster til fro og frukt [Norway’s wild flowers — from bees and flowers to seeds and fruits| (Bk Rev.) 234 Gagea lutea 71, 75, 79, (v.c. 75) 432 Gale, B. A. — Langham Ponds, Surrey (Fld Mtg Rpt) 332 Gale, B. A. — London University Botanic Gar- dens, Surrey (Fld Mtg Rpt) 332 Gale, B. A. — Silwood Park, Berkshire (Fld Mtg Rpt) 332 Galeobdolon argentatum 329; luteum cv. Variega- tum 329 Galeopsis bifida (v.c. 109) 430; ladanum 80 Galinsoga parviflora 329 Galium album 185; aparine 416; cruciata (v.c. 93) 430; debile 329, (v.c. 8) 223; elongatum 13; mollugo 266, 364, 416, (v.c. 79) 223; palustre 13; spurium 75, 76, 80; uliginosum 73, 79, 185; verum 266, 364 Gampsoceras 252 Gardiner, John Campbell (1905-1989) (Obit.) 239-240 Garnock-Jones, P. J., with C. J. Webb & W. R. Sykes — Flora of New Zealand, Volume IV. Naturalised Pteridophytes, Gymnosperms, Dicotyledons (Bk Rev.) 328-329 Gaultheria shallon (v.c. 70) 428 Genista 274; anglica (v.c. 80) 217; tinctoria 263, (v.c. 45) 217, subsp. tinctoria (v.c. 45) 423 Gentiana 247; lutea 247; nivalis 247; pneumo- nanthe 73, 80; verna 327 Gentianella 39, 247; amarella 247; subsp. septentrionalis (v.c. 64) 428; anglica (v.c. 5) 221; campestris 73, 80, 184, (v.c. 12) 221 Geranium macrorrhizum (v.c. 39) 216; pyrenai- cum 79, (v.c. 72) 422; robertianum 388, subsp. celticum (v.c. 99) 422; rotundifolium (v.c. 50) 422; sanguineum 80, 87 Geum 248; urbanum 182, 416 Gifford, E. M. & Foster, A. S.— Morphology and evolution of vascular plants (Bk Rev.) 100 Gilbert, O. L. — Wild figs by the River Don, Sheffield 84-85 Gilbert, O. L. — The vegetation of the River Don, Sheffield (Talk) 117 Gledhill, D. — The names of plants. 2nd ed. (Bk Rev.) 100-101 Glyceria 203; declinata 186; plicata (v.c. 2) 227 Gornall, R. J. — Rev. of A revision of the tribe Antirrhineae 107 Gornall, R. J., with J. E. Wentworth & J. P. Bailey — Contributions to a cytological cata- logue of the British and Irish flora 415-417 Gosler, A. G. —- Introgressive hybridization between Crataegus monogyna Jacq. and C. laevigata (Poiret) DC. in the Upper Thames Valley, England 49-62 Graham, G. G. & Primavesi, A. L. — Notes on some Rosa taxa recorded as occurring in the British Isles 119-124 Grange, W. — Rev. of The flora of Derby 99-100 Gray, A. J. & Benham, P. E. M. (eds) — Spartina anglica — a research review (Bk Rev.) 324— 425 Gray, A. J., with A. F. Raybould, M. J. Law- rence & D. F. Marshall — The origin and taxonomy of Spartina < neyrautii Foucaud 207-209 Gray, A. J., with J. R. Rihan- The distribution of hybrid Marram Grass, * Calammophila bal- tica (Fligge) Brand, in the British Isles 369— 379 Grey-Wilson, C., with M. Blamey - The Illustrated flora of Britain and northern Europe (Bk Rev.) 97 Grime; J. P.; “Hodgson, 'Jc'Gr &- Hunt) RK. — Comparative plant ecology. A functional approach to common British species (Bk Rev.) 230-231 Gulliver, R. L. — The rare plants of the Howar- INDEX TO WATSONIA VOLUME 18 dian Hills, North Yorkshire, 1794-1988 69-— 80 Gymnadenia 317; conopsea (L.) R.Br., A new subspecies ) Of) 2319-320) 9390079087; '317, subsp. borealis (Druce) F. Rose, comb. et stat. nov. 319, (L.) R.Br. subsp. conopsea, Abnormal flowers in Platanthera chlorantha (Custer) Reichenb. and, 316-317, 320, subsp. densiflora 317, (v.c. 105) 433, var. borealis 320, var. densiflora 44, var. insuli- cola 320; odoratissima 320 Gymnocarpium dryopteris 214, 245, (v.c. 28) 419; robertianum (v.c. 96) 214 Habenaria gymnadenia var. borealis 319 Haeupler, H. & Schénfelder, P. — Atlas der Farn- und Bliitenpflanzen der Bundesrepublik Deutschland (Bk Rev.) 101-102 Halerpestes 252 Halliday, G. — Crepis praemorsa (L.) Tausch, new to western Europe 85-87 Halliday, G. — C. C. Haworth’s contribution to the Flora of Cumbria 242 Halliday, G. — Rev. of The living tundra 229-230 Halliday, G. — Rev. of Vegetation of the Soviet polar deserts 229-230 Halliday, G., with H. Hofmann & J. R. Edmond- son — The Cumbrian herbarium of W. H. Youdale 204-206 Hamadryas 252 Hammarbya 238; paludosa 102 Harrison, C. M. — Rev. of The biogeography of the British Isles — an introduction 444-445 Hastorf, C. A. & Popper, V. S. (eds) — Current Paleoethnobotany: analytical methods and cultural interpretations of archaeological plant remains (Bk Rev.) 102 Haufler, C. H., with F. J. Rumsey & E. Sheffield — A re-assessment of Pteridium aquilinum (L.) Kuhn in Britain 297-301 Haworth, C. C. — Six native species of Taraxacum new to the British Isles 131-138 Haworth, C. C. & Richards, A. J.— The lectotypi- fication and revision of Dahlstedt’s species of Taraxacum Weber based on British or Irish plants 125-130 Haworth, Christopher C. (1934-1989) (Obit.) 240-242 | Haworth, C. C., contribution to the Flora of Cumbria 242 Hazeldon, E., Naisbitt, T. & Richards, A. J. — Differential pollination efficiency within a hybrid swarm between Dactylorhiza purpur- ella (T. & T. A. Stephenson) So6 and D. fuchsii (Druce) S06 391-393 Heath, J., with T. Tarpey — Wild flowers of north- east Essex (Bk Rev.) 443-444 Hedera 9, 13-15, 223, 258, 273; algeriensis 7; azorica 14; canariensis ‘Azorica’ 14; colchica 461 7; helix L. and H. hibernica (Kirchner) Bean (Araliaceae) in the British Isles 7-15, 82, 388, (map) 8, (plate) 10, subsp. hibernica 7, 13, 14, var. hibernica 13, 14; hibernica (Kirchner) Bean (Araliaceae) in the British Isles, Hedera helix L. and, 7-15, 82, 273, (map) 8, (plate) 10, ‘Albany’ 13, ‘Deltoidea’ 13, ‘Digitata’ 13, ‘Hamilton’ 13, ‘Helford River’ 13, ‘Hibernica’ 13, 82, ‘Maculata’ 13, ‘Variegata’ 13 Helictotrichon neesii (Steudel) Stace, comb. nov. 413 Helleborus viridis 71,75, 76, 79 Hemerocallis fulva (v.c. 49) 432; lilioasphodelus (v.c. 70) 432 Heracleum mantegazzianum 329; mantegazzia- num X sphondylium (v.c. 75, 99) 220; sphon- dylium 183, 362, var. angustifolium 183 Hesperis matronalis 181, 205 Hieracium 71, 214, 243, 244, 419, 431, 441, 450; Section Alpina 243, 431; Series Alpestria 244; britanniciforme 199, 200; britannicum F. J. Hanb. in Wales 199-200; brunneocroceum (v.c. 94) 432; calenduliflorum (v.c. 90) 431; cymbifolium (v.c. 77) 431; dasythrix (v.c. 77) 431; duriceps (v.c. 93) 224; flocculosum (v.c. 77) 431; graniticola (v.c. 94, 96) 431; han- buryi (v.c. 87) 431; holophyllum 199, var. dentulum 199; holosericeum (v.c. 87, 89) 431; insigne (v.c. 92) 432; macrocarpum (v.c. 92) 431; magyaricum subsp. thaunassium 92; marginatum (v.c. 97) 432; memorabile (v.c. 92) 432; peleterianum subsp. subpeleteria- num 92; pilosella 185; placerophylloides (v.c. 73) 224; reticulatum (v.c. 93) 224; rigens (v.c. 35) 224; rubiginosum 185; sabaudum (v.c. 81, 94) 431; salticola (v.c. 35) 224; sangui- neum (v.c. 77) 431; shoolbredii (v.c. 77) 431; solum 244; sparsifolium (v.c. 75) 224; steno- lepis 199; strictiforme (v.c. 93) 224; subbri- tannicum 199; subcrassum (v.c. 76) 432; tenuifrons (v.c. 88, 92) 432 Hierochloe odorata (v.c. 109) 436 Hill-Cottingham, P. — Somerset ferns, a field guide (Bk Rev.) 231-232 Hippophae rhamnoides (v.c. 73) 426 Hippuris vulgaris 79, (v.c. 45) 220 Hirschfeldia incana (v.c. 38) 215, (v.c. 46, 77) 420 Hobson, D. D.- The status of Populus nigra L. in the Republic of Ireland 303-305 Hodgson, J. G., with J. P. Grime & R. Hunt — Comparative plant ecology. A functional approach to common British species (Bk Rev.) 230-231 Hofmann, H. — Rev. of The herbcrium handbook 324 Hofman, H., Edmondson, J. R. & Halliday, G. — The Cumbrian herbarium of W. H. Youdale 204-206 462 Holcus lanatus 87 Honkenya peploides 270 Hordelymus europaeus (v.c. 53) 228 Hordeum jubatum (v.c. 72) 228 Horsman, F. — A new dactylorchid hybrid 395— 399 Horsman F. — Abnormal flowers in Platanthera chlorantha (Custer) Reichenb. and Gymna- denia conopsea (L.) R.Br. subsp. conopsea 316-317 Horsman, F. — The first record for Vaccinium uliginosum Lx var. pubescens (Wormsk.) Hornem. from the British Isles 87-88 Horsman, F. — Rev. of Plants of Upper Teesdale SZ] Hottonia 248; palustris 71, 79 Houston, L., with L. C. Frost, C. M. Lovatt & A. Beckett — Allium sphaerocephalon L. and introduced A. carinatum L., A. roseum L. and WNectaroscordum — siculum (Ucria) Lindley on St Vincent’s Rocks, Avon Gorge, Bristol 381-385 Hulthemia 247 Humulus lupulus 184 Hunt, A. — Plant names of medieval England (Bk Rev.) 325-326 Hunt, A. — Popular medicine in thirteenth century England (Bk Rev.) 325-326 Hunt, R., with J. P. Grime & J. G. Hodgson -— Comparative plant ecology. A_ functional approach to common British species (Bk Rev.) 230-231 Hyacinthoides hispanica X non-scripta (v.c. 39) 226; non-scripta 117, 388 Hydrilla verticillata 176 Hydrocharis morsus-ranae (v.c. 67) 225 Hydrocotyle (Apiaceae) naturalised in Britain and Ireland, New Zealand species of, 93-95, (key) 94; microphylla 94; moschata 94; novae-zeelandiae 94, var. montana 94, var. novae-zeelandiae 94; sibthorpioides 94; vul- garis 73, 94 Hylm6, B., with K. E. Flinck — Two new species of Cotoneaster 311-313 Hymenophyllum tunbrigense (v.c. 67) 419 Hypericum 441; androsaemum 441, (v.c. 77) 421; x desetangsii Lamotte nm. desetangsii in Yorkshire, with special reference to its spread along railways 63-67; elodes 73, 80; hirsutum 416; humifusum 181, 416; macula- tum 63-66, subsp. maculatum 180, 181, (v.c. 69) 421, subsp. obtusiusculum 64, (v.c. 25) 216, subsp. obtusiusculum X perforatum 63, (v.c. 4, 11) 421; maculatum X perforatum (v.c. 38, 67, 68) 216; montanum 80; perfora- tum 63-66 Hypochaeris 259; radicata 269, 271 Iberis amara 80, (v.c. 12) 216 INDEX TO WATSONIA VOLUME 18 TIfloga 253 Ilex aquifolium 14, 231 Impatiens capensis (v.c. 25) 423; glandulifera 84, 117; parviflora (v.c. 35) 217, (v.c. 42, 94) 423 Ingram, G. I. C. & Dunster, C. W. — A nonre- supinate abnormality in Orchis purpurea Hudson 317-319 Inula conyza 416; helenium 79, 185 Ipomoea pes-caprae 328 Iris foetidissima 416, (v.c. 69) 433; sibirica (v.c. 83) 226; spuria (v.c. 6) 226 Isoetes echinospora 234; hystrix 234; lacustris (v.c. 72) 419 Jalas, J. & Suominen, J. (eds) — Atlas Florae Europaeae. Volume VIII: Nymphaeaceae to Ranunculaceae (Bk Rev.) 326-327 Jasione montana 76, 80, (v.c. 38, 53) 223 Jermy, A. C., with J. W. Clark — Additions to the flora of Mull and adjacent small islands 179- 187 Juncus 441, 448; acutiflorus 164; ambiguus (v.c. 108) 226, (v.c. 46) 433; bufonius 441; capita- tus 441; compressus (v.c. 5, 39) 226, (v.c. 77) 432; maritimus 207; planifolius 441; squarrosus 185; subulatus 441; tenuis (v.c. 7) 226; triglumis 185 Juniperus communis subsp. alpina 180, subsp. nana 180 Jury, S. L. — Rev. of Atlas of the British flora. Repr. of 3rd ed. 441-442 Jury, S. L., with F. J. Rumsey — An account of Orobanche L. in Britain and Ireland 257-295 Kaplopanax 273 Kay, G. M. — Rev. of A guide to some difficult plants 441 Kelly, D. L. — Cornus sericea L. in Ireland: an incipient weed of wetlands 33-36 Kenneth, A. G. — an appreciation 244 Kenneth, Archibald Graham (1915-1989) (Obit. ) 242-244 Kent, D. H. — The Shasta Daisy 89 Kent, D. H. — Rev. of Flora of New Zealand, Volume IV. Naturalised Pteridophytes, Gymnosperms, Dicotyledons 328-329 Kingdonia 252 Kirengeshoma 101 Knautia 249; arvensis 66 Knowltonia 252 Koeleria macrantha 87 Kumlienia 252 Kwasnik, E. I. — Rev. of Plants for people 440-441 Laburnum anagyroides 205 Laccopetalum 252 Lack, A. J. & Diaz, A. — The pollination of Arum maculatum L. — a historical review and new observations 333-342 INDEX TO WATSONIA VOLUME 18 463 Lactuca serriola 444 Lagarosiphon major (v.c. 29) 432 Lamiastrum galeobdolon 117, subsp. argentatum 329, (v.c. 67, 79) 223, (v.c. 77) 430 Lamium album 185; amplexicaule 185; hybridum 185, (v.c. 11) 223 Lancaster, R. — Travels in China. A plantsman’s paradise (Bk Rev.) 102-103 Lang, D. C. — A new variant of Ophrys apifera Hudson in Britain 408-410 Lang, D. — A guide to the wild orchids of Great Britain and Ireland (Bk Rev.) 103 Larix 298; kaempferi 179 Lathraea clandestina (v.c. 4, 35) 223; squamaria 79, 388 Lathyrus aphaca 80; montanus 66; nissolia (v.c. 64, 69, 99) 423; palustris 243; sylvestris (v.c. 43) 217 Lavandula 440 Lawrence, M. J., with A. F. Raybould, A. J. Gray & D. F. Marshall — The origin and taxonomy of Spartina X neyrautii Foucaud 207-209 Legousia hybrida 80 Lemna minuscula 439, (v.c. 38) 227; polyrhiza 80; trisulca (v.c. 73) 227 Leontodon hispidus 87; taraxacoides 185 Lepidium heterophyllum 181, 416; latifolium (v.c. 38) 216; ruderale 329 Leptochloa muelleri (Benth.) Stace, comb. nov. 413 Leucanthemum lacustre 89; maximum 89; X superbum (Bergmans ex J. Ingram) Kent, comb. nov. 89; vulgare 87 Leucocoma 252 Leucojum vernum 185 Lewington, A. — Plants for people (Bk Rev.) 440— 44] Leymus arenarius 372, 416 Libbey, Richard Pearse (1911-1987) (Obit.) 451- 452 Lilium martagon (v.c. 99) 226 Limodorum abortivum 261 Limonium species, Chromosome numbers in two, 82-84; humile 82, 83; vulgare 82, 83 Linaria 107; purpurea (v.c. 72) 429; purpurea X repens (v.c. 46) 429; repens X vulgaris (v.c. 99) 222, (v.c. 77) 429 Linum perenne 80 Liparis loeselii 102 Listera cordata 80; ovata 154, 186 Lithospermum arvense (v.c. 61) 428; officinale 74 Littorella uniflora 73, 79 Logfia 253 Lolium 401, 402 Long, D. G., with M. E. Braithwaite — The botanist in Berwickshire (Bk Rev.) 438-439 Lonicera caerulea (v.c. 57) 224; caprifolium (v.c. 79) 224; grata 264; involucrata 224, (v.c. 70) 430; nitida (v.c. 77) 430; xylosteum (v.c. 50) 223, (v.c. 50, 95, 99) 430 Lotus angustissimus 272; corniculatus 39, 87, 402; subbiflorus 272; uliginosus 66 Lovatt, C. M., with L. C. Frost, L. Houston and A. Beckett — Allium sphaerocephalon L. and introduced A. carinatum L., A. roseum L. and Nectaroscordum siculum (Ucria) Lindley on St Vincent’s Rocks, Avon Gorge, Bristol 381-385 Lowe, C., with M. Lowes & G. Ward — Plants of Upper Teesdale (Bk Rev.) 327 Lowes, M., with C. Lowe & G. Ward — Plants of Upper Teesdale (Bk Rev.) 327 Ludwigia palustris 102 Luzula 441; campestris 441; multiflora 441 Lycium barbarum (v.c. 75) 222, (v.c. 73) 429; chinense (v.c. 12, 77) 222 Lycopodium clavatum 180, (v.c. 52) 214 Lycopus europaeus 184, 416 Lysichiton americanus 97, 179, 186, (v.c. 35) 434 Lysimachia nemorum 416; vulgaris 75 Lythropsis 254; peploides 254 Lythrum 249, 253, 254; biflorum 254; boraei 254; borysthenicum 253; hispidulum 254; loise- leurti 254; nummulariifolium 253; portula 71, 80; salicaria 183, 254 Love, D. (trans.), with V. D. Aleksandrova — Vegetation of the Soviet polar deserts (Bk Rev.) 229-230 Love, D. (trans.), with Y. I. Chernov — The living tundra (Bk Rev.) 229-230 Magnolia 100 Mahonia aquifolium 320, (v.c. 94) 215; aquifo- lium X repens 320; X decumbens Stace, hyb. nov. 320; repens 320 Malus domestica 85 Malva alcea 80; sylvestris 180, 181 Mamillaria 236 Marshall, D. F., with A. F. Raybould, A. J. Gray & M. J. Lawrence — The origin and tax- onomy of Spartina X neyrautii Foucaud 207- 209 Martin, M. H., with A. J. Willis & K. B. Taylor — Orchis purpurea Hudson in the Avon Gorge, Bristol 387-390 Matricaria matricarioides 185 McAllister, H. A. & Rutherford, A. — Hedera helix L. and H. hibernica (Kirchner) Bean (Araliaceae) in the British Isles 7-15 McNeill, J. — Rev. of Plant taxonomy and bio- systematics. 2nd ed. 106-107 Meconopsis cambrica 180 Medicago arabica 416, (v.c. 51) 423; falcata x sanval(vie.! 77) 217; "minima (W.ck 12)°217; sativa 260, subsp. falcata 402, subsp. sativa 402 Meikle, R. D. — Salix X seminigricans A. & G. 464 Camus: a willow hybrid new to the British Isles 410-412 Melampyrum pratense (v.c. 29, 61) 429 Melanium 247 Melica nutans 76, (v.c. 43) 228, (v.c. 93) 435; uniflora 388 Mentha 187; arvensis 184; arvensis X spicata (v.c. 42) 430; nicholsoniana 212; X piperata 184; pulegium 75, 80, (v.c. 12, 25) 430; spicata x suaveolens 212; suaveolens 184; X villosa nm. alopecuroides 184, (v.c. 75) 430, var. nichol- soniana (Strail) R. Harley, comb. nov. 212 Menyanthes trifoliata 154 Mercurialis perennis 11, 364, 388, 416 Mertensia maritima 184, (v.c. 73) 428 Middendorfia 254; borysthenica 254; hamulosa 254 Milium effusum (v.c. 81) 228 Mill, R. R., with K. Tan (ed.) & T. S. Elias — The Davis & Hedge Festschrift (Bk Rev.) 439-440 Miller, A. G. & Morris, M. — Plants of Dhofar (Bk Rev.) 327-328 Millward, D. — A flora of Wensleydale. A centen- nial review of the plants of the Dale (Bk Rev.) 232 Milne-Readhead — The B.S.B.I. Black Poplar survey 1—5 Mimulus L., A new hybrid binomial in, 210-212, 327; ‘A. T. Johnson’ 211; cupreus X luteus var. rivularis 211; guttatus 210-212; guttatus x luteus var. rivularis 211; luteus 210, 211, var. rivularis 211, 212; X maculosus 211; moschatus 184, (v.c. 77) 222; X robertsii Silverside, hybr. nov. 211, 212; variegatus 211 Minuartia hybrida 80; verna 116, 117, 327 Molinia 217; caerulea subsp. altissima (v.c. 35) 224 Monotropa hypopitys (v.c. 39) 221 Montia 248; fontana 201, subsp. fontana 181; perfoliata179, 181, 416; sibirica 181 Moore, D. M. — Rev. of The correspondence of Charles Darwin. Vol. 4, 1847-1850 98 Morris, M., with A. G. Miller — Plants of Dhofar (Bk Rev.) 327-328 Myosotis 97, 113, arvensis subsp. umbrata (v.c. 67, 68) 222, (v.c. 99) 428; ramosissima subsp. globularis (v.c. 68) 222; sylvatica 416 Myosurus 252; minimus 76, 80, (v.c. 54) 420 Myriophyllum 248; aquaticum (v.c. 25, 39, 67) 426; spicatum 79; verticillatum 80 Myrrhis odorata 327 Naisbitt, T., with E. Hazeldon & A. J. Richards — Differential pollination efficiency within a hybrid swarm between Dactylorhiza purpur- ella (T. & T. A. Stephenson) So6 and D. fuchsii (Druce) So6 391-393 Najalis flexilis (v.c. 97, 103) 225 INDEX TO WATSONIA VOLUME 18 Naravelia 252 Nectaroscordum 382; siculum (Ucria) Lindley on St Vincent’s Rocks, Avon Gorge, Bristol, Allium sphaerocephalon L. and introduced A. carinatum L., A. roseum L. and, 381-385, subsp. bulgaricum 382, 383, subsp. siculum 382, 383 Nelson, George A. (1903-1989) (Obit.) 111-112 Neottia nidus-avis 73, 80, 186 Nepeta cataria 75, 80 Newton, A. — Rubus stenophyllus Lef. & Muell. in Britain 89-90 Newton, A. & Porter, M. — Five brambles from Wales 189-198 Newton, J. M. — Rev. of The Macmillan field guide to British wildflowers 104-105 Norman, E. (ed.)—A guide to some difficult plants (Bk Rev.) 441 Nymphoides peltata 75, 79, (v.c. 80) 221 Obituaries 111-114, 238-246, 447-452 Odontites verna 184 Oenanthe aquatica 79; pimpinelloides (v.c. 7) 220; silaifolia (v.c. 54) 427 Oenothera 408; Subsection Oenothera 407; bien- nis 407, 408, (v.c. 75) 426; cambrica Ros- tanski and O. novae-scotiae Gates (Onagra- ceae), The status of, 407-408; erythrosepala (v.c. 80) 220, (v.c. 43) 426; fallax (v.c. 25) 220; novae-scotiae Gates (Onagraceae), The status of Oenothera cambrica Rostanski and, 407-408; parviflora 408; stricta (v.c. 38) 220 Onobrychis victifolia 402 Onoclea sensibilis (v.c. 12, 99) 419 Ononis repens 270; spinosa 66, 416 Onopordon acanthium (v.c. 50) 431 Ophioglossum azoricum (v.c. 11) 215; vulgatum 117, 180 Ophrys 391; apifera Hudson in Britain, A new variant of, 408-410, 75, 79, (illus.) 409, subsp. jurana 409, var. bicolor 410, var. botteronii 409, var. chlorantha 410, var. fla- vescens 410, var. friburgensis 409, var. imma- culata 410, var. trollii 410; insectifera 80, 154; sphegodes 43, 45 Orchis 307, 391; anatolica 307; X braunii var. townsendianus 396; coriophora 307; erice- torum 395; francis-drucei 170; latifolia 395, 409, var. eborensis 162, 170; latifolia-macu- lata 396; majalis subsp. traunsteinerioides 170, subsp. traunsteinerioides var. eborensis 170; mascula 39, 47, 186, (v.c. 72) 433; morio 39, 44, 45, 47, 79, 307; praetermissa 395; purpurea Hudson, A nonresupinate abnor- mality in, 317-319, Hudson in the Avon Gorge, Bristol 387-390, (illus.) 318; traun- steineri 153; traunsteinerioides 153, 170; ustu- lata L. in southern England, The current distribution and abundance of, 37—48, 73, 80, INDEX TO WATSONIA VOLUME 18 102, (map) 38, var. albiflora 40 Oreopteris limbosperma 231, 232 Origanum vulgare 66, 416 Ornithogalum pyrenaicum (v.c. 51) 432; umbella- tum 79 Ornithopus perpusillus (v.c. 98) 423 Orobanche L. in Britain and Ireland, An account of, 257-295, (key) 258-259; Section Oro- banche 258; Section Trionychon 258; aegyp- tiaca 258-260, (illus.) 275, (illus.) 278; alba 184, 258, 259, 263, 264, (illus.) 275, (illus.) 278, (map) 285; amethystea 270, 272; are- naria 261; barbata 272; caerulea 261; caryop- hyllacea 258, 259, 263, 265, 266, 273, 294, (illus.) 275, (illus.) 279, (map) 287; crenata 258, 259,263, 267, 270) 27362742 (alus.) 276, (illus.) 280, (map) 293; elatior 79, 248, 259, 263, 266, 267, 269-271, 273, 294, (illus.) 276, (illus.) 279, (map) 288, f. citrina 267; epithy- mum 263; flava 258, 273, 294, (illus.) 276; hederae 81, 82, 258, 259, 264, 268, 270, 272, 273, (v.c.12) 223, (v.c. 12, 42) 429, (illus.) 277, (illus.) 281, (map) 292, f. monochroma 82, 223; leucorum 266; loricata 259, 265, 267, 268, 272, 273, (illus.) 277, (illus.) 280, (map) 289; lucorum 258, 273, 294, (illus.) 276; major 262, 265-267; maritima 271, 272; minor 257, 259, 261, 264, 265, 267-270, 272, 273, (key) 269-270, var. compositarum 267, 269, 270, (illus.) 277, (illus.) 280, var. flava 269, 271, (illus.) 277, (illus.) 281, var. lutea 271, var. maritima 268, 270-272, (illus.) 277, (illus.) 281, (map) 291, var. minor 270-272, (illus.) 277, (illus.) 280, (map) 290; pallidif- lora 264; picridis 265, 267, 268, var. carotae 274; pruinosa 273; purpurea 258-261, (illus.) 275, (illus.) 278, (map) 283; ramosa 258-261, 273, (illus.) 275, (illus.) 278, (map) 282, subsp. ramosa 258, 260; rapum 262; rapum- genistae 258, 259, 262, 263, 266, 267, 274, 294, (v.c. 12) 223, (illus.) 276, (illus.) 279, (map) 284; reticulata 259, 264, 265, 273, (illus.) 275, (illus.) 279, (map) 286, var. pallidiflora 265, var. procera 265; ritro f. hypochaeroides 271; rubra 263, 264; speciosa 273; vulgaris 265 Orthilia secunda 184 Oxalis 452; articulata (v.c. 75) 217; corniculata (v.c. 73) 423; exilis (v.c. 38, 81) 217 Oxygraphis 252 Paeonia officinalis (v.c. 77) 215, (v.c. 69) 420 Page, C. N. — Herbarium sheets of the rare fern X Asplenophylltis microdon (T. Moore) Alston in Edinburgh 319 Page, C. N. — Ferns: their habitats in the British and Irish landscape. (A natural history of Britain’s ferns.) (Bk Rev.) 103-104 Pankhurst, R. J. (trans.), with F. Weberling — 465 Morphology of flowers and _ inflorescences (Bk Rev.) 236-237 Papaver argemone 76, 79, (v.c. 50) 420; hybridum 80; lecoqti (v.c. 28, 35) 420; orientale (v.c/ 69) 420; somniferum 180 Paphiopedilum charlesworthii 451 Parapholis incurva 323, (v.c. 5, 54) 436 Parentucellia viscosa (v.c. 8) 223, (v.c. 95) 429 Parietaria diffusa 183; judaica 183 Paris quadrifolia 71, 79 Parnassia palustris 73, 79 Parnell, J.— Rev. of Morphology and evolution of vascular plants 100 Paroxygraphis 252 Parthenocissus quinquefolia (v.c. 73) 423 Pastinaca sativa 416 Payne, R. M. — Rev. of Somerset ferns, a field guide 231-232 Pearman, D. — Alopecurus bulbosus Gouan in Dorset 206-207 Pelargonium 273, 274 Pentaglottis sempervirens 184 Peplis 249, 253, 254; australis 254; biflora 254; boraei 254; borysthenica 253; erecta 254; hispidula 254; nummulariifolia 254; portula 254; timeroyi 254; tithymaloides 254 Perring, F. H. — Rev. of Atlas der Farn- und Bliitenpflanzen der Bundesrepublik Deutschland 101-102 Perring, F. H. & Walters, S. M. (eds) — Atlas of the British flora. Repr. of 3rd ed. (Bk Rev.) 441-442 Perring, F. H. & Walters, S. M. — The Macmillan field guide to British wildflowers (Bk Rev.) 104-105 Perry, A. R., with R. G. Ellis — Obit. of Arthur Edward Wade (1895-1989) 113-114 Petasites 258, 274; fragrans (v.c. 77) 224; japoni- cus (v.c. 35, 73) 430 Peucedanum officinale 444; ostruthium 327, (v.c. 79) 220 Phacelia tanacetifolia (v.c. 12) 221 Phalaris arundinacea 187 Phascum cuspidatum 364 Phelipaea ramosa 260 Phelypaea aegyptiaca 260; caerulea 261 Philonotis fontana 201 Phragmites 203, 207, 422 Phyllitis 442; scolopendrium 319 Physocarpus opulifolius (v.c. H36) 217 Picea abies 179 Pickard, R. S. (ed.), with R. Sawyer — Honey identification (Bk Rev.) 106 Picris 259; hieracioides 80, 268 Pilosella officinarum 268, subsp. euronota (v.c. 75) 225, subsp. officinarum 185, (v.c. 75) 225; peleteriana subsp. subpeleteriana (Nae- geli & Peter) P. D. Sell, comb. nov. 92; praealta subsp. thaumasia (Peter) P. D. Sell, 466 comb. nov. 92 Pilularia globulifera 416, (v.c. 53, 75) 215 Pimpinella major 66; saxifraga 66, 87 Pinguicula vulgaris 87, 154 Pinus contorta 179; deodara 92; devdara 92; syl- vestris 180 Piuttia 252 Plagiomnium undulatum 388 Plant Records 214-228, 419-436 Plantago coronopus 416; lanceolata 87, 364, 416; major subsp. intermedia 416, 444, (v.c. 67, 68) 223, (v.c. 39, 73) 430, subsp. major 416; media (v.c. 42, 94) 430 Platanthera 316, 317; bifolia 39, 43, 79, 316; chlorantha (Custer) Reichenb. and Gymna- denia conopsea (L.) R.Br. subsp. conopsea, Abnormal flowers in, 316-317, 186; chloran- tha X Pseudorchis albida 316 Poa angustifolia 186, 234; chaixii 179, 186; praten- sis 11, 87, 364; subcaerulea 186, 234, 444; trivialis 84 Polemonium caeruleum 77 Polycarpon tetraphyllum 272 Polygala 39; amara 87; vulgaris 181 Polygonatum multiflorum 185 Polygonum 250, 251, 351, 357; Section Aconogo- non 356, 357; alpinum 356; amphibium 183; arenastrum (v.c. 50) 427; aviculare 183; bis- torta 180, 183; boreale (v.c. 103) 220; (plate) 355, (plate) 356; campanulatum 183, 351, 353, 354, 357, (v.c. 70) 428, var. campanula- tum 353, var. fulvidum 353, var. lichiangense 351, 354; cuspidatum 183; hydropiper 183; lichiangense W. Smith: rejected as a natura- lized British species 351-358, (plate) 352, (plate) 355, (plate) 356; maritimum (v.c. 1, 11) 427; minus (v.c. 34) 427; mite (v.c. 50) 427; molle 357; polystachyum 351, 353, 354, 356, 357, var. pubescens 351, 353, 354, 357, (plate) 355, (plate) 356; rurivagum (v.c. 7, 46, 73) 220, (v.c. 46) 427; sachalinense 183; weyrichii 354, 356 Polypodium 231; cambricum 234, (v.c. 11) 420; interjectum 234, (v.c. 77) 420; interjectum X vulgare (v.c. 77) 420 Polystichum 231; aculeatum 180; setiferum 180 Popper, V. S., with C. A. Hastorf (eds) — Current Paleoethnobotany: analytical methods and cultural interpretations of archaeological plant remains (Bk Rev.) 102 Populus 44; * canadensis 3, var. serotina 1; X euramericana 305; nigra L. in the Republic of Ireland, The status of, 303-305, 1-3, 5, 184, (plate) 2, (map) 4, (map) 304; tremula 184 Porter, M., with A. Newton — Five brambles from Wales 189-198 Potamogeton 91, 246; Subgen. Coleogeton 90; alpinus * praelongus (v.c. H35) 225; com- pressus 80; crispus X friesti (v.c. 73) 225; INDEX TO WATSONIA VOLUME 18 crispus X perfoliatus (v.c. 73) 225; filiformis Pers. in Anglesey 90-91, (v.c. 80) 225; friesii (v.c. 7, H35) 225; gramineus 79, (v.c. 68) 225; gramineus X natans (v.c. 87, 97) 432; lucens x natans (v.c. 27) 432; pectinatum 90; pecti- natus 90, 117, (v.c. 43) 432; pusillus 79, 432; rutilus (v.c. 103, 108) 225; trichoides (v.c. 2, 54) 225, (v.c. 12) 432 Potentilla anglica (v.c. 7, 79) 218; argentea 80, (v.c. 5, 72) 425; crantzii (v.c. 72) 425; erecta 87, subsp. strictissima (v.c. H40) 218; norve- gica (v.c. 41) 218; reptans (v.c. 95) 425; sterilis 182, 416; tabernaemontani 80 Poterium sanguisorba 39 Prelli, R. — Guide des fougéres et plantes alliées (Bk Rev.) 442 Preston, C. D. — Potamogeton filiformis Pers. in Anglesey 90-91 Primavesi, A. L., with G. G. Graham — Notes on some Rosa taxa recorded as occurring in the British Isles 119-124 Primula 248; farinosa 87, 154, 232; sonchifolia 351; veris 39, 117, 364, 416; vulgaris 416, 442 Proctor, M. C. F. — Obit. of Guy Malcolm Spooner (1907-1989) 245 Prunus armeniacus 101; avium 182; domestica X spinosa (v.c. 5) 218; X fruticans 439; lusita- nica 205; padus 71, 79, 182; serotina (v.c. 46) 426; spinosa 182, 231, 360, 364 x Pseudanthera breadalbanensis 316 Pseudorchis 317; albida 180, 186, 316 Pseudoscleropodium purum 364 Pseudotsuga glaucescens 92; menziesii subsp. glaucescens (Schwerin) P. D. Sell. comb. nov. 92; taxifolia subsp. glaucescens 92 Psilotum 442 Pteridium 297-300, 442; aquilinum (L.) Kuhn in Britain, A re-assessment of, 297-301, 11, subsp: aquilinum 297-300, subsp. aquilinum var. aquilinum 297, 298, subsp. aquilinum var. latiusculum 299, 300, subsp. atlanticum 297, 298, 300, subsp. caudatum 300, subsp. latiusculum 297-300, subsp. latiusculum X aquilinum subsp. aquilinum 299 Puccinellia distans 186; rupestris (v.c. 53) 435 Puccinia 384; allii 384 Pulicaria vulgaris Gaertner in Britain in 1990, The status of, 405—406 Punt, W., Blackmore, S. & Clarke, G. C. S. (eds) — The northwest European pollen flora. Vol. 5 (Bk Rev.) 105 Pyrola minor 79; rotundifolia subsp. maritima (v.c. 50) 221 Pyrus communis 85 Quercus 231, 236; cerris 179; robur 34 Radiola linoides 73, 75, 80 Raghavan, V. — Developmental biology of fern INDEX TO WATSONIA VOLUME 18 467 gametophytes (Bk Rev.) 232-233 Ranunculus 247, 252; L. subgenus Batrachium (DC.) A. Gray, Three natural hybrids in, 139-146, 326, 412; aquatilis 145, (v.c. 94) 215; aquatilis xX fluitans 413; aquatilis x trichophyllus 139; arvensis (v.c. 7) 215; asiati- cus 247; bachii Wirtg., The typification of, 412-413, 142, 144, 145; x bachii 139, 144, 145, 413; baudotii 207, (v.c. 29) 420; bulbo- sus 87; cambricus 145; circinatus (v.c. 81) 215; ficaria 34, 84, 117; fluitans 144, 145, 413, (v.c. 4) 420; fluitans < aquatilis 139, 144, 145; fluitans X peltatus 139, 142, 413, (plate) 143; fluitans X trichophyllus 139, 144, 145; fluitans P bachii 144; X kelchoensis S. Webster, hybr. nov. 142, 144, 413, (plate) 143; lingua 73, 79; lutarius 139; X novae-forestae S. Webster, hybr. nov. 139, (plate) 140; omiophyllus 141; omiophyllus X peltatus 139; omiophyllus x tripartitus 139, (plate) 140; parviflorus 76, 80; peltatus 144, (v.c. 7) 215; penicillatus 144, 145, subsp. penicillatus 144, subsp. pseudo- fluitans 145, (v.c. 95) 420, var. pseudofluitans 420; repens 364, 416; sceleratus 180; tricho- phyllus 145, 413; tripartitus 141 Raphanus raphanistrum 215; sativus (v.c. 50) 215, (v.c. 50) 421 Rapistrum rugosum (v.c. 12) 421 Raybould, A. F., Gray, A. J., Lawrence, M. J. & Marshall, D. F. — The origin and taxonomy of Spartina X neyrautii Foucaud 207—209 Raynes, P., with K. Futter — The flora of Derby (Bk Rev.) 99-100 Rechinger, K. H. — Rumex acetosa L. subsp. biformis (Lange) Valdés-Bermejo & Castroviejo in Britain 209-210 Reports 115-117, 331-332 Reseda alba (v.c. 11) 421 Reynoutria 250; japonica 117, 183; japonica xX sachalinensis (v.c. 11) 220; sachalinensis 183, (v.c. 49, 94) 428 Rhinanthus angustifolius 75, 76, 80; borealis 184; minor 39, 66, 117, 184, subsp. calcareus (v.c. 7)222 Rhododendron ponticum 36, 184 Rhopalopodium 252 Rhus typhina (v.c. 41) 216, (v.c. 41) 423 Rhynchospora alba 186 Rhynia 100, 442 Ribes nigrum 183; rubrum 183; spicatum (v.c. 67, 68) 219 Rich, T. C. G., with D. Bevan — Bevan’s bitter- cress: the small, white-flowered variant of Cardamine X fringsii Wirtgen also present in Britain 403—405 Richards, A. J., with C. C. Haworth — The lectotypification and revision of Dahlstedt’s species of Taraxacum Weber based on British or Irish plants 125-130 Richards, A. J., with E. Hazeldon & T. Naisbitt — Differential pollination efficiency within a hybrid swarm between Dactylorhiza purpur- ella (T. & T. A. Stephenson) So6 and D. fuchsii (Druce) So6 391-393 Rihan, J. R. & Gray, A. J. — The distribution of hybrid Marram Grass, X Calammophila bal- tica (Fligge) Brand, in the British Isles 369- 379 Robertson, S. A. — Flowering plants of Seychelles (Bk Rev.) 105-106 Robinia pseudacacia (v.c. 70) 423 Robson, N. K. B. — Rev. of Morphology of flowers and inflorescences 236-237 Robson, N. K. B. — Rev. of The names of plants. 2nd ed. 100-101 Robson, N. K. B. — Rev. of Travels in China. A plantsman’s paradise 102-103 Rorippa amphibia 79; austriaca (v.c. 77) 421; austriaca X sylvestris (v.c. 98) 216, (v.c. 77) 421; islandica 71, 80; palustris * amphibia (v.c. 38) 216; sylvestris (v.c. 94) 421 Rosa taxa recorded as occurring in the British Isles, Notes on some, 119-124, 231, 247, 360; Group Andegavenses 120, 122; Group Deseglisei 123; Group Dumales 120, 122; Group Lutetianae 120; Group Pubescentes 120; Group Rubiginosae 122; Group Scabra- tae 120, 122; Group Subcaninae 123; Group Subcollinae 123; Group Transitoriae 120; afzeliana 121, 122; agrestis 122, 124; andega- vensis 122; arvensis 122-124; blondaeana 122; caesia 121, 124, (v.c. 41, 75) 425, subsp. caesia 124, subsp. caesia X canina 123, subsp. glauca (Nyman) Graham & Primavesi, comb. nov. 121, 122, 124, (v.c. 75) 425, subsp. glauca X canina 123, (v.c. 41) 425; caesia X canina 121, (v.c. 12) 425; canina 120-124, var. squarrosa 122; canina X mic- rantha (v.c. 11) 425; canina xX obtusifolia 120, (v.c. 41) 425; canina X rubiginosa (v.c. 41) 425; canina X tomentosa 124; coriifolia 121; corymbifera 120, 121; deseglisei 123; dumalis 121, subsp. dumalis 121; < dumalis 121; dumetorum 120, 121, 123, var. deseglisei 123;