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Kevin Walker.Pub Watsonia 26: 291–302RECENT (2007) CHANGES IN THE FLORA OF THE BRITISH ISLES 291 The last thirty five years: recent changes in the flora of the British Isles K. J. WALKER* CEH Monks Wood, Abbots Ripton, Huntingdon, Cambs. PE28 2LS ABSTRACT county of Cambridgeshire (v.c. 29). I do not deal with changes in plant conservation, land In 1970 Max Walters predicted a number of changes use, or the ten activities he proposed for the that have taken place in the British and Irish flora society as these now form the basis of much of over the last 35 years. Although few species became the society’s current work (e.g. monitoring nationally extinct during this period, losses at the common species, research on rare species, local scale were dramatic, particularly in S.E. England where land use pressures were greatest. publication of handbooks). Despite these losses the British flora has become increasingly diverse due to the introduction, natural- 1. FLORISTIC RICHNESS WILL CONTINUE TO INCREASE isation and spread of alien species. Research into their habitats, modes of dispersal, and competitive AS A RESULT OF MAN’S ACTIVITIES inter-actions with native species has provided valuable ecological insights. Hybridisation events As Max predicted species invasions have have produced some of the best studied examples of exceeded extinctions by an order of magnitude hybrid speciation. In the future, such naturalisation leading to a dramatic net increase in the events may also provide important evidence for number of plant species recorded in the British major environmental perturbations such as climate Isles (Fig. 1). Despite localised losses only 21 change. native species have gone extinct nationally since records began in the sixteenth century KEYWORDS: aliens, diversity, extinction, hybrid- (Stace 2002). Nine of these species have been isation, naturalisation, neophytes, urban habitats. lost in the last 35 years, although some are either dubiously native (e.g. Galeopsis sege- INTRODUCTION tum) or are likely to have been over-looked (e.g. Epipogium aphyllum) (Table 1). During In 1970 Max Walters concluded The flora of a the same period an equal number of species changing Britain conference by predicting a have been discovered or newly described of number of changes that were likely to take which only two are of questionable origin place in the British flora over the subsequent (Cystopteris diaphana, Serapias parviflora) twenty-five years (Walters 1970). Although and one (Epipactis youngiana) may be a brief, his paper dealt with floristic richness, recently evolved ecotype (Hollingsworth et al. aliens, extinctions, plant conservation and land 2006). In comparison to these rather modest abandonment. The aim of this paper is to changes to the native flora, there has been a review three of these predictions in view of the dramatic increase in the number of naturalised major changes which have taken place: (1) aliens and casuals (Fig. 1): by the middle of the floristic richness has increased as a result of 1950s the number of recorded casuals exceeded man’s activities; (2) the number of national and the number of natives whereas there were regional extinctions has continued to increase; around 600 naturalised aliens (30% of the and (3) much has been learnt from the native flora). By 2000 the number of casuals naturalisation of alien species. In addressing and naturalised aliens had increased to make up these three predictions I have intentionally 45% and 18% of the entire British flora drawn heavily (but not exclusively) on research respectively. published within this journal, especially the Historically alien plants have arrived review by Brenan (1982) which deals with unintentionally as escapes from botanic gar- many of the same topics. Wherever possible, I dens or as contaminants in the ballast of ships, have also referred to changes in Max’s home grain, animal fodder and waste products of *e-mail: [email protected] 292 K. J. WALKER 3500 Native Naturalised aliens 3000 Casuals 2500 2000 1500 1000 Number ofspecies 500 0 1900 1920 1940 1960 1980 2000 Year FIGURE 1. The number of native taxa, naturalised aliens and casuals in the flora of Great Britain according to various sources (Druce 1908; Druce 1928; Dandy 1958; Kent 1992; Kent & Stace 2000). ‘Native’ includes agamospecies such as Hieracium, Rubus and Taraxacum. Adapted from Stace (2002). TABLE 1. NATIVE SPECIES DISCOVERED OR KNOWN TO HAVE BECOME EXTINCT IN BRITAIN (INCLUDING ISLE OF MAN) SINCE 1970. UPDATED FROM STACE (2002). SPECIES IN THE CRITICAL GENERA ALCHEMILLA, EUPHRASIA, HIERACIUM, LIMONIUM, RUBUS, SORBUS AND TARAXACUM ARE OMITTED First/last Species V.C. Comment record (a) Discovered Epipactis youngiana 1976 67 Probably only an ecotype of E. helleborine Senecio eboracensis 1979 63 Derived from spontaneous hybrid Crepis praemorsa 1988 69 Utricularia stygia 1988 97 Serapias parviflora 1989 2 Native status uncertain Callitriche palustris 2000 99 Cystopteris diaphana 2000 2 Native status uncertain Carex salina 2004 105 (b) Extinct Arnoseris minima 1971 24 Probable archaeophyte. Re-introduced Bromus interruptus 1972 29 Probable neophyte. Re-introduced Schoenoplectus 1972 59 Only native on Jersey. Re-introduced pungens Galeopsis segetum 1975 49 Probable archaeophyte Saxifraga rosacea 1978 49 Still occurs in Ireland subsp. rosacea Crepis foetida 1980 15 Probable archaeophyte. Re-introduced Epipogium aphyllum 1986 22 Unconfirmed records since 1990 Neotinea maculata 1986 71 Occurs in Ireland. Formerly Isle of Man Senecio eboracensis 2000 61 Possibly extinct? Luzula pallidula is included as extinct by Stace (2002) but there are a number of confirmed records from Holme Fen in Huntingdonshire (v.c. 31). RECENT CHANGES IN THE FLORA OF THE BRITISH ISLES 293 TABLE 2. CHANGE IN THE NUMBER OF TAXA RECORDED IN BERKSHIRE (V.C. 22). ADAPTED FROM CRAWLEY (2005) Druce Bowen Crawley % change (1897) (1968) (2005) 1897–1968 1968–2005 Native & archaeophyte 820 883 955 +8 +8 Neophyte 300 629 710 +110 +13 Apomictic 63 84 184 +33 +119 Subspecies & varieties 400 38 41 -91 +8 Hybrids 70 99 181 +41 +83 clothing manufacture (skins, wool, tan-bark) has allowed an assessment of changes in (Lousley 1953). Although introductions via floristic richness over the past 100 years (Table these agencies declined during the second half 2; Crawley 2005). Whereas the number of of the twentieth century (Shimwell 2006), the native and archaeophyte additions has been numbers originating from horticulture, either as rather modest there has been a dramatic escapes or outcasts from gardens, have in- increase in the number of recorded neophytes creased dramatically. For example, the New (Bowen 1968). Similarly, critical and hybrid Atlas showed that eight of the 10 most rapidly taxa have been much better studied since 1968 spreading species since the 1960s are whereas the vast majority of infraspecific taxa neophytes which were originally introduced as formerly recognised by Druce (1897) are not garden plants (Buddleja davidii, Cotoneaster included in the modern floras of Bowen and simonsii, Laburnum anagyroides, Linaria Crawley. purpurea, Lysimachia punctata, Prunus Based on his experiences in Cambridgeshire cerasus, P. laurocerasus, Syringa vulgaris) Max Walters predicted that urban areas would (Preston et al. 2002). accrue more alien species than rural areas and Changes at the vice-county level show a therefore, that urbanised 10-km squares would similar pattern. Berkshire (v.c. 22) has one of be the richest. However, unlike continental the best studied floras in the country, parti- Europe, British urbanised tetrads are not cularly with regard to alien species, and this significantly richer overall than their rural 500 150 (a) (b) 400 100 300 200 50 100 Neophyte/casual Native/archaeophyte 0 0 0.1 1.0 10.0 100.0 0.1 1.0 10.0 100.0 -1 Population density (ha ) Population density (ha-1) FIGURE 2. The relationship between human population density (displayed on a log scale) and (a) the number of natives and archaeophytes and (b) neophytes and casuals recorded in Cambridgeshire parishes (v.c. 29) between 1987–2000 (n=151). Human population densities were calculated from Population Census Returns for 2001. 294 K. J. WALKER counterparts, although they do support a east of a line running from Poole in Dorset to greater number (and proportion) of introduced Goole in S.W. Yorkshire (‘the Poole to Goole alien species (Roy et al. 1999). The results for line’; Table 3). The vast majority are Northern agriculturally ravaged Cambridgeshire were as (Boreal) species at the southern edge of their predicted by Max with more densely populated range in lowland England that disappeared (and therefore urbanised) parishes tending to be following the widespread loss of lowland marginally more species-rich (R2=4·2%, F=6·5, heaths and bogs (e.g. Diphasiastrum spp., p <0·05) whereas differences in the number of Huperzia selago, Gymnocarpium dryopteris, natives and archaeophytes were not significant Listera cordata, Pseudorchis albida, Pyrola due to large variation in numbers of species media, Scheuchzeria palustris, Selaginella between parishes (R2=1·9%, F=2·9, p=0·09; selaginoides, Trientalis europaeus, Vaccinium Fig. 2a). In contrast, urbanised parishes have vitis-idaea). Others have been lost due to significantly more neophytes and casuals with drainage of fenland (Tephroseris palustris), several of the ten richest parishes being almost hybridisation (Schoenoplectus triqueter), entirely urban (R2=16·9%, F=30·3, p <0·001; ploughing of calcareous grassland (Cirsium Fig. 2b). The slight discrepancy between tuberosum) and changes to aquatic habitats Cambridgeshire and the national picture (i.e. (Elatine hydropiper, Luronium natans) or more species in urban areas) may be due to simply because their extreme rarity in S.E. local factors. Some urban parishes in the England has made them highly susceptible to county have been extraordinarily well recorded other factors (Asplenium obovatum, A. viride, by resident botanists (e.g. Cambridge, Chester- Genista pilosa, Mertensia maritima, ton, Ely, Whittlesey) whereas the majority of Orobanche alba, Vicia orobus).
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