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Botanical Society of the British Isles

Patron: Her Majesty Queen Elizabeth the Queen Mother

Applications for mem bers'*» sould be addressed to the Hon. General Secretary,
‘atural History Museum, Cromwell Road, London,

ae Society’s Prospectus may be obtained.

» £69
Ae 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. R. Mill, 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).

Watsonia, 19, 1-19 (1992) ee

Distribution of Pteridophyta in Wales

G. HUTCHINSON and B. A. THOMAS

Department of Botany, National Museum of Wales, Cardiff, CF1l 3NP

ABSTRACT

New 10-km square maps are presented for pteridophytes in Wales. The time-spans used for these maps are
pre-1950, 1950-69 and 1970-1990.

INTRODUCTION

An interest by botany staff of the National Museum of Wales in recording the distribution of
pteridophytes in Wales has been evident since H. A. Hyde and A. E. Wade wrote the original edition
of Welsh Ferns. This descriptive handbook, published jointly by the National Museum of Wales and
the University of Wales in 1940, was written in response to a perceived need for a simple and
inexpensive text. Its usefulness outside the Principality was anticipated correctly by the inclusion of
brief accounts of British species not occurring in Wales. Notes were given on the distribution of the
species in Wales and lists included of specimens in the Welsh National Herbarium. Specimen lists
for the very common species were impractical so only vice-counties were listed.

This system was followed in the next three editions, published in 1948, 1954 and 1962, although
the third edition was largely a lithographical reprint of the second. There were, however, several
changes in the fourth edition which reflected an increased knowledge of fern distribution in Wales.
' The three species of Polypodium, the two subspecies of Asplenium trichomanes, and hybrids of
Asplenium and Polystichum were included for the first time. Details of Dryopteris hybrids were
expanded. The varieties of Asplenium, Athyrium, Ceterach, Cystopteris and Dryopteris, which had
been included earlier, were omitted.

The larger fifth edition (1969) was revised and expanded by S. G. Harrison, to include the
clubmosses, quillworts and horsetails. The sixth edition (1978) gave many more descriptions of
hybrids with corresponding details of their known distributions in Wales.

Throughout these six editions there was a marked increase in the number of listed herbarium
specimens, thereby giving greater information on species, subspecies and hybrid distributions in
Wales. For example, the number of listed specimens for Phegopteris connectilis increased from 20 to
51 and for Hymenophyllum wilsonii from 22 to 66. There is a clear increase in the number of sites
listed for many taxa and therefore a greater known distribution for them. More species lists simply
became too large so they were omitted when every vice-county in Wales was represented. The
authors also recorded the overall increase in the total number of specimens held in the herbarium.
Numbers of British ferns were given as 1,660 in 1940, 1,990 in 1948, 2,200 in 1954 and 2,260 in 1962,
then the number of British Pteridophyta as 3,200 in 1969 and 3,900 in 1978. The herbarium now
includes 6700 British Pteridophyta of which 3900 are Welsh.

Meanwhile, there was the general move towards recording the distribution of plants by the grid
. square system, rather than by vice-counties, and the publication of the Adlas of the British flora
(Perring & Walters 1962). The pteridophyte maps in the latter were, however, widely accepted to be
imperfect because they went to the printers within six years of the start of the mapping scheme
(Jermy et al. 1978). This was rectified considerably when the Atlas of Ferns of the British Isles was
published (Jermy ef al. 1978). Even so, some of these later maps were clearly still incomplete,
especially those of the commonly less well understood species, hybrids of Polypodium and
Dryopteris, and the subspecies of Asplenium trichomanes and Dryopteris affinis.

It was against this background of recording the distributions of pteridophytes that we commenced
work on a new edition of Welsh Ferns. Clearly, the idea of incorporating lists of herbarium

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. R. Mill, 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).

Watsonia, 19, 1-19 (1992) | es

Distribution of Pteridophyta in Wales

G. HUTCHINSON and B. A. THOMAS

Department of Botany, National Museum of Wales, Cardiff, CF1 3NP

ABSTRACT

New 10-km square maps are presented for pteridophytes in Wales. The time-spans used for these maps are
pre-1950, 1950-69 and 1970-1990.

INTRODUCTION

This system was followed in the next three editions, published in 1948, 1954 and 1962, although
the third edition was largely a lithographical reprint of the second. There were, however, several
changes in the fourth edition which reflected an increased knowledge of fern distribution in Wales.
. The three species of Polypodium, the two subspecies of Asplenium trichomanes, and hybrids of
Asplenium and Polystichum were included for the first time. Details of Dryopteris hybrids were
expanded. The varieties of Asplenium, Athyrium, Ceterach, Cystopteris and Dryopteris, which had
been included earlier, were omitted.

Throughout these six editions there was a marked increase in the number of listed herbarium
specimens, thereby giving greater information on species, subspecies and hybrid distributions in
Wales. For example, the number of listed specimens for Phegopteris connectilis increased from 20 to
51 and for Hymenophyllum wilsonii from 22 to 66. There is a clear increase in the number of sites
listed for many taxa and therefore a greater known distribution for them. More species lists simply
became too large so they were omitted when every vice-county in Wales was represented. The
authors also recorded the overall increase in the total number of specimens held in the herbarium.
Numbers of British ferns were given as:1,660 in 1940, 1,990 in 1948, 2,200 in 1954 and 2,260 in 1962,
then the number of British Pteridophyta as 3,200 in 1969 and 3,900 in 1978. The herbarium now
includes 6700 British Pteridophyta of which 3900 are Welsh.

Meanwhile, there was the general move towards recording the distribution of plants by the grid
. $quare system, rather than by vice-counties, and the publication of the Adlas of the British flora
(Perring & Walters 1962). The pteridophyte maps in the latter were, however, widely accepted to be
imperfect because they went to the printers within six years of the start of the mapping scheme
(Jermy et al. 1978). This was rectified considerably when the Atlas of Ferns of the British Isles was
published (Jermy et al. 1978). Even so, some of these later maps were clearly still incomplete,
especially those of the commonly less well understood species, hybrids of Polypodium and
Dryopteris, and the subspecies of Asplenium trichomanes and Dryopteris affinis.

It was against this background of recording the distributions of pteridophytes that we commenced
work on a new edition of Welsh Ferns. Clearly, the idea of incorporating lists of herbarium

I oy HUTCHINSON AND B. A. THOMAS

nor is it a suitable method of portraying the distribution of taxa. We
"yom9 } bution on a 10-km grid square system. With the completion of the
[/) { x. 987-88) and talk of a new atlas of the British flora, although funds
x )0 is a convenient date to end the current time span. These maps we
ene here, in advance ot a new Welsh Ferns, to ensure their early availability to a wide audience in
the hope that these will stimulate further interest in Welsh pteridophytes and their distribution.

COMPILATION OF MAPS

The base-maps used were those of Perring & Walters (1962) and Perring & Sell (1968) with
nomenclature modified according to Derrick et al. (1987). To these were added records held by the
Biological Records Centre, Monks Wood Experimental Station in November 1986 which included
all the records of Jermy et al. (1978). Information from all the specimens at NMW was checked as
was that from specimens of the more critical taxa at BM. Appropriate literature sources were also
checked. The records in the Welsh Plant Records data-files held at NMUW were added (including
records received which applied up to the end of 1990).

Updated maps were sent, for modification and comment, to all Welsh vice-county recorders and
several interested members of both the B.S.B.I. and the British Pteridological Society. However,
maps of those taxa considered to be rare were only sent on a vice-county basis to the relevant vice-
county recorder(s). Several records were added at this stage and recorders were also able to point
out former mapping errors. A healthy correspondence on queries followed, which included
obtaining more details from records at the Biological Records Centre. Generally, the wishes of vice-
county recorders were complied with, especially the removal of doubtful records. Untraceable
records were not used for the maps and on the boundary with England only records from the Welsh
side of the 10-km square were included. The 10-km square records which appeared in the Aiélas of
the British flora as 1930 onwards that could not be traced are given a separate symbol because they ~
cannot be assigned to either side of the 1950 time span boundary used in our maps.

Following this stage of the work it has been possible to add numerous records that resulted mostly
either from field work associated with current county flora surveys or determinations of new
specimens sent to NMW. Finally, the pteridophyte maps compiled from the B.S.B.I. Monitoring
Scheme (1987-1988) were added.

The maps are presented in Figs 1-84, arranged in alphabetical order. Symbols used for the maps
are as follows:—

@ 1970-1990

@ 1930-1969 untraced Atlas of the British flora record
© pre-1950

@ Introduced 1970-1990

<> Introduced pre-1970

Critical taxa have been looked at very carefully and many specimens of them pe ommine dl Maps for
the main ones, which are outlined here, are still considered incomplete.

Asplenium trichomanes L. Since the Atlas of the British flora, a new subspecies (A. trichomanes
subsp. pachyrachis (Christ) Lovis & Reichst.) and a nothosubspecies (A. trichomanes nothosubsp.
staufferi Lovis & Reichst.) have been identified for Wales (Rickard 1989). A. trichomanes subsp.
trichomanes is considered under-recorded especially in the southern half of Wales. New records for
this area were only accepted after microscopic measurement of the exospores according to the
method of Reichstein (Hegi 1984). The A. trichomanes distribution map is identical with that for A.
trichomanes subsp. quadrivalens D. E. Meyer emend. Lovis, and is omitted.

Dryopteris affinis (Lowe) Fraser-Jenkins. An understanding of Dryopteris affinis as comprising
three subspecies in the British Isles has only emerged in the second half of the 1980s (Fraser-Jenkins
1987). Var. paleaceo-lobata sensu Fraser-Jenkins is included under subsp. affinis and var. robusta
(Oberholzer & von Tavel ex Fraser-Jenkins) Fraser-Jenkins & Salvo under subsp. borreri
(Newman) Fraser-Jenkins, but records of these varieties were almost zero.

Polypodium vulgare L. agg. The records of Polypodium vulgare were treated with caution as many
applied to P. vulgare sensu lato. Where accurate identification could not be established records were

DISTRIBUTION OF PTERIDOPHYTA IN WALES S

treated as P. vulgare agg. For P. X font-queri Rothm. and P. x shivasiae Rothm. only records
confirmed by R. H. Roberts are mapped. Many of the records of the commonest hybrid, P x
mantoniae Rothm., have also been confirmed by R. H. Roberts.

Ophioglossum azoricum C. Presl. Only the records recognized by Paul (1987) are included.
Trichomanes speciosum Willd. The distribution map includes only records for sporophytes. Other
sites where only gametophytes have been found are not included.

ACKNOWLEDGMENTS

We thank all Welsh vice-county recorders, other interested members of the B.S.B.I. and the British
Pteridological Society for adding to the updated base-maps, for pointing out erroneous records and
answering numerous queries. C. D. Preston allowed access to the record-card files at the Biological
Records Centre, Monks Wood Experimental Station, provided a set of pteridophyte maps resulting
from the B.S.B.I. Monitoring Scheme (1987-1988) and answered numerous queries about the
records. A. C. Jermy allowed us to examine specimens in the cryptogamic herbarium, BM. Thanks
are due to referees for determinations and help with the taxonomy, especially C. R. Fraser-Jenkins
(Dryopteris affinis) and R. H. Roberts (Polypodium), and A. C. Jermy. We also thank the
numerous recorders who sent in specimens for determination and J. G. Gavan for preparing the
maps.

REFERENCES

Derrick, L. N., Jermy, A. C. & PAut, A. M., eds (1987). Checklist of European Pteridophytes. Sommerfeltia 6:
i-xx, 1-94.

FRASER-JENKINS, C. R. (1987). Dryopteris, in Derrick, L. N. et al., Checklist of European Pteridophytes.
Sommerfeltia 6: x—xiil.

Hea, G. (1984). Illustrierte Flora von Mitteleuropa, 3rd ed., 1(1): 215-216. Berlin & Hamburg.

Hype, H. A. & Wane, A. E. (1940, 1948, 1954, 1962). Welsh Ferns, 1st—4th eds. Cardiff.

Hype, H. A., WabE, A. E. & Harrison, S. G. (1969, 1978). Welsh Ferns, 5th & 6th eds. Cardiff.

Jermy, A. C., ARNOLD, H. R., FARRELL, L. & PERRING, F. H,. (1978). Atlas of Ferns of the British Isles. London.

Pau , A. M. (1987). The status of Ophioglossum azoricum (Ophioglossaceae: Pteridophyta) in the British Isles.
Fern Gazette 13: 173-187.

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.

RickarD, M. H. (1989). Two spleenworts new to Britain — Asplenium trichomanes subsp. pachyrachis and
Asplenium trichomanes nothosubsp. staufferi. Pteridologist 1: 244-248.

(Accepted August 1991)

APPENDIX: CHECKLIST OF WELSH PTERIDOPHYTES

Adiantum capillus-veneris L.
Asplenium adiantum- “nigrum L.
Asplenium adiantum-nigrum L. x A. septentrionale (L.) Hoffm. (A. X contrei Callé, Lovis & Reichst.)
Asplenium billotii F. W. Schultz
Asplenium ceterach L.
Asplenium marinum L.
Asplenium ruta-muraria L.
- Asplenium scolopendrium L.
Asplenium septentrionale (L.) Hoffm.
Asplenium septentrionale (L.) Hoffm. subsp. septentrionale xX A. trichomanes L. SSS. trichomanes (A. X
__ alternifolium Wulfen nothosubsp. alternifolium)
Asplenium trichomanes L.
Asplenium trichomanes L. subsp. pachyrachis (Christ) Lovis & Reichst.
Asplenium trichomanes L. subsp. pachyrachis (Christ) Lovis & Reichst. x A. trichomanes L. subsp.
quadrivalens D. E. Meyer emend. Lovis (A. trichomanes L. nothosubsp. staufferi Lovis & Reichst.)
Asplenium trichomanes L. subsp. quadrivalens D. E. Meyer emend. Lovis

2 G. HUTCHINSON AND B. A. THOMAS

specimens is no longer practical, nor is it a suitable method of portraying the distribution of taxa. We
decided instead to record distribution on a 10-km grid square system. With the completion of the
B.S.B.I. Monitoring Scheme (1987-88) and talk of a new atlas of the British flora, although funds
have not yet been allocated, 1990 is a convenient date to end the current time span. These maps we
present here, in advance of a new Welsh Ferns, to ensure their early availability to a wide audience in
the hope that these will stimulate further interest in Welsh pteridophytes and their distribution.

COMPILATION OF MAPS

The base-maps used were those of Perring & Walters (1962) and Perring & Sell (1968) with
nomenclature modified according to Derrick et al. (1987). To these were added records held by the
Biological Records Centre, Monks Wood Experimental Station in November 1986 which included
all the records of Jermy et al. (1978). Information from all the specimens at NMW was checked as
was that from specimens of the more critical taxa at BM. Appropriate literature sources were also
checked. The records in the Welsh Plant Records data-files held at NMW were added (including
records received which applied up to the end of 1990).

Updated maps were sent, for modification and comment, to all Welsh vice-county recorders and
several interested members of both the B.S.B.I. and the British Pteridological Society. However,
maps of those taxa considered to be rare were only sent on a vice-county basis to the relevant vice-
county recorder(s). Several records were added at this stage and recorders were also able to point
out former mapping errors. A healthy correspondence on queries followed, which included
obtaining more details from records at the Biological Records Centre. Generally, the wishes of vice-
county recorders were complied with, especially the removal of doubtful records. Untraceable —
records were not used for the maps and on the boundary with England only records from the Welsh
side of the 10-km square were included. The 10-km square records which appeared in the Adlas of
the British flora as 1930 onwards that could not be traced are given a separate symbol because they _
cannot be assigned to either side of the 1950 time span boundary used in our maps.

The maps are presented in Figs 1-84, arranged in alphabetical order. Symbols used for the maps
are as follows:—

@ 1970-1990

@ 1930-1969 untraced Adlas of the British flora record
© pre-1950

@ Introduced 1970-1990

> Introduced pre-1970

Critical taxa have been looked at very carefully and many specimens of them examined. Maps for
the main ones, which are outlined here, are still considered incomplete.

Polypodium vulgare L. agg. The records of Polypodium vulgare were treated with caution as many
applied to P. vulgare sensu lato. Where accurate identification could not be established records were

DISTRIBUTION OF PTERIDOPHYTA IN WALES 3

treated as P. vulgare agg. For P. X font-queri Rothm. and P. X shivasiae Rothm. only records
confirmed by R. H. Roberts are mapped. Many of the records of the commonest hybrid, P x
mantoniae Rothm., have also been confirmed by R. H. Roberts.

ACKNOWLEDGMENTS

REFERENCES

Derrick, L. N., JErMy, A. C. & PAut, A. M., eds (1987). Checklist of European Pteridophytes. Sommerfeltia 6:
i-xx, 1-94. :

FRASER-JENKINS, C. R. (1987). Dryopteris, in Derrick, L. N. et al., Checklist of European Pteridophytes.
Sommerfeltia 6: x—xiil.

Heai, G. (1984). Illustrierte Flora von Mitteleuropa, 3rd ed., 1(1): 215-216. Berlin & Hamburg.

Hype, H. A. & Wane, A. E. (1940, 1948, 1954, 1962). Welsh Ferns, 1st-4th eds. Cardiff.

Hype, H. A., Wapve, A. E. & Harrison, S. G. (1969, 1978). Welsh Ferns, Sth & 6th eds. Cardiff.

Jermy, A. C., ARNOLD, H. R., FARRELL, L. & PERRING, F. H. (1978). Atlas of Ferns of the British Isles. London.

Pau, A. M. (1987). The status of Ophioglossum azoricum (Ophioglossaceae: Pteridophyta) in the British Isles.
Fern Gazette 13: 173-187.

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.

RickarD, M. H. (1989). Two spleenworts new to Britain — Asplenium trichomanes subsp. pachyrachis and
Asplenium trichomanes nothosubsp. staufferi. Pteridologist 1: 244-248.

(Accepted August 1991)

APPENDIX: CHECKLIST OF WELSH PTERIDOPHYTES

Adiantum capillus-veneris L.

Asplenium adiantum-nigrum L.

Asplenium adiantum-nigrum L. x A. septentrionale (L.) Hoffm. (A. X contrei Callé, Lovis & Reichst.)

Asplenium billotii F. W. Schultz

Asplenium ceterach L.

Asplenium marinum L.

Asplenium ruta-muraria L.

_ Asplenium scolopendrium L.

Asplenium septentrionale (L.) Hoffm.

Asplenium septentrionale (L.) Hoffm. subsp. septentrionale x A. trichomanes L. suey trichomanes (A. X
alternifolium Wulfen nothosubsp. alternifolium)

Asplenium trichomanes L.

Asplenium trichomanes L. subsp. pachyrachis (Christ) Lovis & Reichst.

Asplenium trichomanes L. subsp. pachyrachis (Christ) Lovis & Reichst. x A. trichomanes L. subsp.
quadrivalens D. E. Meyer emend. Lovis (A. trichomanes L. nothosubsp. staufferi Lovis & Reichst.)

Asplenium trichomanes L. subsp. quadrivalens D. E. Meyer emend. Lovis

+ G. HUTCHINSON AND B. A. THOMAS

Asplenium trichomanes L. subsp. quadrivalens D. E. Meyer emend. Lovis x A. trichomanes L. subsp.
trichomanes (A. trichomanes L. nothosubsp. lusaticum (D. E. Meyer) Lawalrée)

Asplenium trichomanes L. subsp. trichomanes

Asplenium viride Hudson

Athyrium filix-femina (L.) Roth

Azolla filiculoides Lam.

Blechnum spicant (L.) Roth

Botrychium lunaria (L.) Swartz

Cryptogramma crispa (L.) Hook.

Cyrtomium falcatum (L. f.) C. Presl

Cystopteris fragilis (L.) Bernh.

Diphasiastrum alpinum (L.) Holub

Dryopteris aemula (Aiton) Kuntze

Dryopteris affinis (Lowe) Fraser-Jenkins

Dryopteris affinis (Lowe) Fraser-Jenkins subsp. affinis

Dryopteris affinis (Lowe) Fraser-Jenkins subsp. borreri (Newman) Fraser-Jenkins

Dryopteris affinis (Lowe) Fraser-Jenkins subsp. cambrensis Fraser-Jenkins

Dryopteris affinis (Lowe) Fraser-Jenkins x D. filix-mas (L.) Schott (D. X complexa Fraser-Jenkins)

Dryopteris carthusiana (Vill.) H. P. Fuchs

Dryopteris carthusiana (Vill.) H. P. Fuchs x D. dilatata (Hoffm.) A. Gray (D. x deweveri (Jansen) Jansen
Wachter)

Dryopteris dilatata (Hoffm.) A. Gray

Dryopteris dilatata (Hoffm.) A. Gray x D. expansa (C. Presl) Fraser-Jenkins & Jermy (D. < ambroseae Fraser-
Jenkins & Jermy)

Dryopteris expansa (C. Presl) Fraser-Jenkins & Jermy

Dryopteris filix-mas (L.) Schott

Dryopteris filix-mas (L.) Schott X D. oreades Fomin (D. X mantoniae Fraser-Jenkins & Corley)

Dryopteris oreades Fomin

Dryopteris submontana (Fraser-Jenkins & Jermy) Fraser-Jenkins

Equisetum arvense L.

Equisetum arvense L. X E. fluviatile L. (E. X litorale Kuhl. ex Rupr.)

Equisetum fluviatile L.

Equisetum hyemale L.

Equisetum palustre L.

Equisetum palustre L. x E. telmateia Ehrh. (E. x font-queri Rothm.)

Equisetum sylvaticum L.

Equisetum telmateia Ehrh.

Equisetum variegatum Schleicher ex Weber & Mohr

Gymnocarpium dryopteris (L.) Newman

Gymnocarpium robertianum (Hoffm.) Newman

Huperzia selago (L.) Bernh. ex Schrank & C. F. P. Mart.

Hymenophyllum tunbrigense (L.) Sm.

Hymenophyllum wilsonii Hook.

Isoetes echinospora Durieu

Isoetes lacustris L.

Lycopodiella inundata (L.) Holub

Lycopodium annotinum L.

Lycopodium clavatum L.

Matteuccia struthiopteris (L.) Tod.

Onoclea sensibilis L.

Ophioglossum azoricum C. Presl

Ophioglossum vulgatum L.

Oreopteris limbosperma (All.) Holub

Osmunda regalis L.

Phegopteris connectilis (Michx) Watt

Pilularia globulifera L.

Polypodium cambricum L.

Polypodium cambricum L. X P. interjectum Shivas (P. X shivasiae Rothm.)

Polypodium cambricum L. X P. vulgare L. (P. X font-queri Rothm.)

Polypodium interjectum Shivas

Polypodium interjectum Shivas X P. vulgare L. (P. X mantoniae Rothm.)

Polypodium vulgare L.

DISTRIBUTION OF PTERIDOPHYTA IN WALES

Polypodium vulgare L. agg.

Polystichum aculeatum (L.) Roth
Polystichum aculeatum (L.) Roth x P. setiferum (Forskal) Woynar (P. x bicknellii (Christ) Hahne)
Polystichum lonchitis (L.) Roth
Polystichum setiferum (Forskal) Woynar
Pteridium aquilinum (L.) Kuhn
Selaginella kraussiana (Kuntze) A. Braun
Selaginella selaginoides (L.) Link
Thelypteris palustris Schott

Trichomanes speciosum Willd.

Woodsia alpina (Bolton) Gray

Woodsia ilvensis (L.) R. Br.

7 6 i 2 4 6
+ +— —
| S/H | ie. S\J |
IE ! | sts t 9
v.c. 52, Ang. 3 : Sar
eer RS ae,
| v.c. 51, Flints:.
Wel 49, Caerns v.c.|50, Denbs.
4
all: = a
vic. 48,| Mer
a al at
{ a y
ae ail =| _| oO}
vic. 47) Monts.
af
| s/O |
S|N
+ t | 7
v.c.| 46, Cards. v.c.|43, Rads.
b +
S|M
+ + + +
4 v.c, 45, Pembs. v.c.|42, Brecs.
2 | yc. = Carme.
eS
e v.c.|35, Mons,
SR ly h i vic. 41,] Glam.
Tr ir
SiS |
i 1 d L | ! 7
4
0, a all H
SiT j
Sava shilel eel ipiak pup) aL 469)

Map of Welsh vice-counties and 10-km squares.

6 G. HUTCHINSON AND B. A. THOMAS

Ficures 1—6: 1. Adiantum capillus-veneris; 2. Asplenium adiantum-nigrum; 3. Asplenium adiantum-nigrum X A.
septentrionale (A. X contrei); 4. Asplenium billotii; 5. Asplenium ceterach; 6. Asplenium marinum.

DISTRIBUTION OF PTERIDOPHYTA IN WALES fl

au
| |

Ficures 7-12: 7. Asplenium ruta-muraria; 8. Asplenium scolopendrium; 9. Asplenium septentrionale,
10. Asplenium septentrionale subsp. septentrionale x A. trichomanes subsp. trichomanes (A. X alternifolium
nothosubsp. alternifolium); 11. Asplenium trichomanes subsp. pachyrachis; 12. Asplenium trichomanes subsp.
pachyrachis x A. trichomanes subsp. quadrivalens (A. trichomanes nothosubsp. stauffert).

8 G. HUTCHINSON AND B. A. THOMAS

Fig. 13

FicureEs 13-18: 13. Asplenium trichomanes subsp. quadrivalens; 14. Asplenium trichomanes subsp. quadrivalens
A. trichomanes subsp. trichomanes (A. trichomanes nothosubsp. lusaticum); 15. Asplenium trichomanes
subsp. trichomanes; 16. Asplenium viride; 17. Athyrium filix-femina; 18. Azolla filiculoides.

DISTRIBUTION OF PTERIDOPHYTA IN WALES

Figures 19-24: 19. Blechnum spicant; 20. Botrychium lunaria; 21. Cryptogramma crispa; 22. Cyrtomium
falcatum; 23. Cystopteris fragilis; 24. Diphasiastrum alpinum.

10 G. HUTCHINSON AND B. A. THOMAS

FicuREs 25-30: 25. Dryopteris aemula; 26. Dryopteris affinis; 27. Dryopteris affinis subsp. affinis; 28. Dryopteris
affinis subsp. borreri; 29. Dryopteris affinis subsp. cambrensis; 30. Dryopteris affinis x D. filix-mas (D. X
complexa).

DISTRIBUTION OF PTERIDOPHYTA IN WALES : 11

Ficures 31-36: 31. Dryopteris carthusiana; 32. Dryopteris carthusiana x D. dilatata (D. Xx deweveri);
33. Dryopteris dilatata; 34. Dryopteris dilatata x D. expansa (D. X ambroseae); 35. Dryopteris expansa;
36. Dryopteris filix-mas. —

12 G. HUTCHINSON AND B. A. THOMAS

Fig. 37

FiGuREs 37-42: 37. Dryopteris filix-mas x D. oreades (D. X mantoniae); 38. Dryopteris oreades; 39. Dryopteris
submontana; 40. Equisetum arvense; 41. Equisetum arvense Xx E. fluviatile (E. X litorale); 42. Equisetum
fluviatile.

DISTRIBUTION OF PTERIDOPHYTA IN WALES 13

2 3 4

FicurEs 43-48: 43. Equisetum hyemale; 44. Equisetum palustre; 45. Equisetum palustre x E. telmateia (E. X
font-queri); 46. Equisetum sylvaticum; 47. Equisetum telmateia; 48. Equisetum variegatum.

14 G. HUTCHINSON AND B. A. THOMAS

Figures 49-54: 49. Gymnocarpium dryopteris; 50. Gymnocarpium robertianum; 51. Huperzia_ selago;
52. Hymenophyllum tunbrigense; 53. Hymenophyllum wilsonii: 54. Isoetes echinospora.

DISTRIBUTION OF PTERIDOPHYTA IN WALES 15

Fig. 55

Figures 55-60: 55. Isoetes lacustris; 56. Lycopodiella inundata; 57. Lycopodium annotinum; 58. Lycopodium
clavatum; 59. Matteuccia struthiopteris; 60. Onoclea sensibilis.

16 G. HUTCHINSON AND B. A. THOMAS

FicuRES 61-66: 61. Ophioglossum azoricum, 62. Ophioglossum vulgatum; 63. Oreopteris limbosperma;
64. Osmunda regalis; 65. Phegopteris connectilis; 66. Pilularia globulifera.

DISTRIBUTION OF PTERIDOPHYTA IN WALES 17

Fig. 68

@8000000
nd ep ©0880

FIGURES 67-72: 67. Polypodium cambricum; 68. Polypodium cambricum x P. interjectum (P. X shivasiae);
69. Polypodium cambricum X P. vulgare (P. X font-queri); 70. Polypodium interjectum; 71. Polypodium
interjectum X P. vulgare (P. X mantoniae); 72. Polypodium vulgare.

18 G. HUTCHINSON AND B. A. THOMAS

Fig. 73

()
®
®
@
®
®
pOQO
O8

Ficures 73-78: 73. Polypodium vulgare agg.; 74. Polystichum aculeatum; 75. Polystichum aculeatum X
P. setiferum (P. X bicknellii); 76. Polystichum lonchitis; 77. Polystichum setiferum; 78. Pteridium aquilinum.

DISTRIBUTION OF PTERIDOPHYTA IN WALES

Fig. 79

Figures 79-84: 79. Selaginella kraussiana; 80. Selaginella selaginoides; 81.
82. Trichomanes speciosum, 83. Woodsia alpina; 84. Woodsia ilvensis.

Thelypteris palustris;

Watsonia, 19, 21-37 (1992) 21

Peroxidase isoenzyme and morphological variation in Sorbus
L. in South Wales and adjacent areas, with particular
reference to S. porrigentiformis E. F. Warb.

M. C. F. PROCTOR and A. C. GROENHOF

Department of Biological Sciences, University of Exeter, Hatherly Laboratories, Prince of Wales
Road, Exeter EX4 4PS

ABSTRACT

The delimitation of microspecies of Sorbus L. (Rosaceae) occurring in South Wales, the Wye Valley, the Bristol
area and Mendip is considered in relation to their peroxidase phenotypes and leaf, fruit and growth-habit
characters. On isoenzyme evidence, S. minima (A. Ley) Hedl., S. leyana Wilmott, S. anglica Hedl. and S.
bristoliensis Wilmott all appear essentially uniform and well delimited. S. xX vagensis Wilmott is variable in
peroxidase phenotype, as expected of a sexual hybrid. S. leptophylla E. F. Warb. appears genetically uniform,
but plastic in habit and leaf size. Its established distribution is limited to two localities in Brecs. (v.c. 42);
peroxidase evidence and fruit characters confirm a probable relationship to plants on Craig Breidden, Monts.
(v.c. 47). S. eminens E. F. Warb. from near Symonds Yat is genetically different from plants in the Bristol —
Mendip area, which show similarities in peroxidase, leaf, fruit and growth-habit characters to trees on the Menai
Straits and to S. hibernica E. F. Warb. Plants which have been named S. porrigentiformis E. F. Warb. in South
Wales include: (a) a widespread plant, of uniform peroxidase phenotype, with obovate leaves and broad crimson
fruits, typically forming a twiggy shrub with rather slender shoots (the holotype appears identical with this); and
(b) a much more local plant in the Mynydd Llangattock — Cwm Clydach area and the Black Mountains, of
different peroxidase phenotype, commonly growing into a small tree, with more oblong biserrate leaves and
larger subglobose scarlet fruits. Plants from near Symonds Yat growing with and somewhat resembling S.
porrigentiformis, differ from both (a) and (b) in peroxidase phenotype. Like S. porrigentiformis, they appear
more closely related to S. graeca (Spach) Kotschy than to S. aria (L.) Crantz, sensu stricto; they need further
study.

INTRODUCTION

The British species of Sorbus L. (Rosaceae) comprise three sexually reproducing diploids, S.
aucuparia L., S. aria (L.) Crantz and S. torminalis (L.) Crantz, and a number of polyploids all of
which are probably normally apomictic. These form populations of essentially identical individuals
which can be recognized as distinct microspecies (Warburg 1952, 1962; Perring & Sell 1968). The
microspecies distinguished in the British Isles fall into three groups. The largest number (making up
the S. aria aggregate) appear to be derivatives of diploid S. aria sensu lato. A group including S.
anglica Hedl. show characters intermediate between S. aria sensu lato and S. aucuparia; the S.
latifolia group are similarly intermediate between S. aria sensu lato and S. torminalis.

All but four of the described microspecies of Sorbus in the British Isles occur in the area around
the Bristol Channel. The sites at which they grow fall into three broad groups. In an eastern group of
sites, on the Carboniferous Limestone of Mendip, the Avon Gorge and the Wye Valley, the
apomicts occur in contact with sexual S. aria as well as S. aucuparia and S. torminalis. The other two
groups, on the Carboniferous Limestone crags of Breconshire and West Glamorgan and on the Old
Red Sandstone of the North Devon and West Somerset coast, are beyond the western limit of S. aria
and the Whitebeam populations appear to be wholly apomictic.

Sorbus in the Bristol Channel area thus presents a picture of some complexity, and potentially of
great interest as a model for studying the evolution and ecology of an apomictic group. Proctor et al.
(1989) investigated populations in Devon and West Somerset, and showed the usefulness of
peroxidase isoenzyme phenotypes in helping to delimit and characterize apomictic Sorbus
populations. Isoenzymes provide additional characters and make possible an interplay of hypothesis

22 M. C. F. PROCTOR AND A. C. GROENHOF

and test between biochemical and morphological evidence; they also extend the possibilities for
assessing genetic relationships between populations.

Warburg (1952, 1957) indicated the occurrence of some variation within S. anglica, S. eminens
E. F. Warb. and S. porrigentiformis E. F. Warb. The experience of a number of field botanists has
suggested that there are real difficulties in the characterization and identification of several of the
species, especially S$. porrigentiformis in South Wales. Warburg (1952) gave chromosome numbers
of 2n = 51 and 2n = 68 in that species. In this paper we present the results of a survey of peroxidase
phenotypes in apomictic Sorbus populations in South Wales and neighbouring areas, and consider
the results in relation to some morphological characters of the plants, and their geographical
distributions.

MATERIALS AND METHODS

PEROXIDASE ISOENZYMES

Shoot samples were collected in the field in the summers of 1989 and 1990, and brought back fresh to
the laboratory. The extracts used for electrophoresis were of ‘bark’ tissue scraped from (usually)
second-year shoots. They were made as soon as possible after collection, but we found that even in
summer shoots remained in good condition for a week or more if kept in polythene bags in a
refrigerator. The extracts were run on vertical polyacrylamide slab gels. Details of the methods used
are given by Proctor ef al. (1989), and the conventions used to describe peroxidase phenotypes here
are the same as in that paper. The bands obtained on the gels can conveniently (but arbitrarily) be
divided into six groups (Fig. 1). Two slow-running groups (A and B) and a fast-running band (F)
tend to be rather diffuse and present in most or all samples, so are of little taxonomic interest. Bands
in groups C and E are well defined, and occur in varying combinations in all the British apomictic
Sorbus species. One or two bands of group D, characteristic of S$. aucuparia, occur in the apomicts
of the S. anglica group.

LEAF, FLOWER AND FRUIT CHARACTERS

We collected herbarium material from most of the populations we examined; the specimens have
been retained by M.C.F.P. In addition, we collected and pressed a number of samples of
- representative individual leaves from short shoots; these were photocopied as a convenient means
for record and comparison. The colour of both leaf surfaces was recorded from some of the material
while still fresh by matching with the R.H.S. Colour Chart (Royal Horticultural Society 1966).
Fresh pollen samples from flowers collected in June 1989 were examined microscopically after
staining in aceto-carmine, and samples from the same inflorescences were tested for germination in
15% sucrose solution. Fruit was collected in September and early October. Length and breadth of
samples of well-formed fruits were measured to the nearest 0-2 mm using a sliding caliper-rule. To
provide a reproducible and quantitative basis for recording fruit colour, fresh fruits were matched
with a limited sub-set of the shades in the R.H.S. Colour Chart, chosen so that their coordinates in
the system of the Commission Internationale d’Eclairage (C.I.E.) formed a reasonably evenly
graded series. Individual fruits tend to run through the whole or a part of this series as they ripen;
broad differences between species are easily established, but for rigorous comparisons fruits need to
be at the same stage of ripeness. The C.1I.E. chromaticity and reflectance coordinates for the shades
used are given in Appendix 1.

RESULTS

The peroxidase phenotypes found in our material are summarized diagrammatically in Fig. 1, and —
the fruit measurements in Table 1.

S. ANGLICA AND ITS ALLIES
Samples of S. anglica from the Mynydd Llangattock area, the Wye Valley, the Avon Gorge,

PEROXIDASE ISOENZYME AND MORPHOLOGICAL VARIATION IN SORBUS L. 23

2 x2) ® ® @
3 S i elf i Ain
o =~ 3 one 3 Ke)
= oe = =X x x
ax =] (One) fe) fe)
— — - o fox a= _ P=
= = ® jor ro) > 0 Co) Co)
> > 2 iS Da Eo a a.
o © fo) = = Te! fe) fe)
— = G) o id wm om -
© o 2 gS Py oe oy ae)
> > 5 E a a) SS Shika pee
“ Be = S = & ® Woe sees
o 2 aS c Ss = es = CS Os Ora Osa. Ore
c iss] = aS) WY 7) c © ® O MGR iS ES ees D>
© en) = £ c c o > PN OL 1 UO! a Bio
8 o = 79 a 5 © o c S Ce) | SES 2 Sy Sis tay te
ro) BY £ Oo 5 2 S & = na fra ® ®
= = 2 3 £ .S & Le es | Si, 6 O oe oO
ee ee 8 Ree Gass as os
S x e iss 2 = o o : 5 Sa aay : mt :
, : : ; ; : roo) “ oO st Lo o
[i ee ee ey ee in Rees ee ee a ee
Py re ge ee eg i ol eM ee
C1 aE | Ge GE GE | GD) GED EER c=
= — Gp — eee | —— ——> soe eC CS
C4Au Pe eS => =>
x E3 a Eas | ee Pres a aR

Ficure 1. Schematic diagram of peroxidase phenotypes of apomictic Sorbus populations from South Wales and
adjoining areas. Conventions follow Proctor et al. (1989).

Cheddar, Breidden and Llangollen all show the same peroxidase phenotype as that found in a plant
of Avon Gorge origin by Proctor et al. (1989). The trees varied slightly in leaf shape but in general
had somewhat more angular and deeply incised lobes than plants from Devon (Fig. 2). There
appears also to be a consistent difference in fruit shape between trees on Breidden and those farther
south (Table 1).

The geographically very restricted S. minima (A. Ley) Hed1. (five trees) and S. leyana Wilmott
(three trees) show consistent and distinctive peroxidase phenotypes. In S. minima Proctor et al.
(1989) indicated a pair of bands in positions D2 and E1; the latter generally stains more strongly and
the two are often not clearly resolved. Pollen of both species showed virtually no germination in
15% sucrose solution, and a large proportion of misshapen and empty grains, suggesting that both
are probably triploid (Liljefors 1953).

THE S. LATIFOLIA GROUP
Four trees of S. bristoliensis Wilmott from the Leigh Woods side of the Avon Gorge were consistent
" in their peroxidase phenotype, erroneously shown by Proctor et al. (1989) as including band C4; this
is absent in S. bristoliensis, although the weak band CS is present. S. croceocarpa (Sell 1989), of
which there are planted and naturalized trees around Bristol, has a peroxidase phenotype differing
from that of S. devoniensis E. F. Warb. most conspicuously in the very much stronger band C3; the
same phenotype is found consistently in material from the type area in Anglesey and at Nantporth
on the Bangor side of the Menai Straits.

Four trees of S. X vagensis Wilmott all gave different peroxidase phenotypes, a result consistent
with the supposition that this is a sporadically occurring sexual hybrid.

24 M. C. F. PROCTOR AND A. C. GROENHOF

TABLE 1. FRUIT DIMENSIONS AND COLOURS FROM SORBUS POPULATIONS, SEPTEMBER-
OCTOBER 1989-90
Fruit dimensions are given for individual trees; n is the number of fruits measured. Length/breadth ratios were
calculated individually for each fruit. Fruit colour assessments are based on all the material available from a

Species and locality

S. aucuparia

Countisbury, N. Devon
(v.c. 4)

Lynmouth, N. Devon
(v.c. 4)

S. anglica
N. Whilborough Common,

S. Devon (v.c. 3) (i)
(ii)
Woody Bay, N. Devon
(v.c. 4)
Cheddar, N. Somerset (v.c. 6)
(i)
(ii)

Lover’s Leap, Mons. (v.c. 35)
Cwm Clydach, Glam. (v.c. 41)
Breidden, Monts. (v.c. 47)

(i)

(ii)

(iii)

(iv)

S. aria
Cheddar, N. Somerset (v.c. 6)
(i)
(ii)
(iii)
(iv)
Seven Sisters Rocks,
Herefs. (v.c. 36)

S. leptophylla
Craig-y-Rhiwarth, Brecs. (v.c. 42)
(i)

(iv)
Craig-y-Cilau, Brecs. (v.c. 42)

S. aff. leptophylla
Breidden, Monts. (v.c. 47)

S. wilmottiana
Bristol, W. Gloucs. (v.c. 34)

S. eminens

Bristol, W. Gloucs. (v.c. 34)

Leigh Woods, N. Somerset
(v.c. 6)

Worlebury, N. Somerset
(v.c. 6)

Length

mean (s.d.)

(mm)

10-2 (0-51)
9-9 (0-86)
11-0 (0-43)

10-8 (0-64)
10-9 (0-56)
10-7 (0-63)
10-9 (0-68)

10-1 (0-48)
10-6 (0-22)
11-7 (0-52)
11-2 (0-88)

12-4 (0-43)
14-1 (0-77)
12-3 (0-39)
11-4 (1-09)

12-8 (0-79)

14-7 (0-56)
14-3 (0-73)
14-4 (0-32)
13-3 (0-68)
14-8 (0-71)

14-3 (0-30)

10-7 (0-54)

13-8 (0-77)
12-0 (0 92)

12-7 (0-81)

Breadth

mean (s.d.) Length/breadth

(mm)

11-0 (0-46)
10-6 (0-50)
12-4 (0-62)

11-8 (1-07)
11-9 (0-29)
11-6 (0-94)
11-6 (0-63)

10-1 (0-44)
10-6 (0-61)
11-3 (0-50)
11-4 (0-68)

12-7 (0-56)
13-6 (0-68)
13-5 (0-55)
11-3 (0-97)

11-6 (1-25)

13-8 (0-64)
13-0 (0-61)
13-1 (0-52)
12-5 (0-43)
14-3 (0-63)

11-6 (0-75)

10-4 (0-61)

15-4 (0-64)
13-8 (0-91)

14-0 (0-62)

particular locality at the time of scoring.

mean (s.d.)

0-93 (0-036)
0-93 (0-051)
0-89 (0-045)

0-91 (0-039)
0-92 (0-038)
0-92 (0-024)
0-95 (0-080)

1-00 (0-037)
1-00 (0-047)

1-04 (0-059)
0-98 (0-052)

0:98 (0-035)
1-04 (0-048)
0:91 (0-045)
1-01 (0-034)

1-09 (0-079)

1-23 (0-067)

1-03 (0-035)

0-89 (0-029)
0-87 (0-045)

0-91 (0-057)

Fruit colour
(R.H.S. Colour Chart)

44A(-45A)

33A

444-454

(444-454

45A(-46A)

45A
44A(-45A)

(28A-)32A(-33A)

(44A-)45A

br)

44A-A5A

- (44A-)45A
(33A-)44A-45A(—46A)

45A+46A

PEROXIDASE ISOENZYME AND MORPHOLOGICAL VARIATION IN SORBUS L. Pre

TABLE 1. continued

Species and locality

Cheddar, N. Somerset (v.c. 6)
(i)
(ii)
(iii)
(iv)
Seven Sisters Rocks,
Herefs. (v.c. 36) (i)

(il)

S. aff. eminens

Bangor, Caerns. (v.c. 49)
(i)
(11)

S. hibernica

Dromineer, N. Tipperary
(v.c. H10)

Rathdrum, Co. Wicklow
(v.c. H20)

S. porrigentiformis
(‘Type (a)’)

Anstey’s Cove, S. Devon (v.c. 3)

Woody Bay, N. Devon (v.c. 4)
Cheddar, N. Somerset (v.c. 6)
Heale Ladder, N. Somerset
(v.c. 6)
Wick Rocks, N. Somerset
(v.c. 6)
Craig-y-Rhiwarth, Brecs.
(v.c. 42) (i)
(ii)
Craig-y-Cilau, Brecs. (v.c. 42)

Aberedw Rocks, Rads.
(v.c. 43)

(‘Type (b)’) |
Daren Disgwylfa, Brecs.
(v.c. 42) (i)
(ii)
(iii)
Blackrock, Glam. (v.c. 41)
Craig-y-Cilau, Brecs. (v.c. 42)
Cwm Clydach, Glam. (v.c. 41)

Taren-yr-Esgob, Mons./Brecs.
(v.c. 35/42)

10
10

Length
mean (s.d.)
(mm)

10-9 (1-12)
14-2 (1-06)
12-0 (0-32)
11-2 (0-38)

10-6 (0-39)
10-2 (0-19)

12-0 (0-66)
11-2 (0-39)

13-8 (0-65)

12-3 (0-46)

10-6 (1-11)
9-1 (0-68)
10-2 (0-55)

9-3 (0-18)
10-6 (0-42)

9-6 (0-50)
9-8 (0-49)
9-0 (0-52)

10-8 (0-62)

11-0 (0-61)
12-2 (0-67)
11-2 (1-15)
11-8 (0-24)
11-4 (0-59)

11-3 (0-70)
11-1 (0-42)
12-2 (0-64)
12-4 (0-75)

11-6 (0-27)

Breadth
mean (s.d.)
(mm)

13-2 (1-01)
16-5 (0-64)
14-3 (0-42)
13-4 (0-61)

11-5 (0-18)
11-6 (0-52)

13-4 (0-46)
13-2 (0-79)

14-8 (0-58)

13-2 (0-39)

132811622)
11-8 (0-58)
13-4 (0-93)

11-0 (0-17)

12-4 (0-38)

11-4 (0-40)
12-3 (0-37)
11-7 (0-40)

13-0 (0-86)

12-4 (0-45)
13-7 (0-48)
12-0 (0-82)
12-0 (0-58)
13-6 (0-40)

12-0 (0-69)
12-7 (0-12)
12-6 (0-40)
12-7 (0-75)

13-6 (0-33)

Length/breadth

mean (s.d.)

0-83 (0-056)
0-86 (0-031)
0-85 (0-030)
0-84 (0-029)

0-97 (0-034)
0-88 (0-043)

0:90 (0-044)
0-86 (0-031)

0-94 (0-032)

0-93 (0-028)

0-78 (0-067)
0-78 (0-045)
0:77 (0-035)

0-84 (0-041)
0-86 (0-033)

0-85 (0-049)
0-80 (0-035)
0:78 (0-034)

0-84 (0-042)

0-90 (0-028)
0:90 (0-042)
0-93 (0-043)
0-99 (0-035)
0-84 (0-026)

0-94 (0-064)
0-88 (0-025)
0-97 (0-037)
0-98 (0-022)

0-86 (0-011)

Fruit colour

(R.H.S. Colour Chart)

(44A-)45A—46A

45A(-46A)

(44A-)45A

46A
45A-46A-53A
46A-53A

(45A-)46A(-53A)
46A(-53A)
45A—46A

ASA

44 A—45A(-46A)

9°

45A
(33A-)44A(-45A)

45A

TABLE 1. continued

M. C. F. PROCTOR AND A. C. GROENHOF

Length Breadth
mean (s.d.) mean(s.d.) Length/breadth Fruit colour
Species and locality n (mm) (mm) mean (s.d.) (R.H.S. Colour Chart)
S. graeca, sensu lato,
Seven Sisters Rocks,
Herefs. (v.c. 36)
Peroxidase phenotype I

(i) 5 10:2 (0:37) 12-6 (0-57) 0-82 (0-039) 45A-46A

(ii) 15 . 10-7 (0:59) 12-9 (0-61) 0-83 (0-036) 46A

(iii) 6 10-9 (0-43) 13-3 (0-43) 0-82 (0-026) 3,

(iv) 5 9-6 (0-33) 12-5 (0-64) 0-78 (0-018) 5%

(v) 10 ~=10-1 (0-63) 12-2 (0-45) 0-83 (0-025) 46A-53A
Peroxidase phenotype II 5 11-0 (0-66) 12-2 (0-58) 0-90 (0-038) (45A-)46A
Peroxidase phenotype III 7. 10-8 (0:41) 11-9 (0-67) 0-91 (0-043) 46A

S. rupicola
Brixham, S. Devon (v.c. 3)

(i) 6 12-8 (0-58) 14-3 (0-50) 0-90 (0-021) (33A—)44A(-45A)

(ii) 10. 12-1 (0-91) 13-6 (0-87) 0-89 (0-026) fe

(iii) 10 ~=—-:11-4 (0-85) 13-5 (0-72) 0-85 (0-029) fe
Trentishoe, N. Devon (v.c. 4) 10 = 13-0 (0-62) 14-9 (0-60) 0-87 (0-041) 45A—46A
Breidden, Monts. (v.c. 47)

(i) 5 11-2 (0-36) 12-6 (0-56) 0-89 (0-015) —

(ii) 5. 10-5 (0:23) 11-7 (0-23) 0-90 (0-023) —

(iii) 5 11-0 (0-70) 12-3 (0-42) 0-89 (0-042) —
Tighnabruaich, Main Argyll

(v.c. 98) 1 12-0 (—) 12-8 (—) 0-94 (—) —

THE S. ARIA GROUP

S. leptophylla E. F. Warb. most characteristically forms a sprawling tree with long more or less
pendulous branches and large leaves (Fig. 3a—b), rooted into shady vertical limestone cliff faces.
Plants of this form grow at both of its two known Breconshire localities, and at both sites have the
same constant peroxidase phenotype. It is probably fortuitous that this matches that of “Taxon D’,
an unnamed apomict from the North Devon — West Somerset coast, resembling S. vexans E. F.
Warb. but with broader and darker red fruits (Proctor et al. 1989), as the two plants are quite
different in habit and leaf and fruit characters. At Craig-y-Rhiwarth S. leptophylla is the
predominant species (Fig. 3c—f), and the erect individuals in full sun at the top of the cliff appear at
first sight very different from those beneath the tree canopy. It is evidently plastic, and small trees
can be difficult to distinguish vegetatively from the biserrate-leaved porrigentiformis-like plant that
accompanies it at Craig-y-Cilau. We have no evidence of its occurrence anywhere in South Wales
other than the two localities mentioned. Pollen samples showed a high percentage of well-formed
grains and good germination in 15% sucrose solution.

Warburg (1957) indicated that S. leptophylla probably occurred also in Montgomeryshire (on
Craig Breidden). Some plants on the west crags of Breidden have leaves similar in shape to typical S.
leptophylla but smaller (Fig. 3g—h). One tree of this type that we sampled appears identical in
peroxidase phenotype with Breconshire S. leptophylla, and has fruits similar in size, shape, colour
and lenticel size and distribution. Specimens collected on the north crags of Breidden by P. J. M.
Nethercott in 1990 may also belong to the same form, their somewhat larger leaves reflecting
response to a shadier and less drought-stressed habitat.

S. wilmottiana E. F. Warb. (Warburg 1962, 1967) is a rare species of the Avon Gorge at Bristol.
We have been able to examine only two trees, which gave a distinctive peroxidase phenotype (C1 C2
El):

PEROXIDASE ISOENZYME AND MORPHOLOGICAL VARIATION IN SORBUS L. Dy

FiGure 2. Silhouettes of representative leaves. (a)—(f) S. anglica (a) N. Whilborough Common, S. Devon, v.c. 3,
(b) Woody Bay, N. Devon, v.c. 4, (c) Cheddar, N. Somerset, v.c. 6, (d) Lover’s Leap, Mons., v.c. 35, (e) Cwm
Clydach, Glam., v.c. 41 ,(f) Breidden, Monts., v.c. 47; (g) S. leyana, Daren Fach, Brecs., v.c. 42; (h) S. minima,
Craig-y-Cilau, Brecs. NECA?

i)
(o6)

M. C. F. PROCTOR AND A. C. GROENHOF

(g) (h)

Figure 3. S. leptophylla: silhouettes of representative leaves. (a) and (b) Craig-y-Cilau, Brecs., v.c. 42; (c)-(f)
Craig-y-Rhiwarth, Brecs., v.c. 42; (g) and (h) S. aff. leptophylla, Breidden, Monts., v.c. 47.

PEROXIDASE ISOENZYME AND MORPHOLOGICAL VARIATION IN SORBUS L. 29

(g) fo) 10 cm

[at ye Oh

Figure 4. Silhouettes of representative leaves of S. eminens sensu lato, and related populations. (a) Seven Sisters
Rocks, Herefs., v.c. 36 (Wye Valley phenotype (i)), (b) Seven Sisters Rocks (Wye Valley phenotype (ii)), (c)
Leigh Woods, N. Somerset, v.c. 6, (d) Bristol, W. Gloucs., v.c. 34, (e) Worlebury, N. Somerset, v.c. 6, (f)
Cheddar, N. Somerset, v.c. 6, (g) Nantporth, Bangor, Caerns., v.c. 49, (h) S. hibernica, Dromineer, N.
Tipperary, v.c. H10, (i) S. hibernica, Rathdrum, Co. Wicklow, v.c. H20.

30 M. C. F. PROCTOR AND A. C. GROENHOF
(c)
(f) 4

(i)

Figure 5. Silhouettes of representative leaves of S. porrigentiformis and related apomictic populations. (a)—(i) S.
porrigentiformis (Type (a)) (a) Babbacombe, S. Devon, v.c. 3, (b) Wick Rocks, W. Gloucs., v.c. 34, (c)
Cheddar, N. Somerset, v.c. 6, (d) Leigh Woods, N. Somerset, v.c. 6, (e) Seven Sisters Rocks, Herefs., v.c. 36,
(f) Nicholaston, Glam., v.c. 41, (g) Craig-y-Rhiwarth, Brecs., v.c. 42, (h) Craig-y-Cilau, Brecs., v.c. 42, (i)
Aberedw, Rads., v.c. 43;

PEROXIDASE ISOENZYME AND MORPHOLOGICAL VARIATION IN SORBUS L. 31

Fig. 5. continued

-(j)-(0) ‘large-leaved porrigentiformis’ (Type (b)) (j) Cwm Clydach, Glam., v.c. 41, (k) Daren Disgwylfa, Brecs.,
v.c. 42, (1) Craig-y-Cilau, Brecs., v.c. 42, (m) Taren-yr-Esgob, Mons./Brecs., v.c. 35/42, (n) Daren Lwyd,
Brecs., v.c. 42, (0) Cwmyoy, Mons., v.c. 35; (p) Bristol Type (c), Leigh Woods, N. Somerset, v.c. 6; (q)-(s) S.
graeca sensu lato, Seven Sisters Rocks, Herefs, v.c. 36, (q) peroxidase phenotype I, (r) peroxidase phenotype II,
(s) peroxidase phenotype III. Leaves (e), (0), (r) and (s) are from material collected at flowering time in late
May; their small size is mainly due to this, rather than inherent difference from other comparable populations.

o>)
bh

M. C. F. PROCTOR AND A. C. GROENHOF

S. eminens E. F. Warb. was described (Warburg 1952, 1957) as occurring in woods on the
Carboniferous Limestone of the Wye Valley and the Avon Gorge. Three trees from shady beech
woodland above Seven Sisters Rocks (Fig. 4a), and a small plant with similarly shaped but smaller
and firmer leaves in nearby open rocky scrub all showed a constant peroxidase phenotype (C1 C2
(C3) El E2); two further trees of similar leaf shape (Fig. 4b) differed in showing a stronger band C3.
Warburg (1957) stated that S. eminens around Symonds Yat has proportionately longer, more
rhomboid and more deeply-toothed leaves than plants from the lower Wye Valley or the Avon
Gorge. We have found no trees of S. eminens matching the Avon Gorge form in the lower Wye
Valley. Herbarium specimens from this area, including Warburg’s type in BM, appear essentially
similar to other Wye Valley material, and agree well with Warburg’s description and illustration.

Samples collected from a uniform population in the shady lower fringe of Leigh Woods in the
Avon Gorge showed a constant peroxidase phenotype (C2 C3 E1 E3). We found the same
peroxidase phenotype in uniform populations of trees in rocky grassland or open scrub near
Weston-super-Mare and in Cheddar Gorge. The Avon Gorge and Mendip populations are similar
in most leaf, fruit and growth-habit characters; the Mendip plants have rather firmer leaves with a
broadly cuneate base and their fruits have larger lenticels. Field observation suggests that the
difference in leaf texture may be mainly due to the more exposed habitat of the Mendip trees, but
that the difference in shape of the leaf base is, at least in part, genetic. Two other Sorbus populations
resemble the Avon Gorge and Mendip ‘S. eminens’ in peroxidase phenotype and leaf characters.
These are a population at Nantporth near Bangor, Gwynedd, which has been named S.
porrigentiformis but is clearly not that species, and the widespread Irish S. hibernica E. F. Warb.
Both have fruits similar in size, shape and colour to the Avon Gorge plant (Table 1).

S. porrigentiformis is by far the most widely recorded species in our area. There are at least four
clearly different peroxidase phenotypes which correlate with morphological characters (Fig. 5,
Table 1), and appear relevant to a satisfactory delimitation of this species.

(a) The most widespread, with peroxidase phenotype C2 C3 El E2, occurs throughout the recorded
range of the species from Torbay and Gower to Radnorshire and the Bristol-Mendip area. It is
typically an open shrub (rarely a small tree) with rather angular and ‘twiggy’ branching, and
relatively slender twigs. The leaves are dark green above (usually 147A on the R.H.S. Colour
Chart), obovate, with an entire cuneate base and almost simple outwardly directed teeth (Fig. 5a—i);
the fruit is broader than long (length/breadth ratio c. 0-75—0-85), and a deep crimson red when ripe.
Samples of pollen from Gower and Craig-y-Rhiwarth showed 72-84% of well-filled grains and
about 20% germination in 15% sucrose solution; Dr Q. O. N. Kay has obtained chromosome counts
of 2n = (67—)68 in plants from Woody Bay (v.c. 4), Wick Rocks (v.c. 34) and Craig-y-Cilau (v.c. 42)
(unpublished). The holotype of S. porrigentiformis in BM appears identical with this form.

(b) The plant which has been called ‘large-leaved porrigentiformis’ from Craig-y-Cilau and some
other crags in the Mynydd Llangattock area has a different peroxidase phenotype (C2 C3 (C4) E2
E3). It readily grows into a well developed tree, usually with rather more robust shoots. The leaves
are somewhat more oblong in outline, and tend to be biserrate (Fig. 5j-o); the fruits are rather
larger, subglobose (length/breadth ratio c. 0-85—0-95) and a brighter red. We have found material
with this peroxidase phenotype and morphological characters on Carboniferous Limestone at
Craig-y-Cilau and crags to the east on Mynydd Llangattock, in Cwm Clydach, and on Old Red
Sandstone at Taren-yr-Esgob, Daren Lwyd and near Cwmyoy in the Black Mountains. Dr Q. O.N.
Kay has obtained chromosome counts of 2n = 68 in plants from Cwm Clydach (v.c. 41) and Daren
Disgwylfa (v.c. 42). The population at Taren-yr-Esgob has leaves with a more narrowly cuneate
base and sharper and more deeply incised teeth (Fig. 5m). The leaf-shape difference appears to be
correlated with a consistently clearer separation between peroxidase bands E2 and E3 in this
population than elsewhere, but both differences are rather slight.

(c) A morphologically uniform population of rather small trees in the Avon Gorge shows
consistently the peroxidase phenotype C2 (C3) (E2) E3. In leaf shape (Fig. 5p) and habit this plant
could be mistaken at first sight for $. porrigentiformis, but it has lighter green leaves (usually 146A
on the R.H.S. Colour Chart) which more nearly resemble a small S. aria. The Bristol ‘porrigentifor- —
mis’ discussed by Proctor et al. (1989) is this plant.

(d) Plants on Seven Sisters Rocks near Symonds Yat in the Wye Valley, growing with S.
porrigentiformis and showing a general resemblance to it in leaf-shape (Fig. 5q—s), habit, and fruit
shape and colour have the peroxidase phenotype Cl C2 (C3) El E2; several morphologically

PEROXIDASE ISOENZYME AND MORPHOLOGICAL VARIATION IN SORBUS L. 33

different groups of individuals differ in the relative intensity of the bands. Similar phenotypes occur
in S. eminens from the same area, and appear to be common in crimson-fruited forms of S. aria
sensu lato, in the Wye Valley generally. These plants cannot be genetically identical with any of the
other S. porrigentiformis-like populations we have examined.

The very widely distributed S. rupicola (Syme) Hed. is rather sporadically scattered in our area.
Samples examined from Gower, Craig-y-Cilau and Craig Breidden agree in peroxidase phenotype
with material from Devon, Yorkshire and Scotland (Proctor et al. 1989 and popup ished data).

ANOMALOUS INDIVIDUALS

We have not searched systematically for aberrant individuals within otherwise uniform apomictic
populations, but a few examples have come to light in the course of our sampling. One individual at
the top of Craig-y-Rhiwarth, in a population mainly of S. leptophylla with a few S. porrigentiformis
(type (a) above), has distinctive ovate leaves and gave a peroxidase phenotype (C2 E1 E3) different
from either. A tree below Leigh Woods, Bristol, with long cuneate-based many-veined leaves,
clearly differing from S. aria or any of the named apomicts, also has a unique peroxidase phenotype.
A single tree with rather narrow rhomboid leaves at Cwmyoy gave a peroxidase phenotype similar
to but not identical with S. wilmottiana.

DISCUSSION

Our results are consistent with accepted taxonomic views for the more geographically restricted
apomictic species, namely S. minima, S. leyana, S. bristoliensis, S. leptophylla and S. wilmottiana.
The peroxidase isoenzyme variation confirms that S. x vagensis is probably a sporadically occurring
sexual hybrid. However, the most interesting findings relate to S. eminens and S. porrigentiformis.

The isoenzyme evidence confirms the difference between S. eminens from the Avon Gorge and
material from the Symonds Yat area (Fig. 6). Indeed the difference in peroxidase phenotype
suggests that the two populations may not be very closely related. On the other hand, the peroxidase
evidence links the Avon Gorge form with trees on Mendip, with the population that has been named
S. porrigentiformis on the Menai Straits, and with S. hibernica. Warburg verbally suggested an
affinity between a specimen from Cheddar and S. hibernica, without identifying it with that species
(P. J. M. Nethercott, in litt.). These plants all show similarities in leaf form and toothing, and in
growth habit, and their relationships need further investigation.

Our observations on S. porrigentiformis and related plants show why there have been problems in
delimiting this species in South Wales, and suggest how these may be resolved. The widespread
plant (type (a) above; Fig. 7) corresponds to the generally accepted concept of S. porrigentiformis
(Warburg 1952, 1957; Butcher 1961) and appears identical with the Wye Valley specimen on which
Warburg typified this species (Offa’s Dyke, Tidenham, 18 September 1933, A. J. Wilmott (no. 4484)
(BM)). It is locally accompanied or replaced in the Mynydd Llangattock-Cwm Clydach area and the
Black Mountains by genetically distinct populations (type (b); Fig. 8) which should be regarded as
an independent taxon. The uniform but probably more local plant in the Leigh Wood quarries (type
(c)) needs further study in the context of other Avon Gorge, Mendip and Wye Valley populations.

Warburg & Karpati (1968) associate S. porrigentiformis with S. graeca (Spach) Kotschy rather
than with S. aria sensu stricto. The S. porrigentiformis-like plants which accompany true S.
porrigentiformis near Symonds Yat are clearly genetically different from any we have examined
elsewhere, but they share S. graeca-like characters (cuneate-based obovate leaves and crimson
subglobose fruits with few lenticels) with S. porrigentiformis and a number of our other apomicts.
Field observations and herbarium material both indicate the existence of much variation in the Wye
Valley. In South Wales, as in South-west England, the peroxidase variation in Sorbus seems nicely
matched to the morphological variation and to the needs of recognizing and delimiting the apomictic
populations. In the Symonds Yat area it appears that this is not so, and that considerable
morphological variation may be accompanied by little or no variation in the peroxidase enzymes,
especially among the more S. graeca-like plants. In general, if two individuals show different
isoenzyme phenotypes under the same conditions, they cannot be genetically identical, but the fact
that two individuals are identical with respect to a particular enzyme system does not necessarily

34 M. C. F. PROCTOR AND A. C. GROENHOF

Ficure 6. Distribution of S. eminens E. F. Warb. in 2-km squares (tetrads) of the National Grid. @ Symonds
Yat-type peroxidase phenotype; A Avon Gorge-type peroxidase phenotype; O other records of S. eminens
sensu lato. The O shows the locality of the holotype specimen.

mean that they are identical in other respects. More work is needed on the Wye Valley populations,
and exploration of other enzyme systems may well prove useful in this.

The apomictic Sorbus species we have considered are all confined to crags and rocky ground, an
essentially disjunct habitat. Their present distribution must reflect an interplay between colonization
and extinction on these ‘islands’ in a ‘sea’ of country they cannot colonize (Macarthur & Wilson
1967). S. rupicola has by far the widest total range (Perring & Sell 1968), in which wide disjunctions
suggest fragmentation of a once more continuous area, perhaps in the more open landscape of the
early Post-glacial (Pigott & Walters 1954; Birks 1973; Boyd & Dickson 1987). Next most widespread
are S. porrigentiformis and S. anglica, both with much more coherent distributions centred on the
South Wales limestone. Some of the more restricted distributions may have relic features, but to a
great extent we are probably looking at rather recent patterns of bird dispersals (compare the
patterns in Rubus of Weber (1987)) from centres of origin concentrated in three main areas, the Avon
Gorge and Wye Valley, the Mynyd Llangattock area, and the Exmoor coast. From the peroxidase
data, it is tempting to speculate that S. porrigentiformis was involved in the origin of S. anglica, S.
leyana, S. minima, S. bristoliensis (Sell 1989) and other apomictic populations within its area (but
excluding S. leptophylla and the North Devon apomicts); farther north, S. rupicola was probably the
S. aria-group parent of S. arranensis and S. pseudofennica (Proctor et al. 1989). However, the
inheritance of peroxidases in Sorbus is unknown, and likely to be complex (Gottlieb 1981), so these
are no more than tentative conjectures which should stimulate search for other evidence.

It was Linnaeus’s precept that the first step in understanding is to know the entities one is dealing
with. However, the interest of Sorbus only begins with the delimitation and enumeration of
apomictic microspecies. A far more interesting challenge is to understand how an apomictic group

PEROXIDASE ISOENZYME AND MORPHOLOGICAL VARIATION IN SORBUS L. 35

Figure 7. Distribution of S. porrigentiformis E. F. Warb. in 2-km squares (tetrads) of the National Grid.
@ peroxidase phenotype (a); O tetrads from which we have seen material which on morphological characters
appears certainly to be this type. O shows the location of the holotype specimen. Outside the area of this map, S.
porrigentiformis confirmed as peroxidase phenotype (a) occurs in v.c. 3 in tetrads SY82.70, 92.62 and 92.64.

of this kind came into being, under what selection pressures, and how it functions over an extended
span of time (Gustafsson 1947; Clausen 1954; Asker 1979). In this quest, fast-growing herbaceous
plants (e.g. Taraxacum (Richards 1970a, b, 1973; Ford 1981)) and the long-lived populations of
Sorbus offer complementary opportunities. In the latter genus, many questions of cytogenetics,
population biology and ecology invite investigation, both in the apomictic populations of South
Wales and South-west England, and in the mixed sexual and apomictic populations of the Wye
Valley, Avon Gorge and Mendip which may well provide models for the kind of situation in which
many of our present-day Sorbus apomicts originated.

ACKNOWLEDGMENTS

This work would not have been possible without the help and cooperation of many people who have
given us information about localities, answered our queries, sent us Sorbus material, or granted us
permission to visit sites, particularly Miss Gill Barter, Mr D. Doogue, Mr T. J. Evans, Dr G.

Howells, Dr Q. O.N. Kay, Dr D. L. Kelly, Dr H. McAllister, Miss Vicky Morgan, Mr P. D. Sell,

Mr A. McG. Stirling, Dr G. B. Wilson and Mr R. G. Woods. We are grateful to them all. Our
especial thanks are due to Mr P. J. M. Nethercott and Mr M. Porter for their generosity in sharing
their knowledge and much time in the field with us, and to Dr Margaret E. Proctor without whose
work on Devon Sorbus this project would never have begun.

36 M. C. F. PROCTOR AND A. C. GROENHOF

e—> f} Ny p

a eS
=

ae /2

sr Bee
Ficure 8. S. porrigentiformis sensu lato. Distribution of peroxidase phenotype (b) (‘large-leaved porrigentifor-
mis’) in 2-km squares (tetrads) of the National Grid. © population at Taren-yr-Esgob.

REFERENCES

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BritisH CoLour Councit (1938). Horticultural Colour Chart. London.

BuTcHER, R. W. (1961). A new illustrated British Flora. London.

CLAUSEN, J. (1954). Partial apomixis as an equilibrium system in evolution. Atti del LX Congresso Internationale
di Genetica. I. Caryologia 6 (suppl.): 469-479.

Forp, H. (1981). The demography of three populations of dandelion. Biol. J. Linn. Soc. 15: 1-11.

Gort .ies, L. D. (1981). Electrophoretic evidence and plant populations. Progress in Phytochemistry 7: 1—46.

Gustarsson, A. (1947). Apomixis in higher plants. II. The causal aspect of apomixis. III. Biotype and species
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Lityerors, A. (1953). Studies on propagation, embryology and pollination in Sorbus. Acta Horti Bergiani 16:
227-329.

MacartTHur, R. H. & Witson, E. O. (1967). The theory of island biogeography. Princeton.

PERRING, F. H. & SELL, P. D. eds. (1968). Critical supplement to the Atlas of the British Flora. London.

Picotr, C. D. & Watters, S. M. (1954). On the interpretation of the discontinuous distributions shown by
certain British species of open habitats. J. Ecol. 42: 95-116.

Proctor, M. C. F., Proctor, M. E. & GRoENHOF, A. C. (1989). Evidence from peroxidase polymorphism on the
taxonomy and reproduction of some Sorbus populations in south-west England. New Phytol. 112: 569-575.

Ricuarps, A. J. (1970a). Eutriploid facultative agamospermy in Taraxacum. New Phytol. 69: 761-774.

Ricuarps, A. J. (1970b). Hybridization in Taraxacum. New Phytol. 69: 1102-1121.

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Ricuarps, A. J. (1973). The origin of Taraxacum agamospecies. Bot. J. Linn. Soc. 66: 189-211.

Roya HortTIcULTURAL SocIETY (1966). R.H.S. Colour Chart. London.

SELL, P. D. (1989). The Sorbus latifolia (Lam.) Pers. aggregate in the British Isles. Watsonia 17: 385-399.

Waresure, E. F. (1952). Sorbus L., in CLAPHAM, A. R., Tutin, T. G. & WARBuRG, E. F. Flora of the British Isles,
Ist ed., pp. 539-556. Cambridge.

Warburg, E. F. (1957). Some new names in the British flora [Sorbus]. Watsonia 4: 43-46.

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Warsurc, E. F. (1967). Sorbus wilmottiana E. F. Warburg, sp. nov., in SELL, P. D. Taxonomic and
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(Accepted April 1991)

APPENDIX

The table below gives the C.I.E. coordinates of the shades from the R.H.S. Colour Chart used in matching fruit
colours for Table 1, and the corresponding colour names from the earlier Horticultural Colour Chart (British
Colour Council 1938). The first two figures of the C.I.E. coordinates define the chromaticity (hue and
saturation) of the colour on a modified colour triangle; the third coordinate measures percentage reflection.
Colours 28A-33A appear in the ‘orange-red group’ and colours 44A-53A in the ‘red group’ of the R.H.S.
Colour Chart; 45A is a nearly pure red. For further explanation see Royal Horticultural Society (1966).

Number in R.H.S. C.I.E. coordinates Reflection
Colour Chart Colour name X y factor (%)
28A Persimmon Orange 0-547 0-391 45-5
32A Indian Orange 0-561 0-370 27-6
33A Capsicum Red 0-567 0-355 23-6
44A [unnamed] 0-592 0-339 14-6
45A Guardsman Red 0-593 0-315 IES
46A Currant Red 0-546 0-310 9-4

53A Cardinal Red . 0-522 0-298 8-3

Saad aaa ith ac

Watsonia, 19, 39-41 (1992) 39

Short Notes

OROBANCHE ALBA STEPH. EX WILLD. IN FIFE (V.C. 85)

Comment is necessary on “An account of Orobanche L. in Britain and Ireland” by Rumsey & Jury
(1991), particularly on the section dealing with O. alba Steph. ex Willd. Here, the authors state that
“it is now believed absent from the east coast of Britain” and lament its disappearance from Fife. I
may have been the source of this belief, as in Ballantyne (1970) I said — rashly as it turns out — that
the species was extinct in all its former v.c. 85 stations.

Happily this is not the case. In view of the confusion, it is as well to document the plant’s localities
in Fife. These are all on the north shore of the Firth of Forth, from west to east:—

GR NT1.8: East of St David’s, 1820 (E); Cliffs below St David’s, ‘fine specimens” (Graham
1840); 1969, J. Carlyle (pers. comm.). This locality is a small rocky bay with cliffs of 15-20 m
between Inverkeithing and Dalgety Bay new town. I did not see O. alba on a visit on 3 August 1979
but as I did not search the area thoroughly, it may well still be extant.

GR NT2.8: (a) Sibbald (1684) stated that he found ‘“‘Orobanche major . . . Broomrape upon the
Buck [back] of Bruntisland’’; in 1710 he rephrased this comment to read: “I found it below a rising
ground upon the north side of the town of Bruntisland” (Sibbald 1803). (The earlier description is
repeated by several authors, e.g. Greville (1824).) Both Sibbald’s localities almost certainly refer to
an area near to or on Burntisland Binn, a low hill some 200 m high overlooking the town. Over two
centuries elapsed before the next record from here — a specimen in E dated 1906 says ‘‘Burntisland’’;
more specifically, a year later another observer stated ‘“‘near the Binn, Burntisland, July 1907 (J.
Anderson) and later, W. E. Evans” (in an annotated copy of Balfour & Sadler (1871)). There seems
to be no further records from this site.

(b) A separate locality from (a) appears to be c.2-2-5 km to its east, between Burntisland and
Kinghorn. This is variously given as follows:— near Pettycur, 1837, KNS (Young 1936); “‘. . . walked
to hills east of Burntisland, . . . picked Orobanche rubra’’, 1854 (Balfour, 1902); near Kinghorn,
1892 (STA); Kinghorn, 1904/07 (E); ‘‘on broken ground between Kinghorn and Burntisland”’, 1907
(E). There are no further records from this area, which is now occupied by a large caravan site,
although neighbouring cliffs (with difficult access) may still provide a haven.

(c) Perhaps the best known site is on top of a grassy cliff at Abden near the old Kinghorn
Poorhouse, about half-way between Kinghorn and Kirkcaldy; this is usually referred to as near
Seafield Tower, which is some 400 m to the north. The first mention is in 1809 (Maughan, 1811 and
in E) and there are records at regular intervals since, mainly in E and BM, to 1903; by this time, it
appeared to be almost extirpated, as described by Blackstock, a local naturalist. Commenting on the
“rooting-out” of Fife’s rare plants, Blackstock (1895) continued ‘Fifteen years ago, Orobanche
rubra {etc.| where plentiful in well-defined stations along the Fife coast. Five years ago, I counted
twenty good specimens of the Broomrape at its station near Kinghorn. . . Last year I saw two plants
— this year [1893] I failed to discover any”’. Ten years later, in 1903, another local botanist said of the
same site: “‘. . . if fortune specially favours us, we may gather a single specimen of the rarest plant in
Fife, the red Broomrape”’ (Young 1903). It was these observations that led me to believe that the
species had disappeared about the turn of the century. However, in 1971 J. E. Lousley (pers.
comm.) wrote to me to say that he had found the plant on 29 July 1938 (specimen in RNG). I again
searched for it, as I had done in the 1960s, without success; then, late in 1985 I was told it had been
seen during that summer, and on 23 July 1986 I came across three stunted stems. I have not had the
opportunity of returning since.

GR NT2.9: An open circle appears on the map of O. alba provided by Rumsey & Jury (1991) for
this square but there is none in the Adlas (Perring & Walters 1962) and I know of no locality. This
record is presumably based on the specimens collected by W. MclIvor from “Kirkcaldy” (MANCH,
1847; STA, undated). Nominally this is in this square but the actual station is almost certainly Abden
(above).

40 SHORT NOTES

GR NO4.0: In the New Statistical Account (1845) for the parish of Kilconquhar the compiler (the
local minister) in 1837 listed several plants from “‘under Kincraig Hill, on rocks”, among them O.
rubra. While this site is quite suitable, there is no other record from a locality which became well
explored. In particular, Charles Howie, who lived not far away and who botanized in east Fife from
c. 1834-84, does not mention the species from here. Additionally the minister’s botanical
knowledge must be questioned as he includes, from elsewhere in the parish, such improbable
species as Silene acaulis, Azalea procumbens and Adiantum capillus-veneris! The record in Wood
(1862/87) is based on the N.S.A., as is the “Elie” of Sonntag (1894). It seems best to discount this
locality.

To summarize, O. alba has been definitely recorded in four separate localities in v.c. 85. In two of
these it is probably extinct; in the other two, it seems to be surviving, although in small numbers. It
needs to be refound in the St David’s station to establish its status there, while the Abden site
requires to be monitored regularly because of the very small size of the population there.

I am grateful to D. R. McKean and A. Angus for checking specimens in E and STA respectively.

REFERENCES

Bacrour, I. B. (1902). Botanical excursions made by Professor John Hutton Balfour, 1846-1878. Notes Roy.
Bot. Gard. Edinb. 2: 175.

Ba.rour, J. H. & SADLER, J. (1871). The flora of Edinburgh, 2nd ed. Edinburgh.

BALLANTYNE, G. H. (1970). The flowering plants of Kirkcaldy and District. Kirkcaldy.

BiackstTock, W.S. (1895). Plants of Kirkcaldy. Proc. East of Scotland Union of Natural History Societies: 61-65.

GRAHAM, R. (1840). Notice of recent excursions in the neighbourhood of Edinburgh. Trans. Proc. bot. Soc.
Edinb. 1: 63.

GREVILLE, R. K. (1824). Flora Edinensis. Edinburgh.

MauGuHan, R. (1811). List of the rarer plants observed in the neighbourhood of Edinburgh. Mem. Wernerian
Nat. Hist. Soc. 1: 234.

New STATISTICAL ACCOUNT (1845). Parish of Kilconquhar (by Rev. W. Fernie) 9: 323.

PERRING, F. H. & WALTERS, S. M. eds. (1962). Atlas of the British flora. London.

Rumsey, F. J. & Jury, S. L. (1991). An account of Orobanche L. in Britain and Ireland. Watsonia 18: 257-295.

SIBBALD, SiR R. (1684). Scotia Illustrata. Edinburgh.

SIBBALD, Sir R. (1803). The history, ancient and modern, of the sherriffdoms of Fife and Kinvois: . . New ed. [of
1710 ed. ] Cupar.

SONNTAG, C. O. (1894). A pocket flora of Edinburgh. Edinburgh.

Woop, W. (1862/87). The East Neuk of Fife. ist & 2nd eds. Edinburgh.

Youna, W. (1903). Botanical Sketch. Davidson’s Illustrated Guide to Kirkcaldy and Neighbourhood: pp.83-87.

Youna, W. (1936). List of the flowering plants and ferns recorded from Fife and Kinross (v.c. 85). Trans. Proc.
bot. Soc. Edinb. 32: 1-173.

G. H. BALLANTYNE
Branksome, 193 Nicol St., Kirkcaldy, Fife, KY1 IPF

RE-ESTABLISHMENT OF A DISCARDED NAME IN CREPIS L. (ASTERACEAE)

While reviewing Linnaean names associated with temperate eastern North American species of

Hieracium and Crepis I chanced upon Crepis vesicaria L. subsp. haenseleri (Boiss. ex DC.) P. D.

Sell (1976), a name in current use (e.g. Clapham, Tutin & Moore 1987). The name is no longer

correct because of modifications made in Art. 57 of the 1983 Sydney Code (cf. Reveal 1983a)

conferring priority and transferability to autonyms, a concept supported by Sell when he reluctantly
proposed this combination. Accordingly, the following change is mandated.

Crepis vesicaria L. subsp. taraxacifolia (Thuill.) Thell. in Schinz & R. Keller, Fl. Schweiz ed. 3, 2:
361 (1914), autonym priority established by C. taraxacifolia subsp. haenseleri (Boiss. ex. DC.)
Nyman, Consp. 459 (1879).

Crepis taraxacifolia Thuill., Fl. Env. Paris ed. 2, 409 (1799).

Barkhausia haenseleri Boiss. ex DC., Prodr. 7: 153 (1838).

C. vesicaria subsp. haenseleri (Boiss. ex DC.) P. D. Sell in Bot. J. Linn. Soc. 71: 254 (1976).

SHORT NOTES 41

C. vesicaria var. taraxacifolia (Thuill.) B. Boivin in Naturaliste Canad. 94: 523 (1967).

It is possible that var. taraxacifolia is not the correct name for the taxon at this rank. Until there is
a catalogue of all infraspecific names for Old World plants (Reveal 1983b) it will be difficult to
ascertain autonymic synonymy as required by the Code.

ACKNOWLEDGMENTS

I wish to thank J. R. Press and E. Groves of The Natural History Museum, London, for their
assistance. Work on the typification of temperate eastern North American plants is supported by
National Science Foundation Grant BSR-8812816. This is Scientific Article A-6199, Contribution
No. 8368, of the Maryland Agricultural Experiment Station and Cooperative Extension Service.

REFERENCES

CrapHaM, A. R., TutIn, T. G. & Moore, D. M. (1987). Flora of The British Isles, 3rd ed. Cambridge.

REVEAL, J. L. (1983a). The Demoulin Rule and newly mandated combinations in Eriogonum (Polygonaceae).
Taxon 32: 292-295.

REVEAL, J. L. (1983b). The need for an index to infra-taxonomic entities. Taxon 32: 622.

SELL, P. D. (1976). Taxonomic and nomenclatural notes on the Compositae subfam. Cichorioideae. Bot. J.
Linn. Soc. 71: 236-267.

J. L. REVEAL
Department of Botany, University of Maryland, College Park, MD 20742-5815, U.S.A.

Watsonia, 19, 43-51 (1992) 43

Book Reviews

Plant form. An illustrated guide to flowering plant morphology. A. D. Bell. Pp. xiiit+341, with
numerous colour photographs and line drawings. Oxford University Press, Oxford. 1991. Price £50
hardback (ISBN 0—19-854279-8); £25 paperback (ISBN 0—19-854219-4).

This book is a mine of information on morphological terms, many of them not well known, at a time
when morphology is seldom taught in detail. There is a good index and an excellent cross-
referencing system, which, however, makes the book harder to read right through.

The author states that he has tried to make the book attractive, “‘the better to woo the budding
botanist and the curious amateur plantsman’’. He has certainly succeeded in his primary aim. The
book is laid out in such a way that a topic is dealt with in one or more double page spreads. Usually
on the left there is a colour photograph and descriptive paragraphs: on the right-hand page are
excellent line drawings by Alan Bryan, relating to the same topic. This results in a strangely shaped
book, wider than it is long. For the most part the colour photographs are excellent, although in my
copy, at least, a few were over-exposed (Figs 62b, 74, 170 and especially 134). In many cases they
could have been made more useful, especially to the novice, by the use of arrows or labels. How will
the “budding botanist’? know, for example, that Epiphyllum in Fig. 294 is the narrow green plant
growing up the tree trunk, and not the much more obvious tree itself? In my opinion, it would also
be helpful to have rather more detailed legends to the line drawings, so that they can be understood
without frequent reference to the text (consider, for example, Fig. 73). Teaching opportunities have
been lost in these ways. A good feature of the book is the provision of examples from all over the
world, often unusual or little known plants.

The second part of the book deals with “Constructional organization’’, especially the architecture
of tropical trees. Although this section is also interesting, I am not entirely convinced that the two
parts of the book successfully complement one another.

The author emphasizes throughout that, in order to understand the adult morphology, it may be
necessary to study development; but it is on developmental topics that the book is weakest. It is
unfortunate, also, that the dynamic aspects of phyllotaxis receive little emphasis, and that the
fractional method of description has been perpetuated.

Although there are a few curious placings — Cyclamen sits oddly with root tubers, for example,
and haustoria under root morphology — there seem to be few errors. There are also few
typographical errors, although the legend to Fig. 205 includes a substantial part of the legend to Fig.
239. The term membraneous is used instead of membranous, and principle instead of principal.

This is a beautifully produced book containing a wealth of information on plant morphology
which it is important should not be lost. It is not entirely clear, however, who its principal readers
will be. Few students of plant science to-day, however regrettably, will be able to spend the time
required to master the whole book (or have the funds to buy their own copy). On the other hand, if
the book is to be used as an illustrated dictionary, as the author suggests, it fulfils its purpose very
well. Anyone who has consulted Willis (A Dictionary of the flowering plants and ferns, 1966) or its
more modern counterparts will find this book extremely useful, as will those who need to describe
plants.

E. G. CUTTER

A colour guide to rare wild flowers. J. Fisher. Pp. 364. Constable, London. 1991. Price £12.95
hardback (ISBN 0—09-470780-4); £11.95 paperback (ISBN 0-09-469190-8).

British and Irish botanists are well served, too well served, with illustrated field guides. Having
recently reviewed our popular field botanical literature (British Wildlife 2: 214-218, 1991), and

44 BOOK REVIEWS

concluded that a substantial body of it is superfluous, my first reaction to another guide was
predictably jaundiced. Nevertheless, the publishers appear to have found a small niche in the
market, and have produced a neat and cheap volume. Some 150 species are featured, grouped by
seven regions, e.g. Home Counties, Isles of Scilly, that are rich in rarer species. Each text entry
consists of an informal description of the appearance and habitat of the plant opposite a full-page
photograph. A miniature map of the region included under each species is of no use at all, other than
a reminder that one is perhaps searching in the Home Counties for Melittis melissophyllum, which
might anyway be better sought in Cornwall — possibly the publishers were unaware of the Alas of the
British Flora? The photographs vary in quality and many do not come up to the standard of those in
recent field guides. The text has a freshness of style, however, which conveys that the author is an
enthusiast, is familiar with the literature and recent research, and knows his plants in the field.

Nevertheless, the contents tested even my robust attitude (pro-picking flowers, hunting,
shooting, etc.) to conservation. The title says it all: the book tells the naturalist, reputable or not,
where to find some of our rarest wild flowers. There will be those who feel that the book thus
provides a service, but this argument has the same suspect basis as that preached by those who
market illustrations of the unclothed human form. I fear that this guide will indeed fall ‘into the
wrong hands’; nor is seeking rarities as an end in itself a healthy intellectual exercise for the botanist.

Several text entries give detailed instructions as to how to reach localities for very rare plants,
including precise road directions and, more disturbingly, six-figure grid references. Fortunately the
concept of rarity employed here seems to be based on unfamiliarity (the book having a distinctly
south-eastern bias), and a good many of the species included are not particularly threatened.
Leucojum aestivum, for instance, flourishes in the suburbs of Reading, and most of the species
included in the section on Scilly are merely locally common weeds. One could include within this
category the native Scillonian rarity, Ornithopus pinnatus, a denizen of the celebrated Tresco
Abbey Gardens’ rockery. Nor will anybody who has seen Oxalis pes-caprae colouring Cretan olive
groves a sickly yellow in spring worry unduly about its possible fate at the remotest extremity of its
range. However, the locations of other species should never have been included, although
mercifully the author is circumspect about some of the choicer rarities.

The information divulged is of a sort best kept within notebooks and amongst friends (see
Obituary of John Codrington, p. 53). The book ends with a list of addresses and telephone numbers
of Wildlife Trusts, and a note on the dust cover suggests that they be approached for permission and
information, including the state of the season. I suspect that these bodies, with often limited
administrative resources, will be less than pleased to receive a deluge of enquiries from rarity
hunters armed with what ought to be confidential data! From my own experiences on excursions, I
know only too well that a minority of botanists do lack self-control and will dig up plants for their
gardens. Furthermore, the rising popularity and fashionability of horticulture, not least its current
fashion for ‘wild flowers’, means that rich pickings are to be had from selling rare plants — and, as
with antiques, some sources of supply are more dubious than others.

J. R. AKEROYD

The families and genera of flowering plants. Edited by K. Kubitzki. Vol. 1. The ferns and
gymnosperms. Edited by K. U. Kramer & P. S. Green. Pp. 430 with 216 figures. Springer Verlag,
Berlin. 1990. Price DM298 (ISBN 3-540-51794—4).

This well produced volume is the first in a proposed series covering the vascular plants. This
ambitious project is competently started, well produced, and written and edited by a selected group
of international experts in the different fields. The first volume of the series covers the ferns and fern
allies (edited by K. U. Kramer) and the gymnosperms (edited by P. S. Green).

The ferns are ably written by a recognized group of experts in the different families. As with all
such projects there is an unfortunate limitation in discussions caused presumably by restrictions in
length. Most of the articles provide an excellent review of the families and genera although, at least
at the Royal Botanic Gardens, Kew, not all the family and generic delimitations will be followed.
The detailed studies of Holttum on the Thelypteridaceae, which resulted in the recognition of many

BOOK REVIEWS 45

clearly defined genera are not followed, the species being placed in a restricted number of genera
rather than the more numerous genera of Holttum which indicate the relationships within the
family.

Classification of the ferns at the level of the family is still in dispute. At Kew we will be following
the families as listed by Brummitt. Thus the Pteridaceae will be treated in several families:
Parkeriaceae, Adiantaceae and Pteridaceae. In addition the genus Nephrolepis will continue to be
treated within the family Oleandraceae rather than in the monotypic family Nephrolepidaceae.
Generic limits outlined for the Polypodiaceae are not accepted and rather than the very broad
concepts of Aglaomorpha we will continue to accept its related genera, Thysanosoria, Merinthosor-
us, etc. which were studied by Roos. Indeed at both generic and family levels little stability in the
application of names can be expected. The limits used in this book are not necessarily widely
accepted.

Probably one of the most difficult aspects of the book is the decision to arrange the families
alphabetically. It is unfortunate that the families were not arranged according to their relationships,
particularly as family limits are still uncertain. This approach will cause major problems within the
dicotyledons.

The detailed treatment of the recognized genera with the references specific to each family will,
despite differences in generic delimitation, provide an indispensible reference book to the student
of the pteridophytes. It is difficult to understand why some chapters, such as Conservation, were
included in the section on ferns particularly with the limitations in space. The author of this section is
a recognized expert in conservation of pteridophytes and the chapter in no way reflects either his
knowledge or the available information.

The gymnosperms were the work of a single author, C. N. Page from Edinburgh. The treatment
provides an up-to-date review of the families and genera of the gymnosperms and the subdivision of
the Podocarpaceae is widely, if not universally, accepted. The high standards of the pteridophyte
section are equalled in the section on gymnosperms.

The series on vascular plants starting with this first volume will provide an indispensible reference
which must find its place in every reference library. It is hoped that the treatments at family level will
be organized according to the relationships between the families as this will greatly increase the use
of the volumes to the student of botany. The cost of these volumes will unfortunately restrict their
availability in the libraries of the developing world where they could be extensively used as a
reference for teaching University courses. This volume sets a high standard for the remainder of the
series and will undoubtedly be a classic of this decade. A low priced edition for the developing world
would be a great service to taxonomy.

R. J. JOHNS

Flora of eastern Saudi Arabia. J. P. Mandaville. Pp. x+482; 268 colour plates. Kegan Paul
International, London. 1990. Price £95 (ISBN 0—7103—0371-8).

This exemplary regional Flora fills a notable gap in the Arabian floristic literature. The author, an
American employed in the oil industry, has spent the past 25 years exploring the Eastern Province of
Saudi Arabia and has also made a major contribution to our knowledge of the flora of Oman.
Although the area covered is not rich in species, nor for that matter in families (73 are recognized in
the Flora), there were many thorny taxonomic problems to be confronted in preparing this first
- critical treatment of an area where the Saharo-Arabian element of the flora intermingles with
Mediterranean and subtropical elements.

The book starts with a short history of botanical investigations of the Province; several notable
travel writers are among those mentioned, including Harold St John Philby, Bertram Thomas and
Wilfred Thesiger. The geography and climate of the area are described, and a significant extension
of subtropical (Sudanian) vegetation into central Arabia is highlighted in the chapter on
phytogeography. The region’s palaeo-environments are also described, and the introduction ends
with a short section on vegetation.

The Flora proper is set out in the order of Stebbins’ sequence of 1974, which is close to that of

46 BOOK REVIEWS

Cronquist. Families, genera and species are keyed and described, and detailed distribution
summaries are augmented by citations of the author’s and other specimens. Vernacular names,
collected by the author in the field, are cited when available. His scholarly approach to etymology is
coupled with terse observations on economic uses; under Deverra triradiata, for instance, he notes
that ‘‘“Many Bedouin herdsmen note the camel’s particular fondness for this aromatic shrub.”

Selected species, such as three of the four Stipagrostis, are provided with dot distribution maps,
and there are 268 colour photographs, most of very high quality, which should broaden the sales
appeal of the book. A glossary of botanical terms, a gazetteer of geographic names cited in the text,
an eight-page bibliography, and two indexes of vernacular names (one in Arabic order, the other in
transliteration) precede the general index. The choice of a close-up of Aegilops kotschyi to illustrate
the dust jacket is a bold one, yet it epitomises a ‘desert plant’ and also represents the area’s largest
family, the Gramineae.

The author’s notes provide some original insights into the many taxonomic problems of the flora.
They are invariably helpful and perceptive, displaying a profound field knowledge of the Arabian
flora as well as setting out pointers to the need for further work. His extremely tactful phraseology
when referring to Migahid & Hammouda’s Flora of Saudi Arabia (2nd ed., 1978) as being ‘“‘designed
to be Kingdom-wide in scope . . . although the basis for its attribution of taxa to the Eastern
Province was somewhat unsure” will hardly offend anyone. But in practice Mandaville’s Flora will
be of far greater value to users outside the Eastern Province, as I have found when working on
collections from the United Arab Emirates, and it will be of great relevance throughout the
peninsula. The hopes expressed in the Preface that the book may “assist to some extent in the
development and conservation of natural resources in these lands which have been my home for 40
years” deserve to be fulfilled. Saudi Arabia now has a Flora, as well as a flora, of which to be proud.

J.R. EDMONDSON

The Cornish flora supplement 1981-1990. L. J. Margetts & K. L. Spurgin. Pp. vi+119, with
frontispiece line drawing and endpaper map. The Trendrine Press, Zennor, St Ives, Cornwall. 1991.
Price £14 (ISBN 0—-9512562-2-X).

In Margetts & David’s A review of the Cornish flora 1980, one of the authors forecast that “further
investigations would proceed vigorously” but he would not have foreseen that enough records
would accumulate within the next ten years to justify the publication of a Supplement. Knowledge of
the Cornish flora has been greatly increased by individual recorders and various surveys. These
include the Lizard project directed by Dr L. C. Frost of Bristol University, the monitoring scheme
and meetings of the Botanical Society of the British Isles, and mapping on a 1-km square basis in
S. E. Cornwall by the Caradon Field and Natural History Club. Not only have many areas been
studied in greater detail since the Review but more attention has been given to introduced species
ranging from aliens to garden outcasts.

The arrangement of families, genera and of the species (with a few exceptions) is the same as in
the Review, apart from the garden escapes and casuals which were listed at the end. In the
Supplement these are included in the main list. As in the Review, localities for the less common
species are listed under the number of the appropriate 10-km square but species that are common or
frequent and widespread were not treated in detail there and are not referred to in the Supplement.
In general the nomenclature, English names and distribution follow the Review closely, making it a
pleasure to use the two volumes side by side.

The Supplement is strongly bound, well laid out and clearly printed with the minimum of
typographical errors. Improvements over the Review include the greater distinction between the
generic name and its authority, and the placing of the Dandy (or other) number after rather than
before the specific name. The use of bold type-face for the word ‘introduced’ in place of a sign, and
for the vice-county ‘West’ and ‘East’ and for ‘Scilly’ make the text much easier to use. These three
words are succeeded rather than preceded by the star used to indicate a new vice-county or Scilly
record.

The authors have kept up with contemporary studies of various genera and mapping of critical

BOOK REVIEWS 47

species, for example in Rosa, Taraxacum and Rubus. The last-named genus includes the
appropriately named R. metallorum L. J. Margetts, which is frequent in the old metalliferous
mining areas in W. Cornwall. A description and Latin diagnosis of this new species is given in the
appendix.

The large amount of information in this Supplement includes the up-dating of some records,
reports of over 180 new vice-county records (including introductions) and of a number of exciting
discoveries. No one who has a copy of the Review can afford to be without this excellent
Supplement. It should also enhance sales of the Review, which is still available (see Watsonia 14:
293-294, 1983).

J. A. PATON

Flora of the Outer Hebrides. R. J. Pankhurst & J. M. Mullin. Pp. 171, with 6 maps, 3 transect
outlines and 4 half-tones. Natural History Museum, London. Price £19.95 (ISBN 0-565-01121-9).

This is a provocative Flora, being something less and something more than it pretends to be. In
appearance and content it is much of what we have come to expect in a modern county Flora. The
cover is immediately appealing. There are introductory chapters by specialists on geography,
geology, geomorphology and soils, climate and vegetation, floristic and vegetational history,
vegetation, botanical history and plant lore of the Outer Hebrides. In addition there is an index of
place names with six-figure grid references, the bibliography is cross referenced with particular
islands, and there are lists of S.S.S.I.s and N.N.R.s and of collectors.

The flora list is arranged in two column format which makes it compact and easy to scan though
the absence of an index of genera may be a continuing aggravation to users for whom plant
classification is not a daily routine. There are excellent keys to Euphrasia, Rubus, Hieracium and
Taraxacum which should tempt field botanists to plunge deeper into these troubled but well charted
waters and promote more intelligent collecting. The use of English and Gaelic names recognizes a
cultural dimension to local botany and the importance of vernacular names in education and
conservation. Ancient links with Ireland are seen in our mutual celebration of Cu Chulainn, Patrick
and Colm Cille in the Gaelic names. Compilers of the standard list of Irish plant names used in the
Census Catalogue of the flora of Ireland looked at times to Dwelly and Cameron for inspiration and
the compilers of the present list of Gaelic names have found inspiration in Irish examples, though
the comprehensive list in the second edition of the Census Catalogue was apparently not available in
time. While both compilations might have benefitted from closer co-operation, the Gaelic names
selected by Clark and MacDonald will go a long way towards the production of a standard list of
Gaelic names for plants. The Gaelic Books Council assisted with the expenses of the Flora. Molaim
an saothar agus an taca. [I praise the labour and the support — Ed. ]

There are few typographical or editorial errors though reference to a Salicornia perennis
community on p. 39 may cause some confusion.

The authors had to deal with an unusually high number of unsatisfactory records including
unsubstantiated literature records, field records not refound, student hoaxes and specimens of
doubtful provenance or identification. In this matter they have adopted the role of reporters rather
than commentators allowing themselves only expressions of mild scepticism or faint hope that
improbable records might be confirmed. Outright dismissal is delivered at times in the words of third
parties, as A. J. Wilmott’s, ‘‘I don’t believe it’’, on hearing a report of Cerastium arcticum. Users of
.the Flora will need to read the chapter on the history of recording and the introduction to
Potamogeton to get a flavour of the rivalries and indiscretions which hampered progress with the
Outer Hebrides flora for decades.

It would have been difficult to select from among the unconfirmed records all those which ought
to be disregarded and consigned to an appendix of errors; the baby might have been thrown out with
the bath water. As it is, the flora list is unsatisfactory. Further work on the confirmation or otherwise
of the doubtful records is needed. A conservative view ought to be taken of what is probable or even
possible in the islands.

The authors have acknowledged the considerable work done by their predecessors, notably

48 BOOK REVIEWS

Professor J. W. Heslop-Harrison, Miss M. S. Campbell and A. J. Wilmott. It is their wish that the
book should form a basis for a future, more detailed and comprehensive Flora. Meanwhile we are
indebted to the authors and their team of workers and writers and to the Natural History Museum
for an important and long overdue Flora of these fascinating islands.

D. SYNNOTT

Crucifers of Great Britain and Ireland. B.S.B.I. Handbook No. 6. T. C. G. Rich. Pp. 336. Botanical
Society of the British Isles, London. 1991. Price £10 (ISBN 0—901158—20-8).

This latest of the B.S.B.I.s monographic handbooks is a worthy addition to an indispensible and
acclaimed series. Crucifers, i.e. the family Cruciferae or Brassicaceae, covers “‘the 138 species most
likely to be found in the field” in Britain and Ireland, and sets a high standard for future titles and
revisions of earlier handbooks. Written with a blend of diligence and enthusiasm, which just
occasionally merges into whimsy, it is a more substantial volume than the others, but retains a neat,
compact feel in the hand. Although Tim Rich is now among our Society’s more familiar figures, the
B.S.B.I. was brave to commission one of its then younger members to write a handbook to this large
taxonomic group. Nevertheless, the risk paid off handsomely, and nobody doubts the wisdom of the
Publications Committee’s decision. Crucifers is a most useful contribution to the literature on this
family, and will be of value to all European as well as to British and Irish botanists.

The descriptions are thorough and based for the most part on living material, and under each
species there are substantial observations on taxonomy, variation, ecology and, where data are
available, biosystematics. A 10-page bibliography at the end reflects the author’s broad study of the
family. Introductory sections on taxonomy and identification are for the most part excellent. The
keys to genera and species are clear and should not be too hard to follow. The very first key directs
the reader to sections A—H of the main key, thus avoiding a lengthy crawl through the whole key.
The main key is embellished with small line drawings of morphological features, something that I
would not normally favour, but here they do not clutter and help to explain terminology to an
unfamiliar reader. It is a feature that will be popular with the Field Studies Council and others who
teach groups of students. Generic keys follow and there is an informal synoptic key to provide a
short cut and aide-memoire. I should have liked to see more in the introductory section on the
economic aspects of this important family that provides such a significant proportion of the
vegetable crops, salads and arable weeds of Europe. ;

The coverage is thorough, with the inclusion of all native and established adventive species,
together with several persistent casuals. This adds greatly to the value of the book, as it is likely that
certain species will become more widespread, not least as a result of the consequences to trade of a
post-1992 E.C. free market and increased contact with eastern Europe. Recent years have seen a
considerable expansion of the range of Hirschfeldia incana, for example, in both Britain and
Ireland, especially in London and Dublin where it is now rather common. Indeed, Tim Rich was the
first to report its occurrence in Dublin (/r. Nat. J. 22: 531-2, 1988), where too many of us had
dismissed it as Brassica nigra. Had this present work been available, we might have been
encouraged to note the seeds in the beak of the fruit! Conversely, Camelina sativa has decreased
markedly in Britain and Ireland during this century with the decline of the flax fields which were its
characteristic habitat. However, the revival of flax, or rather linseed, cultivation in Norfolk and
elsewhere, together with other new crops, may perhaps lead to an increase in the frequency of this
and other crucifers.

For two critical genera, the author has called on the services of other specialists. A highlight of the
book is the account of Rorippa, including hybrids, written jointly with B. Jonsell. More
controversial is the account of Cochlearia, by K. H. Dalby, which takes a distinctly narrow view of
specific limits, albeit one closer to the view of some continental workers. Two other departures from
convention, at least from Flora Europaea (including the as yet unpublished revision of Volume 1),
are the inclusion of Cardaria in Lepidium, with the treatment of the two European subspecies of L.
draba at specific rank, and the inclusion of Cardaminopsis petraea within Arabis. These small

BOOK REVIEWS 49

matters should not upset too many botanists; synonymy is nothing new in Cruciferae, a family of
economic importance that has thus been over-classified and now has too many genera!

The general appearance of the book, alas, falls down somewhat. The overall standard of the
copious line drawings suffers from their being the work of a number of artists, as they vary in quality
as well as style. The use of a single artist would have ensured a more consistent, professional-looking
product. This ought to be borne in mind for future handbooks, since they are a flagship publication
of our Society. Although the fact that the artists have provided their services free is to be applauded
and the great majority of the drawings are of a high standard, the end-product may support
accusations from some quarters that the B.S.B.I. is too often amateurish in its approach. The author
has obviously bravely coordinated the efforts of the various artists, making sure for example that
floral parts were consistently illustrated. The use of 10-km dot distribution maps has greatly
enhanced the value of the later handbooks, although in this case there are rather too many maps —
they are surely not necessary for rarer species — and sometimes too many symbols have been used for
clarity. The numbering of the maps, based on that used for species in the text, is confusing. In many
cases, I should have preferred a more detailed geographical description in the text itself.

Crucifers of Great Britain and Ireland is an essential book for the field botanist and should
encourage more of us to tackle them, especially the despised ‘yellow crucifers’. It would be good if
this handbook were to reach a wider public, both as a standard reference and as publicity for the
B.S.B.1., but the Society does maintain a rather cryptic publication policy.

J. R. AKEROYD

British Plant Communities. Vol. 1: Woodlands and scrub. Edited by J. S. Rodwell. Pp. x +395, with
25 line drawings and 25 floristic tables with accompanying distribution maps. Cambridge University
Press, Cambridge. 1991. Price £70 (ISBN 0—521—23558-8).

With this book the editor presents us the first volume of a planned 5-volume series. For the authors it
is the most difficult volume, since here the principles and methods must be laid out. Moreover, it
also encompasses the most difficult of the plant communities, the woodlands. Already at a first brief
look, it is apparent that it is scientifically a complete success, and that it is also an important work for
plant sociology. That it is being reviewed by a Continental European plant sociologist indicates that
it also has great value extending well beyond the British Isles. Indeed, it fills a painfully felt gap for
us. Descriptive vegetation science depends in the long run on regional comparison, and until now
such comparisons were difficult, if not impossible. To be sure, for Ireland and Scotland there were
several individual surveys available (viz. White & Doyle, McVean & Ratcliffe, Birse), but for
England and Wales, despite a wealth of synecologically and syndynamically valuable studies, a
comprehensive systematic review of vegetation types was lacking. In addition, the organization of
such a magnum opus is of interest, since only 15 years have passed since the beginning of the actual
field work — a short time for an enterprise based in an inductive way on circa 35,000 vegetation
relevés!

This undertaking received its initial impulse from the urgent need of nature conservation for a
systematic overview of the objects to be protected, valid on a large scale and based on uniform
standardized methods. Hence there was first of all a need for exactly described plant communities
(vegetation types), which at the same time form the biotopes for animals. Thus it is understandable
that the old discussions of methods far removed from actual practice did not break out again, rather
“one proceeded pragmatically — the success of the result shows that this was good. Although four
universities took part, it was possible quickly to agree upon a procedure and that despite different
evaluation methods, a self-consistent picture emerged. Dr J. S. Rodwell was primarily responsible
for this course of events. (Here I would have liked to have learned more about the methods and
techniques of the individual research groups involved.) In any case, one strived more for
ecologically valid statements than for difficult mathematical algorithms — much as on the Continent.
The British modus operandi (can one say: the modern British school?) has many features in
common with this Continental school, the Braun-Blanquet concept, which today has gained
worldwide acceptance. Indeed, certain passages could be incorporated in a Continental European

0 BOOK REVIEWS

textbook: common principles are for example taking into account all species, according to their
frequency of occurrence, the strict uniformity of relevé plots and last, but not least, the crucial
criterion for ordering called “frequency and abundance’, which evidently corresponds quite well to
the term ‘Treue’ of the Braun-Blanquet school. Even though no formal hierarchy was aimed for, as
it turns out three categories have been introduced and proved useful: community, sub-community
and variant. The parallel to associations is of course obvious. The authors then also draw parallels
between groups of communities and the Continental ‘Verbande’. One could now wish further to
have a general survey of woodlands in the form of a table of ‘Stetigkeit’ or degree of presence (as we
say instead of frequency).

All together, 25 communities have been described, which in turn are placed into six principal
groups, the largest one being the Mixed deciduous and Oak-Birch woodlands. The greatest part of
the book is dedicated to the presentation of the communities and their subunits. This is done in a
consistent way throughout, in the sequence: Synonyms — Constant species (i.e. those of high degree
of presence) — Rare species — Physiognomy — Sub-communities — Habitat — Zonation and succession
— Distribution (inside Britain) with dot-distribution map — Affinities (also with the Continent) —
Floristic table (with degree of presence and abundance). A list of literature references of around 400
items closes this work, an effort which lays out the basic aspects of the vegetation in a most
exemplary way, while offering a wealth of detailed information as well.

We congratulate our British colleagues, students, nature conservationists and floristically
interested active amateurs on this foundation for the plant sociology of the British Isles!

O. WILMANNS

The Orchids of Suffolk: an Atlas and History. M. Sandford. Pp. 123. Suffolk Naturalists Society,
Ipswich. 1991. Price £15 (ISBN 0-9508154-3-8).

Like its companion The Butterflies of Suffoik this is a well-produced, readable, well-illustrated and
informative book. Its 123 pages cover the 32 species that have been recorded from the county, only
twelve of which are now present in anything like reasonable numbers. The book also covers eight
hybrids. But the book is much more than just species accounts, for there are also interesting
chapters on ecology, variation, habitats, soil regions, conservation, the Suffolk Orchid Survey, and
a fascinating contributed chapter by Francis Simpson on “‘Suffolk orchids — half a century of change”’
which gives us one man’s historical perspective and brings out the joys of botanizing.

But it is the catalogue of species that most readers will use most and which occupied more than
half the book. The accounts have detailed information on the discovery and subsequent fortunes of
each species, along with ecological information, taxonomic problems, variations, as well as
anecdotal snippets from older publications. The conservation status is discussed but, here and in the
general chapter, the reader is faced more by a catalogue of woes rather than with ideas for the
future. Even allowing for some of the more intractable problems, more could have been done to lay
out the practical steps that could be taken to ensure that yet more local extinctions do not occur.
Related to this, it would have served the local and wider botanical community well if a list of needed
research had been included, perhaps with some idea of priority. How many more populations will
disappear because we failed to understand what simple management actions were required to save
them?

The reference list is complete, but the list of useful addresses is remarkable for its omission of the
Nature Conservancy Council (now English Nature) and its regional offices.

It is certain that this book will encourage increased local interest in and concern for this
fascinating group, and it is hoped that this will in future be reflected in improvements in their
status.

A. J. WHITTEN & L. FARRELL

BOOK REVIEWS 51

Recovery; a proposed programme for Britain’s protected species. A. J. Whitten. Nature Conser-
vancy Council, C.S.D. Report, No. 1098. Peterborough. 1990.

The bulk of this work is a compilation of proposals for active conservation to ensure the continued
survival in the wild in Britain of each of our endangered species, and it is a mine of information
about these species. For each of the 217 or so species treated, information is given on their
distribution, ecology, reasons for rarity and suggested management which could increase their
frequency to such an extent that the species would no longer be endangered. Each account ends with
costings for the parts of each proposal; site management, translocations, enforcement, research and
the monitoring which will be required over the next 15 years. The total budget required to carry out
all these proposals is around £800,000 over the 15 years, at 1991 prices.

In addition to the individual species accounts, the introductory 27 pages and a series of appendices
(72 pages) discuss the rationale behind the proposals and arrange the species of each category
(plants, bats, Lepidoptera, etc.) in order of degree of threat, recovery potential and the budget
required. English, Welsh and Scottish plants are considered separately and separate lists are also
given for each habitat.

It is interesting to see how little is known about most of our native rare plants (and the majority of
our common ones as well), so I strongly support Dr Whitten’s appeal for more autecological studies.
For most species we don’t even know whether they are usually self-compatible or self-incompatible
and only rarely are the conditions required for seedling establishment known. Many species appear
to require disturbance and therefore active management to provide the open habitats required for
seedling establishment (e.g. Stachys alpina, S. germanica, Teucrium botrys, Damasonium alisma,
etc.), such disturbances naturally having been provided by large mammals, perhaps especially wild
boar, a species eliminated from Britain by Man.

Gardeners may not be as surprised as Dr Whitten that some species which grow poorly in the wild
grow vigorously in cultivation when they are adequately watered and competition is removed (see
comment under Carex depauperata). Relieved from competition, many species of poor competitive
ability (e.g. Equisetum arvense, Poa annua) can thrive and grow all too well.

Through pointing out the gaps in our knowledge this book should stimulate many research
projects in addition to the conservation work proposed. How many of the proposals will be
implemented is rather uncertain in the present political and financial climate, but it is reassuring to
note that, though species-centred, the significance of habitat conservation is stressed, ‘honeypots’
with many rarities and great species diversity being mentioned as of particular importance.

The positive approach proposed in this work may not appeal to those who prefer ‘laisser-faire’
conservation, but Man has already done so much damage that many habitats and populations will
never recover without active intervention. We may think the situation bad in overcrowded Britain,
but it is much worse elsewhere, especially in relic communities and islands (e.g. New Zealand, see
New Scientist, 20th April 1991). While conserving Britain’s habitats, fauna and flora we must never
forget our responsibility as a rich country to the rest of the world and remember that if Britain were
to be wiped off the face of the Earth very few clearly distinct species would become extinct.and it
would make very little difference to the biodiversity of the Earth. This book is a model for similar
projects throughout the world and it clearly demonstrates the value in Britain of the specialist
societies and County naturalists’ trusts.

With many people having been involved in the compilation, errors are bound to have crept in. On
page 280 the Natterjack Toad is referred to as a “‘lizard’’. Arabis alpina is described as annual,
though all Floras and experience show it to be perennial, and there is no mention of the fact that
Cotoneaster integerrimus is apomictic, a major consideration when deciding how it should be
- propagated as it grows in the presence of other Cotoneaster species and is difficult to root from
cuttings. These, however, are very minor criticisms. It is perhaps a pity that bryophytes are not
covered as some primarily western oceanic and montane species and habitats are probably of
greater world significance than any of the British populations of vascular plant or animal species
dealt with here. A follow-up work on bryophytes and lichens would be welcome, though recovery
would depend almost wholly on habitat management, as we have not yet developed the knowledge
and skills to-cultivate them.

H. A. McALLISTER

PEW AES

me
Lind

Watsonia, 19, 53-66 (1992) 53

Obituaries

JOHN ALFRED CODRINGTON
(1898—1991)

The varied aspects of that endearing character, Lt-Col. John Codrington, have been well set out in
obituaries in The Times and The Daily Telegraph of 23 April, The Independent of 3 May, and The
Rutland Times, supplemented by further notices in The Times. These cover his extraordinary
career, but hardly mention his life-long absorption in plants.

He painted four remarkable sets of flowers, dated 1905, 1905, Spring 1906 and Summer 1906, 19
in all, when he was 6 and 7. In July 1908, aged 9, he was given a nicely bound copy of the Rev. C. A.
Johns’ Flowers of the Field, and he was growing alpines when still young. The interest never left him,
and was kept up even during his 20 years with his father’s regiment, the Coldstream Guards. In
1920, when quartered at Wimbledon, he noted Potentilla norvegica. He sent a long letter from
Salzburg to the Wild Flower Magazine dated September 1922, recording several plants; and another
from Constantinople in June 1923.

He wrote a beautifully clear set of “botanical maps and directions” for over 300 scarce and rare
plants in England and Scotland, labelled ‘“‘confidential’’, with an index. An O.S. map is noted for
each and the grid reference. Most are dated 1953, but they extend from 1951 until 1969. Some dozen
are marked “most secret”’, and as many “‘secret’’, but not all will still be valid, e.g. for Agrostemma
githago. This set has been gifted to the Botany Library of the Natural History Museum, London.

He kept up his painting all his life, almost all of landscape or gardens, including garden designs —
his first essay in design was a knot garden at Rockingham Castle in 1916, which still survives. The
only later painting of a flower I have seen is of Cypripedium calceolus in 1957, which is with his
directions for finding that species, and is frankly a poor depiction. In all, something between 2,000
and 4,000 of his paintings survive.

The Royal Horticultural Society awarded him its Veitch Memorial Medal in gold in 1989, “‘living
proof that the finest gardeners never age”’ as the President put it. There is a good account of him in
The Garden for November 1987.

I first met him soon after the last war, and we had kept in regular touch ever since, including
on the tours of the International Dendrology Society, of which he was a faithful member. At
both the houses he eventually lived in, in Pimlico and Rutland, he grew remarkable plants. The
Ranelagh Cottage house reverts to the Grosvenor Estate, which will try to get a suitable tenant
for the garden. What happens to Stone Cottage, at Hambleton, Rutland, is currently uncertain.
Its garden he described in The Englishman’s Garden of 1982, and in The Garden for November
1988.

He worked with Sir Alexander Korda for some years after the war (with Merle Oberon too) at
London Films. On one occasion he noticed on a film set that the flowers were wrong for the time of
year the action was taking place. Sir Alexander bet him that no-one would complain, and no-one
did. Among the plants that interested him most and he distributed widely were Bupleurum falcatum
from Essex stock, which seeded freely in his chalky garden, Smyrnium perfoliatum, Bowles’ Golden
Grass Milium effusum ‘Aureum’, and white-flowered Herb Robert (Geranium robertianum). He
grew many native and naturalized plants in his ‘“‘mad wild jungle” (as he put it) at Hambleton,
Sisymbrium strictissimum doing particularly well; and there were others he had collected from
exotic places. He took seed of a cypress from the Holy Land in 1922, two of the progeny being still in
the churchyard at Oakham and others in that at Preston, nearby, all family territory.

His name will be found as having made contributions to publications such as The Hand List of the
Plants of the London Area of 1951, The Atlas of the British Flora of 1962 and the Flora of Essex of
1974. He was the unquenchable enthusiast, pestering me to go and see how the London Rocket
(Sisymbrium irio) was on Tower Hill (which he had seen during 1951-6 at least) until a year or so
before he died.

Apart from the accounts already mentioned, I know of nothing that he put into print about plants,
except occasional letters to Country Life. This is sad, because he had travelled the world endlessly

54 OBITUARIES

and saw much, and had written a manuscript autobiography. I think that he did not consider himself
a botanist, but was just a devoted, clubbable plantsman, with a wealth of stories. But what a
plantsman!

D. McC inTock

NORAH DAWSON
(1913—1991)

Norah Dawson, B.S.B.I. Recorder for County Armagh, died on 4 March 1991. A daughter of
Armagh, the ‘City of Saints and Scholars’, Norah was one of the most modest and yet competent of
amateur botanists. A history graduate of The Queen’s University of Belfast, she trained as a
librarian, and spent her working life in the employment of Armagh County Council, where she was
for many years the County Librarian.

Norah’s interest in botany developed through the Armagh Field Naturalists’ Society, of which she
became one of the founder members in 1952. At first, the Society was a small band of enthusiastic
novices. Norah, because of difficulties with her sight, preferred to concentrate on plants, whereas
birds absorbed most of the attention of other members. Perhaps rather unusually for her time, she
had studied German rather than Latin at school, but nevertheless soon mastered the use of Latin
names for plants.

With encouragement from the late Pat Kertland, Norah became interested in recording plant
distribution during the years leading up to the publication of the B.S.B.I.’s Atlas. Later, during the
1960s, the Armagh Field Naturalists along with the Belfast Naturalists’ Field Club began a project to
record the Co. Armagh flora in 5-km squares, and Norah became the stalwart of this project.

On her retirement, she took on the honorary secretary’s post in the Field Naturalists’ Society, a
role she carried out with singular efficiency and good humour for several years. As the Society’s
chairman and a near neighbour, I came to know Norah and her constant companion Burren well at
this time. Burren, a red setter, was ironically not named after the botanists’ Mecca in western
Ireland, but after a little-known townland of that name in Co. Down. In fact, Burren became known
under another name to my family rather earlier than Norah, on account of the memorable sight of
him emerging from another neighbour’s back door and careering across the unfenced back gardens
of our recently built estate with a string of sausages hanging from his jaws. Burren was temporarily
known as ‘The Sausage Stealer’; quite unfairly as it turned out, as we later discovered that the
neighbour in question was a butcher with a surplus of out-of-date sausages.

The early years of her retirement were the time when Norah was at her most botanically active,
most days in summer being spent recording along the by-ways of the County, and also further afield
in Ireland, both on her own or with the Irish Biogeographical Society. As befitted a professional
librarian, her plant records were kept in a magnificent state of order, and she also kept meticulous
notes on butterflies in Co. Armagh. Her role as a B.S.B.I. Recorder was largely one of collating her
own records, the county at that time being sparsely populated by resident botanists and seldom
visited by outside ones.

A modest and unassuming person, Norah could always be relied on to do anything she undertook
in a thorough and reliable way. She always felt that a little more work was needed before the records ©
for her county would be in a fit state for publication. It was a great shame, therefore, that health
problems led to premature decline in her recording activities and, as the major contributor to the
stock of botanical records in Co. Armagh, that she did not live to see them through to publication in
a County Flora.

2 . S. FAULKNER

OBITUARIES 55

JOHN GEORGE DONY
(1899—1991)

John Dony M.B.E., B.Sc., Ph.D., Hon. F.L.S., who died in Luton on 24 March 1991, was one of
the best-known British field botanists of this century. He was essentially interested in the
distribution, ecology and features which were susceptible to statistical analysis rather than the
systematics of plants. His Flora of Bedfordshire was widely recognized as a model of how such things
should be done and truly, as the late Sir E. J. Salisbury said, “takes a worthy place amongst its
predecessors and contemporaries’, and so does its author amongst his. His efforts for conservation
in the county in which he was born and died were sustained and successful, resulting in a well-
merited M.B.E. in 1983.

John was born in Luton on 8 August 1899 in Court Road by the parish church in a road of typical
mid-Victorian terraced houses, now totally destroyed. A great fondness for the church, indeed a
magnificent building, remained with John throughout his life, although he had no time for religious
doctrine. His father, an engineer, was a well-known Sunday School teacher and a very active
member of the Chase Street Mission, Luton from 1901 until its closure in the early 1940s. His great-
grandfather was a Cornish tin miner who changed the name from Doney. His mother’s family were
local and mainly connected with the hat industry. His grandfather was a founder member of the
Luton Industrial Co-operative Society in 1883, and John himself was an ardent Fabian. His
education at Surrey Street School, like that of so many of his contemporaries, was restricted to
elementary school (but from what I know of its products it must have been immeasurably superior to
that of the present day). In 1913 he became an apprentice at Hayward Tyler, an engineering firm
renowned for their hydraulic pumps, where he worked on the bench. He stayed in the firm until
1920, his service there broken by World War I, when he joined the Royal Navy Volunteer Reserve
from June to December 1918, although he did not see active service. From 1920 to 1922 John
worked as a draughtsman in London and Cardiff but then after matriculating at London University
in 1922 became an uncertificated school teacher. His working in Cardiff probably explains his
support of Glamorgan County Cricket Club. A suggestion that his first teaching post was in Cardiff
appears to be inaccurate as there is no other mention of this, and his first school post was at Norton
Road School, Leagrave. He obtained an Acting Teachers’ Certificate in 1924, the last time a teacher
could become qualified without going to college. He moved to Queen’s Square School from 1925

56 OBITUARIES

until 1931, then to a school in Kentish Town for nine years. Although he had attended W.E.A.
evening classes in engineering, mathematics and Spanish years before, his attempts to better his lot
now started in earnest. He attended the University Extension Class, 1931-32, in Economics at
Toynbee Hall with W. Milne-Bailey as tutor, for whom he wrote a long essay on the hat industry
which was shown to J. J. Mallon, the Warden of Toynbee Hall, and Barbara (later Baroness)
Wootton, both of whom encouraged him to continue. He enrolled as a student of the London
School of Economics, working in the evenings until he gained his B.Sc. in 1936. He had married
another teacher in the 1920s but this marriage was dissolved in 1931.

The beginnings of the straw hat industry in Luton go back to visits by James I, whose mother
Mary, Queen of Scots had brought over to Scotland some Lorraine straw-plaiters — or so the story
used to go. Actually the industry may have had more local origins. However it started, it eventually
dominated the town, which became a main centre in Europe for the trade. The extent of its
importance is now inconceivable to a generation most of whom do not even own a hat. My own
paternal grandfather, a Belgian hatter, came to Luton in the 1890s when it was the Mecca for hat-
workers of all sorts. John had decided to work on and obtain a doctorate and the hat industry was an
obvious choice of subject for a Lutonian and he gained a Ph.D. in 1941 for his thesis on its history
and economics. To tell the truth he used modern pressurised interrogation techniques on the
surviving hat firms of the day by bombarding them with innumerable queries and wanting to see
their records. From his thesis he produced a book, A history of the straw hat industry (1942), which
became accepted.as the standard, if not the only, work on the subject. The industry rapidly declined
until it became a fragment of the town’s economy but John’s desire to investigate this was never
accomplished. His teaching career blossomed and after a short spell at the North Western
Polytechnic, he became in 1941 History and Economics Master at Luton Modern (later Grammar)
School and remained there until his retirement in 1964. There he was much loved and universally
known by the nickname ‘Doc’. ,

It is always rather difficult to pin down accurately the start of an interest, but John as a schoolboy
belonged to The Band of Hope, a society for promoting temperance principles among the young
(although John was not averse in later life to a modest drink in convivial company). This worthy
body gave a series of books as prizes for an essay on ““The evils of strong drink” and John, a winner,
chose The Works of Shakespeare and J. Saunders’ Field Flowers of Bedfordshire, the latter an
austere work scarcely likely to inspire a youngster. When he was 15 he made a collection of insects
and plants and received a commendation for the flowers. It is likely that Frederick Mander, (later
Sir Frederick, General Secretary and President of the N.U.T. and Chairman of Bedfordshire
County Council), keen on orchids and a teacher at Surrey Street School, encouraged him most of all
to let his innate interest in natural history develop. Mander’s name appears several times in the
Orchidaceae in John’s Flora. John’s early interest in botany relaxed during the time he was
obtaining qualifications but was rekindled in 1935 and developed considerably during the War
years. Once he had resolved to write a new Flora of his county, a group of like-minded friends
(particularly Horace Souster, perhaps his most constant companion). gathered together and an
immense amount of field work was undertaken. How far this had advanced was evident from a very
successful exhibition entitled ““Wild Flowers in Bedfordshire” which was mounted at the Luton
Museum some years before the Flora was published. The covering booklet for this was of course
written by John and for many Lutonians it was the first intimation of the project. This sustained
labour by John and many helpers culminated in his Flora of Bedfordshire in 1953, without doubt his
finest work. This was a subscriber publication and was produced for the very modest sum of “42s
nett”. I was already an old Africa hand by then and received my copy in Nairobi together with a
letter from John dated Christmas Day, 1953 (he wrote in a small but very legible hand which
achieved maximum information per square inch of paper). Almost unbelievably the 1000 copies cost
only £1271 to produce and an expected deficit of £600 was soon turned into a small profit. John was
extremely sensitive about the reception of this work and he awaited comments eagerly. I think that
they were without exception extremely favourable, as indeed they should have been — John had a
natural bent for accuracy and his publications were accordingly much more polished than those of
many a professional scientist. The 20 pages of historical introduction to Bedfordshire botany make
fascinating reading, combining as they do John’s twin interests of botany and local history. The
geographical index is a feature I still constantly use. Today this 42 shilling book usually fetches about
£18—25, which seems rather low bearing in mind inflation. It was reissued in 1978 in a much less

OBITUARIES 57

pleasing format. John then moved on to writing the Flora of Hertfordshire, which was published in
1967 by the Hitchin Urban District Council, again with an introduction by Sir E. J. Salisbury.
Although only 14 years later than the Bedfordshire Flora, the changes in book production gave it an
altogether different look. There was also the novelty of 47 pages of tetrad dot maps. These two
Floras show to a remarkable degree John’s capability of marshalling an enormous amount of varied
information and presenting it in an orderly form; this was perhaps his dominant characteristic; it is
of course a prerequisite of a good historian.

John developed a deep interest in conservation but was not unreasonable. He recognized that
progress, whatever we might individually think about it, was inevitable and did not make foolish
demands. His well-reasoned pleas produced more success as a result. Perhaps all this was started as
a result of a disgraceful act he witnessed as a youngster — the demise of Grass of Parnassus (Parnassia
palustris) in Bedfordshire. The last population was uprooted in his presence by an amateur botanist
who proudly gloated: ‘“That is the last time anyone will find that here!”’. John mentions this in his
Flora of Bedfordshire but did not hint that he witnessed it. Perhaps it is as well that I have forgotten
the name of the perpetrator. The designation of Knocking Hoe, Pegsdon as a National Nature
Reserve in 1958 was a major success for John.

Like many on the political Left, he seemed to actively enjoy being on committees and helping to
found new societies; he did valuable work for a surprising number of them. Early on, his political
affiliations developed (he had long warned of the dangers of fascism) and he became a local
secretary of the Left Wing Book Club and a founder member of the Luton Branch of the Fabian
Society. He became its chairman when it re-formed in 1967 after ten years of lack of public interest.
He was Honorary Keeper of Botany at Luton Museum from 1935 until 1988 during which time he
built up an admirable local herbarium and library; from 1955 he undertook the same duties at
Hitchin Museum. Already a Member of the Wild Flower Society, he joined the Botanical Society of
the British Isles (then the Botanical Exchange Club) in 1937 and after 1947 held office continually
- aS, successively, a Council Member, its Honorary Field Secretary (later renamed Honorary
Meetings Secretary) (1949-56), Honorary General Secretary (1956-64), a Vice-President (1965-66)
and finally President in 1967-69. After this he was made an Honorary Member. He helped found the
Bedfordshire and Huntingdonshire Naturalists’ Trust in 1961 and was also a founder member in
1962 of the Hertfordshire and Middlesex Trust. He was almost a founder member of the
Bedfordshire Natural History Society but missed the first meeting ( I fear I used to gloat over this);
he was, however, intimately associated with it for over 40 years and served as President in the early
1960s. He also served as President of the Hertfordshire Natural History Society and of the South
Bedfordshire Association of National Trust Members. In 1954 he was elected a Fellow (Honoris
Causa) of the Linnean Society of London. Other posts which he held were Chairman of the South
Bedfordshire Preservation Society (which he helped form in 1948) and the presidency of the Luton
Workers’ Educational Association which he took over from Lord Hill (‘The Radio Doctor’) in 1977
and retained for ten years. Towards the end of the 1980s John relinquished his various duties and
one of his last acts was to grace the opening of the John Dony Field Centre by the Mayor of Luton in
October 1990.

He was extremely interested in alien plants, particularly those resulting from the use of wool
shoddy; a remarkable number grew in Bedfordshire and in some places such as Flitwick railway
sidings one could be forgiven for thinking one was in Australia rather than Britain.

Anxious that every record should be as accurate as possible, he gradually got to know all the
botanists of his day, particularly the specialists on tricky groups. Many of them accompanied him on
field trips. Whilst Edgar Milne-Redhead was stationed at Dunstable during World War II, a great
friendship sprang up between them and they travelled by bicycle and train over much of the county
collecting records. A particular friend was V. H. Chambers who died in 1984. He had begun a
herbarium when only twelve years old and continued to list his records until 1931. Later he
specialized in the Hymenoptera and became a recognized authority on certain groups. He
collaborated with John and was in fact the stimulus which persuaded John to undertake the arduous
task of writing a proper Flora. Vic had a car early on and they undertook numerous joint
expeditions; Vic shared with me the almost total inability to ride a bicycle. Many young botanists
owe much to John’s encouragement. Peter Taylor and I were both protegés during the war and we
both decided eventually to become professional botanists. Peter was very neat and tidy (like John)
and did very much more to help John with recording and indeed wrote the Section on the Hepaticae

58 OBITUARIES

in the Flora of Bedfordshire. | already had many zoological interests which interfered with botanical
recording. My father viewed my interest in botany with dismay, particularly as I had a degree in
physics, and was certain I would never get a job as a biologist (they were rare in those days) and it
was John who showed it was a possibility, and in fact made it a respectable option. Without his help
and introductions to other people, probably neither Peter nor I would have ended up at Kew. One
of my main memories of him during the war years was the speed with which he walked despite his
short stature — like a galleon in full sail bent into the wind — and this continued well into later life. In
those early days transport was scarce and we had no car — a large part of most expeditions was done
on foot which of course helped us to find more.

John got on well with others, whether peasants or aristocrats. During the gathering of records for
the Flora he needed to examine the flora of many private and government properties. This was
usually arranged in advance, nearly always without difficulty, but occasionally had to be done off the
cuff, so to speak. He had a formula for this: when the door opened he would say ““We are botanists”
in much the same way as one might announce that one was the Governor of the Bank of England!
Nearly always this resulted in permission being given — only occasionally was the reply the
equivalent of ‘get lost’! John had a quite authoritarian air about him which somewhat overawed the
opposition.

A group of us often ended up in a pub (as often as not the Speed the Plough just north of Barton
and actually mentioned by name on sheet 95 of the old one inch map) on our way back from a
meeting in Bedford for a modest drink. He ate very frugally, often only an apple for lunch during a
field trip; which fruit he was proud of being able to split into two pieces by a deft twist of the hand.

Although as a schoolboy he disdained history, a W.E.A. evening course on economic history he
attended in 1917-18 kindled a strong interest —- so much so that history ultimately became his bread
and butter. He became particularly fascinated by local history, about which he gained an
encyclopaedic knowledge. It is an interesting coincidence that the road of his birth long ago formed
part of the estates of Sir John Rotherham, one of the oldest parts of Luton. He was part author of
The Story of Luton (1964) (reprinted 1966 and 1975) and wrote many other historical papers
besides. John’s total output of articles was quite considerable, many being in the Bedfordshire
Magazine and The Bedfordshire Naturalist, also Watsonia and its predecessors. He was an excellent
writer of obituaries.

In 1971 he joined the élite band of amateur naturalists to have received the Bloomer Award of the
Linnean Society (H. H. Bloomer, 1866-1960, was himself an amateur who mainly studied bivaive
molluscs). John married for a second time in 1962, Christina Mayne Goodman, a keen Birmingham
botanist and former England hockey international, who proved the perfect companion. Most of
John’s work after this date was in the nature of a joint effort. He is survived by her and a son by his
previous marriage. Undoubtedly the secret of John’s success was hard work and determination
coupled with enthusiasm and orderly methods, all in fact attributes of the Victorians, one of whom
he could just claim to be. He could certainly have been a successful politician and probably a
business man but he chose to become a schoolmaster don who will long be remembered for his
attainments.

PUBLICATIONS OF J. G. DONY*

MAJOR WORKS

1942 A history of the straw hat industry. Luton.

1946 The hat industry, in H. A. SILVERMAN, ed. Studies in industrial organisation, pp. 155-198.
London.

1953 Flora of Bedfordshire. Luton (reprinted 1978. Wakefield).

1964 (With J. Dyer and F. Stygall) The story of Luton. Luton. (2nd ed., 1966). (With J. Dyer, 3rd
ed., 1974).

1967 Flora of Hertfordshire. Hitchin.

1970 A history of education in Luton. Luton.

*Compiled mainly from a list produced by himself. It excludes detailed elaboration of recorder’s reports.
Some very short notes, exhibition reports, etc. are also omitted.

1974

e976
1984
1991

OBITUARIES 59

(With F. H. Perring and C. M. Robb) The English names of wild flowers. London. (With F.
H. Perring and S. L. Jury, 2nd ed., 1986).

Bedfordshire Plant Atlas. Luton.

The story of High Town. Bedfordshire County Library. (2nd ed., 1985).

(With C. M. Dony) The wild flowers of Luton. Luton.

~ MAJOR WORK EDITED

1975

A view from the alley (by A. S. Darby). Luton.

| ARTICLES

1946

1947
1948
1948
1949
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/
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1950

Additions and emendations to the Comital Flora for v.c. 30 (Bedfordshire). Rep. botl Soc.
Exch. Club Br. Isl. 1943-4: 803-14.

What Bedfordshire is. J. Beds. Nat. Hist. Soc. 1: 8-12.

Bedfordshire naturalists 1.: William Crouch (1818-1846). Ibid.: 50-52.

The case for nature reserves. Bedford. Mag. 1: ie

Wild orchids in Bedfordshire. [bid.: 145-8.

(With K. Piercey) Nature reserves. Bedford. Nat. 2: 19-21.

Bedfordshire naturalists 2.: James Saunders (1839-1925). Ibid.: 58-61.

A Bobart herbarium at Bedford. Bedford. Nat. 3: 15-16.

Bedfordshire Naturalists 3.: Charles Abbot (1761-1817). Ibid.: 38-42.

Hats and headgear. In Chambers’ Encyclopaedia.

A contribution to the flora of Huntingdonshire. Watsonia 1: 301-7.

Bedfordshire Naturalists 4.: William Hillhouse (1850-1910). Bedford. Nat. 4: 40-2.
Problems of distribution raised in the compilation of a county flora, in J. E. Lousley, ed. The
study of the distribution of British plants, pp. 69-74. London.

The distribution of Bunium bulbocastanum. Ibid.: 74-76.

(With J. E. Lousley) The travels of plants. Bedford. Mag. 3: 185-9.

Wool aliens in Bedfordshire, in J. E. Lousley, ed. The changing flora of Britain, pp. 160-3.
London.

Cyperus rotundus. Proc. botl Soc. Br. Isl. 1: 159.

The position with regard to the conservation of nature in Bedfordshire. Bedford. Nat. 8: 12-
13:

Hordeum leporinum Link, H. glaucum Steud. and H. pusillum var. pubens Hitchcock. Proc.
botl Soc. Br. Isl. 1: 323-24,

Notes on the Bedfordshire railway flora. Bedford. Nat. 9: 12-16.

The drawings of Caroline Gaye. Bedford. Nat. 10: 14-15.

The place of the local flora in the study of the British flora, in J. E. Lousley, ed. Progress in
the study of the British flora, pp. 30-9. London.

Some Hertfordshire plant records. London Nat. 38: 74-6.

Nature conservation in Bedfordshire. Bedford. Nat. 14: 19-24.

Flowers of the roadsides. Bedford. Mag. 8: 34-7.

The expectation of plant records from prescribed areas. Watsonia 5: 377-85.

Counting the cost, in P. J. Wanstall, ed. Local Floras, pp. 87-94. London.

Botanists in Hertfordshire. Hertfordshire Past and Present 3: 39-44.

The Botanical Society of the British Isles. Wild Flower Mag. 339: 5-6.

The Hertfordshire Natural History Society. Wild Flower Mag. 345: 6-7.

A Bedfordshire botanist and schoolmaster. Bedford. Mag. 11: 69-72.

(With J. Godber) Bedfordshire. Encyclopaedia Britannica.

Nature reserves in Bedfordshire. Bedford. Mag. 11: 153-6.

Presidential Address. The B.S.B.I. in a changing Britain. Proc. botl Soc. Br. Isl. 7: 311-23.
Additional notes on the flora of Bedfordshire. [bid.: 523-35.

Charles Abbot’s ‘Flora Selecta’. Bedford Nat. 23: 27-9.

Notes on the flora of Hertfordshire. Trans. Herts. Nat. Hist. Soc. 27: 23-4.

Straw hats — a bibliography. Costume 1: 10-11.

Plant notes for 1969. Trans. Herts. Nat. Hist. Soc. 27: 83.

60 OBITUARIES

1970 The journals of the Botanical Society of the British Isles. Watsonia 8: 1-2.

1970-1 The wild flowers of the river. . . of the riverside . . . of the Ouse meadows and gravel pits.
The trees of the river. The Lock Gate 3: 58-60; 69-70; 98-100; 107-8.

1971 The need for a recommended list of English names for British plants. B.S.B.I. Recorders’
Conf. Rep., pp. 87-94.

— Plant Notes for 1970. Trans. Herts. Nat. Hist. Soc. 27: 119-20.

1972 Opening up the Bedfordshire countryside. Bedford. Mag. 13: 184-90.

— A new system of recording for Bedfordshire. Bedford. Nat. 26: 12-15.

1973 The straw-plaiting industry in Bedfordshire and Hertfordshire. J. Watford & Distr. Ind. Hist.
Soc. 3: 34-42.

1974 Changes in the flora of Hertfordshire. (Presidential Address). Trans. Herts. Nat. Hist. Soc.
27: 255-64.

— Some problems of a railway flora. Herts. and Middlesex Trust for Nature Conservation
Symposium 2: 2-5.

1975 The Bedfordshire Natural History Society. Wild Flower Mag. (Spring Issue): 11-12.

— Putting a quart into a pint pot. B.S.B.I. Recorders’ Conf. Rep. (Attingham Park).

1976 How Luton became a borough. Bedford. Mag. 15: 135-40.

— (With K. Williamson) The natural history of the Ashridge Estate. National Trust.

— Plant Notes, 1971-1974. Trans. Herts. Nat. Hist. Soc. 27: 323-4.

1977 (With C. M. Dony) Some additional wool aliens in Bedfordshire. Watsonia 11: 428.

— Species-area relationships in an area of intermediate size. J. Ecol. 65: 475-84.

— §Species-area relationship in Bedfordshire. Bedford. Nat. 30: 61-2.

— Change in the flora of Bedfordshire, England, from 1798 to 1976. Biol. Conserv. 11: 307-320.

— Plant Notes, 1975-1976. Trans. Herts. Nat. Hist. Soc. 28: 26-7.

1978 The 1919 Peace Riots in Luton. Publications of Beds. Hist. Rec. Soc. 57: 205-33.

— Flowering Plants, in D. Shirley, ed. Hertfordshire, a guide to the countryside, pp. 34-7.

1980 Some wild flowers of the Trust’s reserves in Bedfordshire. Ardea: 37-42.

— Wool aliens. Watsonia 13: 73-4.

— Melampyrum arvense L. - a native or alien species? [bid.: 166.

1981 The natural vegetation of Dunstable and Whipsnade Downs. Bedford. Nat. 35: 43-54.

1982 (With C. M. Dony) Childing Pink in Bedfordshire. Watsonia 14: 228.

1985 (With I. Denholm) Some quantitative methods of assessing the conservation value of
ecologically similar sites. J. appl. Ecol. 22: 229-38.

— Introduced plants, in Wildlife in towns and cities (Country Life Guides): 32-38.

1986 Introduction to Mate’s Illustrated Guide to Luton. Luton.

—— (With C. M. Dony) Further notes on the flora of Bedfordshire. Watsonia 16: 163-172.

1987 Naturalists in the past; Changes in the Flora, in B. S. Nau, C. R. Boon & J. P. Knowles, eds.
Bedfordshire Wildlife, 13-25 & 79-83.

— Nature reserves in Bedfordshire. Bedford. Mag. 20: 323-7.

PAMPHLETS AND PRINTED REPORTS

1943 Report on the Luton hat industry. (To Nuffield College social reconstruction survey.)

*1948 Wild flowers in Bedfordshire. (An exhibition at Luton Museum and Art Gallery.) [undated].

1947-86 Reports of Recorder for Botany (J. Beds. Nat. Hist. Soc., later Bedford. Nat.)

1948 Report of field meeting: Bedford and district. Rep botl Exch. Club Br. Isl. 1946-7: 220-3.

*1948 Bedfordshire Wild flowers. Luton Museum.

1950 Report of field meeting: Huntingdonshire. B.S.B.J. Year Book 1950: 51-3.

1970 (With S. Cowdy and P. D. Rixon) The natural history of Milton Keynes country zone. (To
Countryside Commission.)

1971 Species-area relationships. (Mimeo. to Natural Environment Research Council.)

— A report on sites of natural history interest in Bedfordshire. (To Bedfordshire County
Council.)

2 The classification and assessment of mires in Bedfordshire. British Ecological Society.

*There appear to be two separate items involved. I have a copy of a white undated 16-page pamphlet with a
photograph of Alisma on the front cover; C. Boon has a green pamphlet dated 1948 with a similar photograph
but with 20 pages.

OBITUARIES 61

OBITUARIES
1963 Francis Morland Day, M.A. (1890-1962). Proc. botl Soc. Br. Isl. 5: 194.
1965 Wendell Holmes Camp (1905-1963). Ibid. 6: 99.
— Charles E. Freeman (1906-1965). Bedford. Mag. 10: 4-6.
1968 Gordon Mark Leo Haskell (1920-1967). Proc. botl Soc. Br. Isl. 7: 496.
1976 Ray Palmer (1896-1975). Bedford. Nat. 29: 66-7.

_— Thomas Wyatt Bagshawe — an appreciation. Bedford. Mag. 15: 185-7.
1977. J. Edward Lousley (1907-1976). Watsonia 11: 282-6.
1980 Sir Edward J. Salisbury C.B.E., D.Sc., F.R.S., F.L.S. (1886-1978). Trans. Herts. Nat. Hist.

Soc. 28:2.

_— Harold Owen White — an appreciation. Bedford. Mag. 17: 227-8.

_ 1985 Victor Horace Chambers (1911-1984). Bedford. Nat. 39: 3-5.
1990 John Campbell Gardiner (1905-1989). Watsonia 18: 239-240.

1964 Flora of Nottinghamshire (R.C.L. & B. M. Howitt); A contribution to the flora of Merioneth
(P. M. Benoit & M. Richards). Proc. botl Soc. Br. Isl. 5: 277-8.
1965 The concise British flora in colour (W. Keble Martin). Proc. botl Soc. Br. Isl. 6: 198-9.
1966 A guide to herbarium practice (J. W. Franks). Ibid.: 299.
1968 Britain’s Green Mantle (A. G. Tansley), 2nd ed. Bedford. Mag. 11: 271.
1969 A history of Bedfordshire (J. Godber). Bedford. Mag. 12: 128-30.
— Flora of Derbyshire (A. R. Clapham). Museums J. 69: 76-7.
1972 Flora of Monmouthshire (A. E. Wade); Flora of Rutland (G. Messenger). Museums J.
F233.
_— Watsonian Vice-counties of Great Britain (J. E. Dandy). Watsonia 9: 49-50.

1973 A computer-mapped flora (D. A. Cadbury, J. G. Hawkes & R. C. Readett). Ibid.: 282-5.
1975 Friends in Bedfordshire and west Hertfordshire (J. Godber). Bedford. Mag. 15: 126.
1976 The minutes of the first Independent Church at Bedford (H. G. Tibbutt). [bid.: 305-6.
, 1977 Atlas of breeding birds of Great Britain and Ireland (J. T. R. Sharrock); Atlas of non-marine

| mollusca of the British Isles (M. P. Kerney). Watsonia 11: 407.

-—— A nature conservation review (D. A. Ratcliffe). Bedford. Nat. 31: 9.

re John Howard the philanthropist (J. Godber). Bedford. Mag. 16: 82-3.

_1980 An ecological flora of Breckland (P. J. O. Trist). Watsonia 13: 153-4.

, 1983 Bird habitats in Britain (R. J. Fuller). Brit. Birds 76: 55-6.

— Simpson’s Flora of Suffolk. Watsonia 14: 437.

/1986 The book of Dunstable and Houghton Regis (V. & L. Evans). Bedford. Mag. 20: 170.

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BOOK REVIEWS
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B. VERDCOURT

JOHN DONY, HISTORIAN AND TEACHER

John Dony was a most remarkable man with limitless interests and boundless energy. He was a true
self-made man — he left school at 14, completed an engineering apprenticeship, and then, without
any full-time attention to study, qualified as a teacher, earned a respectable degree in Economics,
-and then secured a Doctorate. The record does not end there for, and this is well documented in the
‘bibliography above, there is a prodigious list of publications devoted to his two academic loves of
Botany and History.

‘| Although he was a prolific historian, I would think that his national reputation is greater in
Botany and that may be because much of his historical work focusses on Luton. However, even a
parochial interest must not be dismissed for what he produced is of considerable value and not just
to local people.

John was best known for his work on the Straw Hat industry (the subject of his Ph.D. thesis)
which was originally inspired more by the issues of labour relations than by the product itself. This
work grew out of an essay on trade unions and the hat industry first composed in 1931 for his
Economics tutor. Much encouraged by J. J. Mallon and Barbara Wootton, the subject was later to

62 OBITUARIES

be enlarged and developed into his doctoral thesis. John had no direct connection with the hat
industry but, as it was an important and lively economic activity in Luton, it offered a ready and
accessible research topic.

In concert with two others (Dyer and Stygall) he produced the definitive local history under the
title The Story of Luton. As a schoolboy it is known that he had little taste for history but his
enthusiasm was kindled by a W.E.A. class and he took to the subject with a passion, and it was that
very passion that made him an effective and entertaining teacher, for he made history challenging,
alive, and, importantly, relevant.

John was very much a son of Luton and he came from a well-known local family which lived
almost in the centre of the town. John’s grandfather supported the Co-operative cause and was a
founder-member of the Luton Industrial Co-operative Society. John’s share number, which he
inherited via an uncle, was 2. He was thus ‘related’ to the development of what was once a significant
social movement.

We were fortunate at Luton Grammar School to have been taught by scholars, but most were
drawn from relatively comfortable middle class backgrounds. John was not of that mould. Although
he was a success, he had had to struggle to be so. He was his own man and he brought to his many
activities a very marked sense of reality doubtless drawn from his experiences. For a man otherwise
destined for the machine-shop John won his place in the pantheon of great teachers. He was a man
who bore his achievements modestly and without display; he gave much of himself and was always a
ready friend and counsellor. His like is rare enough and it is an honour to be one of his ‘old Boys’.

M. BUTCHER

JOHN DONY AT THE LUTON MUSEUM

When I started work at Luton Museum as assistant to the curator in 1938 we were the only full-time
curatorial staff, but we had an Hon. Keeper of Botany, John Dony. On Saturday mornings and in
the school holidays he darted about the museum, working on the herbarium with a great sense of
purpose. He remained Hon. Keeper of Botany for the next 50 years. No museum can have had more
valuable and devoted service than he gave to Luton. He appeared only at these times because he was
teaching in Kentish Town and also working for a Ph.D. in Economics, having taken his first degree
two years before, by evening study after a full day’s teaching. He gained his doctorate for his thesis
on the straw hat industry in 1941. In the same year he took a post at Luton Grammar School,
teaching history and economics, and remained there until he retired, as Head of History, in 1964.

He was born, brought up and educated in Luton; worked in his teens as an apprentice engineer
there until 1920; and began his teaching career, first in Luton and then in Kentish Town (but still
living in Luton). His chosen study, the straw hat industry, was not just a subject for a thesis. He had
grown up in a town which had been shaped by this industry, which took from it its urban pattern, its
social nuances. It was the social and economic complexities of the industry which fascinated him.
His studies in local history and politics were never those of an academic specialist, but, as in his.
botanical work, were ‘habitat studies’.

His historical writings are mostly about Luton: A History of the Straw Hat Industry (1942); The
Story of Luton (1964); A History of Education in Luton (1970), enriched by his own experience as-
pupil and teacher there in the early years of this century; and The 1919 Peace Riots in Luton (1978),
in a volume presented to another Bedfordshire historian, Joyce Godber. Again he wrote from first
hand knowledge, a mildly participating witness of this tragi-comedy of local politics, in an essay
which shows how well he understood the character of the town he had lived in, worked in and served
all his life.

A. BucK

JOHN DONY AND THE B.S.B.1I.

John Dony’s influence on the Society’s development was second only to that of Ted Lousley in the
early post-War years. Throughout those two formative decades, the 1950s and 1960s, he was

OBITUARIES 63

continually in office: initially as the first-ever Field Secretary; in due course, for eight years, as
General Secretary; and ultimately as the third amateur to fill the Presidential chair in the new post-
War amateur/professional alternation. While, unlike Lousley, his involvement in the Society’s
affairs never extended to administering its finances and he was content to leave the publications side
of its activities to others, the nine years he devoted to organizing and overseeing its annual meetings
programme — a much more onerous responsibility than is generally appreciated — proved an
excellent preparation for running the Society as a whole, so that when Lousley stepped down as
General Secretary in 1956 no succession to that office can ever have seemed so natural and
appropriate.

Though very different personalities, the two of them worked together in harmony, united by a
common commitment to efficiency and shared battle scars from the endless in-fighting that had
characterized the régime that preceded theirs. John in particular never forgot or forgave, to the end
of his days, the airy casualness with which the supposed joint organizer had left him to cope alone,
almost at the last minute, with the arrangements for the all-important 1950 Conference at which the
Atlas of the British Flora had its original, carefully stage-managed conception. Nothing of the kind
ever happened again once the Society’s meetings programme was entrusted to his hands alone. Ina
very short time, indeed, the atmosphere was transformed: the conflicts and cantankerousness for
which some of the first post-War field meetings were notorious at once became a thing of the past, as
John brought to these occasions his chuckling sociability, his supreme meticulousness and his
unending tact. They were qualities from which the Society was to benefit more generally in his
subsequent spell as General Secretary and which rendered the team who served with him in those
years a memorably happy and smoothly-functioning one.

The meticulousness which he brought to the Society’s administration was the hallmark of his
personal botanical work too. Delighting in precision, with that tiny, figure-like handwriting
characteristic of the mathematically inclined, he loved nothing more than working carefully-
delimited areas with steadily increasing intensiveness. Deeply rooted in his home town of Luton
and surrounding countryside, he had the compiling of local Floras as his manifest botanical destiny.
And not just their compiling: his training in economics additionally gave him a taste for just those
aspects from which local Flora-writers typically tend to shrink, and he would spare no trouble in
working out costings and gauging market potential, matters on which he became the acknowledged
national expert and was ever-pleased to be asked for advice by fellow authors. It irked him greatly
that the handsome profit that the first of his Floras had unexpectedly made all went into the coffers
of its local government sponsor, yielding him no personal benefit financially in return for all the
effort he had put in, not least in earlier raising the money that had made publication feasible; and he
was determined ever afterwards to assume the risk of publication himself, convinced by that first
experience that works of this type were sound commercial propositions provided the production and
marketing received the careful attention they deserved. Not everyone felt able to share his optimism
on that score, which assumed an input of time and labour which few if any others were likely to be
prepared to contemplate (it extended, for example, to acting as your own publisher’s representative
and hawking copies in person to every bookshop in the county); but he was a lasting force for good
in making Flora-writers think more carefully and constructively about the stages that must come
after the accumulating of the records. The series of habitat studies which formed a major and novel
feature of the first of his Floras has had many copiers, while his switch from the traditional octavo
format to a quarto one for the succeeding Flora of Hertfordshire, to accommodate the printing of the
systematic list in double columns, which he perceived as dictated by cost-cutting logic, was so much
admired by K. G. Messenger that he modelled his Flora of Rutland exactly upon it.

Above all, John was identified with tetrad-mapping. If not quite the first to employ this method
(E. S. Edees anticipated him by just one year), he was certainly to the fore in exploiting it and in
promoting its adoption generally. The work that he undertook in this direction for his Hertfordshire
and Bedfordshire plant atlases so fascinated him that he even talked for a time of going to live in Co.
Carlow on his retirement and submitting that underexplored, similarly-sized county to the same
exhaustive treatment. If that was to remain but a pipe-dream, it was happily a different story with his
gradually-developing wish to research more deeply into the classically Watsonian question of the
number of plant species to be expected in an average stretch of ground — the subject of an impressive
paper that he contributed to Watsonia in 1963, in which his early statistical training came into its
own. A grant secured from the Natural Environment Research Council (a signal achievement in

64 OBITUARIES

itself) not only made that possible, but also largely allayed the financial anxieties that had began to
prey upon him as his time to retire approached.

Captivated by grid-square mapping and cursed with a home territory in which the administrative
boundaries had been subject to numerous and substantial changes, John became a dedicated
opponent of the long-standing vice-county system, repeatedly drawing attention to its drawbacks at
conferences and in informal discussion. Eventually he took the opportunity of his Presidential
Address in 1968 to advocate its wholesale abandonment and the adoption of 50-kilometre squares
as the Society’s unit for recording in its stead. No one could have put the case better, and he was
undoubtedly more disappointed than he ever let on that his arguments failed to win enough converts
to bring about that change.

It was probably the only major disappointment in a notably productive botanical life, in which he
raised to a new high level of achievement the long and proud tradition in these islands of producing
local Floras. It is as a local Flora-writer that he would surely wish to be remembered above all and
the capacity in which his contributions are most obvious and likely to prove most enduring. But the
B.S.B.I. more generally has cause to mourn the passing of a man who left his own very special mark
on it, in the course of the many years in which he served it with singular selflessness, endearing
himself to all of us who had the good fortune to know him.

DE: ALLEN

MARY PATRICIA HAPPER KERTLAND
(1902—1991)

Miss Pat Kertland had an enlightened upbringing and education which was Edwardian in the best
sense. In her youth she travelled widely, including a trip to the Far East, went up to her local
university and earned a higher degree. Her M.Sc. thesis, entitled ““The ecology of Divis”, was
written while she was Hugh Wisnom Scholar at the University and formed the basis of a major paper
in J. Ecol. 16: 301-322 (1928).

She was blessed with the ability to use ‘family money’ without ostentation and was able to live
much of her life in the way she wished, without the absolute requirement to hold the highest possible
salaried post, though by training and intellect she would have had little difficulty in securing a senior
position in any University.

She worked in the Botany Department of The Queen’s University of Belfast, in which she had
been a student, from 1929 to 1937 as Honorary Herbarium Assistant, as Demonstrator in Botany
from 1937 to 1945 and as Curator of the Herbarium from 1945 to 1967. Within these few lines is
encompassed a vast amount of work both personal and in the inspiration of generations of students,
for although the Department was never large, it produced a number of outstanding botanists over
the years on all of whom she had an influence; B. E. S. Gunning, M. Morrison and J. S. Pate
immediately coming to mind. She was a colleague of Professor James Small and his successors Jack
Heslop-Harrison and Eric Simon. The Herbarium collection was greatly enhanced and completely
reorganized during her Curatorship. Her last great task was to oversee its transfer to the Ulster
Museum (BEL) in the spring of 1968, though, technically, she had retired in the autumn of 1967.
Her taxonomic work was meticulous and through it she got to know a large group of experts across
the world.

From 1951, she was formally involved with the /rish Naturalists’ Journal; at first as Assistant
Editor with A. W. Stelfox for two years, then as Editor until 1976, when Mrs Elizabeth Platts
succeeded her. After her retirement she continued to give both of her time and material resources
and remained on the management committee until her death. A short appreciation of her period as
Editor appeared in Jr. Nat. J. 19: 1-2 (1977) and, with it, a reproduction of the pencil portrait
created in 1973 by Raymond Piper which shows her in repose.

Through the Belfast Naturalists’ Field Club she reached a wide audience as excursion leader and
lecturer. Through the B.S.B.I. she became known to a wide spectrum of botanists and was unfailing
in her assistance to members, at least to those of whom she approved, in the discovery, or
rediscovery, of rare species across Ireland.

OBITUARIES 65

The Jrish Naturalists’ Journal fitted well into Pat Kertland’s persona: it gave her the opportunity
to do good by stealth. As Editor she encouraged the amateur naturalist in every way possible, often
taking the barest bones of a scientific note and converting it to an acceptable technical standard.
One recent project dear to her heart was the creation and publication of the Flora of Lough Neagh
by John Harron (1986). This book, dedicated to her, reflects the encouragement of others and the
drive which epitomised her life. She recognized the value of the data, and associated record
specimens, which the author was collecting and, over a long period, catalysed a team to help him
convert this raw material into the finished product.

I was privileged to be one of Pat Kertland’s students in the early 1950s at a time when she took on
extra duties after Dr Mary Lynn’s early retirement due to unstable health (though happily she is still
with us today). I can vouch for her ability as a lecturer, though, on field trips, her propensity to take
both hands off the steering wheel to point out features of interest was disconcerting, to say the least.

In recent years ill-health struck cruelly; firstly increasing deafness and then sudden total
blindness. This did not break her spirit: she moved to a nursing home, organized readers and
continued to make her distinctive contribution to life.

I can do no better than end with a quotation from the Address by the Rev. Dr David Lapsley
given at her funeral in late February 1991: “Most of all she is held in deep respect and affection
because of her interest in people. She was a source of encouragement to others, this honest woman,
with a manner that could be formidable and brusque. She appeared not to suffer fools gladly, but
her words and deeds were constantly flavoured by kindness and generosity.”

W. D. LINTON

EVAN ROBERTS
(1906—1991)

When Evan Roberts died on 15 May 1991, at the age of 84, Wales lost one of her most outstanding
field botanists of this century, a man whose special interest was the distribution and ecology of the
arctic-alpine plants of Snowdonia and who became the acknowledged authority on them.

He was born in Capel Curig, the little village in which he was to spend the whole of his life. Even
when he married, he went to live only a mile away, and following the death of his wife, Mabel, in
1969, moved back to the house where he was born, to spend his latter years with one of his sons.

On leaving the village school at 14, he found employment at the local slate quarry and worked
there almost continuously until it closed in 1953. The work was hard, often dangerous, and poorly
paid. It was these circumstances which eventually drove Evan on to the course which was to give his
life a new dimension and himself so much pleasure and satisfaction. For, having married when he
was 23, the increasing needs of his young family forced him to give up his treasured motor-cycle and
consequently spend more time on the hills around Capel Curig.

As he was fond of recalling, one Saturday afternoon during this period proved particularly fateful.
He took the path up Moel Siabod, the hill immediately behind his home, and, just before reaching
the summit, came across a plant with beautiful purple flowers which he had never seen before. It was
only after several inquiries that he found its name: the plant which had captured his imagination was
Saxifraga oppositifolia. For him it proved to be the key to a new world and from then on Evan spent
much of his spare time searching for it on other hills around Capel Curig. He got hold of a copy of
J. E. Gniffith’s Flora of Anglesey and Carnarvonshire and thereafter searched for other arctic-alpine
plants such as Lloydia serotina, Polygonum viviparum and many others.

A chance meeting with E. Price Evans, the notable ecologist, gained Evan a friend who had
considerable influence on him. For Price Evans urged him to keep a full list of all the plants he
found, with details of their locality, altitude, aspect and the nature of the local rock. Thus began the
most thorough and systematic account ever made of Snowdonia’s arctic-alpine plants and all that
grew with them. Years of careful and laborious work went into the survey, so that at its completion
the plant life of this area was better known than it had ever been before. Rare plants thought to be
extinct were rediscovered; others were found in new localities where they were formerly unknown,
such as, for instance, his discovery of Dryas octopetala on the remote cliffs of Creigiau Gleision,

66 OBITUARIES

8 km from its previously known site above Cwm Idwal. When he took Price Evans there to see it
some time later, they found Carex capillaris near by, a new record for Wales and an interesting
addition to the mountain flora of Snowdonia. Happily, both plants still survive in this remote spot,
where they may be seen together with Saxifraga oppositifolia, Polystichum lonchitis and many other
species found here by Evan.

In addition to his dedication and enthusiasm (and in spite of a defect in one eye) Evan was well
qualified for the work he undertook — he was hardy and a skilful climber. During World War II he
assisted in training Commando units in the techniques of rock-climbing. It was therefore no surprise
that when the Nature Conservancy acquired Cwm Idwal as its first National Nature Reserve in
Wales, in 1954, Evan was appointed to be its Warden. He was later promoted to be the Chief
Warden in North Wales, a position he held until his retirement.

In this new post his unfailing good nature, unruffled manner and natural courtesy, especially
when dealing with those who sometimes transgressed the code of conservation in the Reserves,
made him universally respected. His generosity and readiness to share his knowledge of the
mountains and their flora with all who sought his help, made him many friends over the years. His
services were in great demand to lead parties of students and others around the Reserves and,
indeed, anywhere in Snowdonia. The illustrated lectures he gave during the winter months (both in
his native Welsh and in English) became very popular and did much to further an understanding of
the conservation. movement among the general public in North Wales.

Evan’s concern for conservation was apparent long before he joined the staff of the Nature
Conservancy. I recall his deep disappointment when a small bog near Capei Curig, in which, in
1946, he had found Hammarbya paludosa, was destroyed by forestry operations a few years later, in
spite of his appeal for its preservation. But I have happy memories, too, of many botanical
excursions with him on fine summer days to Cwm Glas and Clogwyn y Garnedd, where he could
show me most of the rare plants as unerringly as if they were in his own back garden; and of one
occasion when he took me to a rock crevice high up on Snowdon to see the very rare Woodsia alpina.
To be with him for a day on the mountains was an inspiration, for he knew Snowdonia, its cliffs and
crags, gullies and ledges like the palm of his hand.

Although he published very little, his detailed records were made availne to the Nature
Conservancy. Copies of them have also been deposited in the Library of the National Museum of
Wales in Cardiff.

The value of his work on the mountain flora of Snowdonia was recognized by the University of
Wales, in 1956, when it awarded him the richly deserved honorary degree of M.Sc. He was later to
receive the M.B.E. for his services to conservation over a long period, often beyond the call of his
official duties. He was, indeed, a remarkable man and his loss is irreplaceable. He will be
remembered with affection and gratitude by those who knew him.

We extend our deep sympathy to his three sons and their families in their bereavement.

R. H. RoBerts

Watsonia, 19, 67-71 (1992) 67

Report

ANNUAL GENERAL MEETING, 4 MAY 1991

The Annual General Meeting of the Society was held at the Merseyside Maritime Museum,
Liverpool, at the invitation of the National Museums and Galleries on Merseyside, at 11.15. 95
members were present and Mr J. Ounsted, Vice-President, took the Chair.

Apologies for absence were read and Minutes of the 1990 Annual General Meeting, as published
in Watsonia 18: 331-332 (1991), were approved and signed by the Chairman.

REPORT OF COUNCIL

The adoption of this report, which had been circulated to members, was proposed from the Chair,
seconded by Dr E. C. Nelson and accepted nem con.

TREASURER’S REPORT AND ACCOUNTS

The Hon. Treasurer, proposing the adoption of his Report and Accounts, offered to explain any
points. Dr T. C. G. Rich asked for clarification of the terms of covenant of subscriptions. Mr
Walpole replied that the covenants must be for a minimum of four years. The adoption of the
Report was seconded by Mr R. G. Ellis and Dr P. Macpherson, who coupled this with a vote of
thanks to the Treasurer. This was carried unanimously with applause.

SUBSCRIPTION INCREASES FROM JANUARY 1992

Following the general rise in costs, particularly of postage, Council had proposed the following rates
from 1992:

Ordinary £12.50 to £15.00

Family £1.00 to £2.00
Junior £5.00 to £6.00
Senior £7.50 to £9.00

Institutional £12.50 to £15.00

The Hon. General Secretary noted that there had been no increase in subscriptions since 1986.
Mrs M. Lindop, relaying a query from a member unable to be present, asked for clarification of the
Senior rate. The Hon. General Secretary read the definition as in B.S.B.I. Rule 24: “Persons over
60 who have been members of the Society for at least 10 years and who are no longer in full time
employment may elect to pay an annual reduced subscription at such rate as from time to time shall
be decided by Council”. It was agreed to republish this rule in B.S. B.J. News for the information of

‘members. The adoption of the new rates was proposed by the Hon. Treasurer, seconded by Mrs A.
Lee, and carried with none against.

ELECTION OF PRESIDENT

The election of Dr P. Macpherson F.R.C.P., F.R.C.R., F.L.S., as nominated by Council, was
proposed from the Chair and carried unanimously with acclamation. Dr Macpherson was then
welcomed and took the Chair. Thanking Mr Ounsted, he also thanked members for his election,
saying that this leap to President had in no way been anticipated, but that he was humbly and deeply

68 REPORT

appreciative. The President then called on Dr J. R. Akeroyd to thank the retiring President,
Professor D. A. Webb Sc.D., F.M.L.S., who was unfortunately unable to be present at the meeting.

Dr Akeroyd, recalling that he and Professor Webb had shared the same biology teacher at
Charterhouse, the late Percy ‘Cheese’ Chapman, reminded us that Professor Webb was first a
marine biologist, whose research over the years had ranged from the blood pigments of sea-squirts
to Saxifraga. Professor Webb, who had been one of the driving forces of Flora Europaea and had
dominated Irish botany since the death of Praeger, had been an active President of this Society,
frequently making the long journey to attend meetings and supporting all the B.S.B.I. activities
during his term of office. He thanked Professor Webb on behalf of the Society and this was
acclaimed with applause.

ELECTION OF VICE-PRESIDENTS

The election of Mr P. S. Green, Dr G. Halliday and Mr A. C. Jermy, as nominated by Council, was
proposed by Dr R. J. Pankhurst, seconded by Mr R. M. Burton and carried unanimously.

RE-ELECTION OF HON. GENERAL SECRETARY AND HON. TREASURER

Proposing the re-election of Mrs M. Briggs, the President recalled that, when invited to accept
nomination for President, he had asked if a medical certificate ensuring the good health of the Hon.
Secretary was available. He particularly thanked the Hon. Secretary for the Annual Report of
Council and this year for the first of a new series of the Year Book which had been widely acclaimed
by members. Proposing the re-election of Mr Walpole, the President adding to his earlier vote of
thanks, referred to the considerable workload of the Treasurer for the Society. The elections were
formally seconded by Mrs A. Lee and Mr R. G. Ellis and carried unanimously with applause.

ELECTION OF COUNCIL MEMBERS

In accordance with Rule 10, nominations had been received for Dr J. R. Akeroyd, Mr J. M.
Montgomery and Dr T. C. G. Rich. Their election was proposed by Mr R. G. Ellis, seconded by Dr —
R. J. Pankhurst and passed unanimously.

MINOR AMENDMENTS OF THE RULES OF THE SOCIETY

Some changes in the wording of eight of the Society’s Rules, to incorporate changes of Rules
approved by Annual General Meetings since 1968, and to consolidate the wording, had been
discussed and approved by Council. Copies of the amended Rules were available at the meeting,
and had been offered to members by post beforehand on request. The amendments were accepted
without comment.

ELECTION OF HONORARY MEMBERS

Mr R. G. Ellis, proposing Mr M. Walpole F.C.A., F.L.S., spoke of him as a keen supporter of
conservation and active member of his local Wildlife Trust, and as a botanical book collector with
the finest private collection in this country. For the Society Mr Walpole was a decisive Chairman of
the Publications Committee, and, most significantly, as Hon. Treasurer he has efficiently handled
B.S.B.I. finances for 20 years.

Proposing Mr A. O. Chater, Mr Walpole noted that some societies awarded medals to individual
members as a token of appreciation of distinction in service to the society. Mr Chater, a competent
Secretary of the Publications Committee for nine years, was co-author of Sedges of the British Isles
and had also been very involved in the publication of five other titles in this series of B.S.B.I.

REPORT 69

Handbooks. His general botanical advice has been valued by the officers and members alike, and as
a Carex Referee and the Specialist on Nomenclature his adjudications are generously offered and
highly respected.

Before the meeting neither recipient knew that they had been nominated by Council; both were
unanimously elected with warm applause.

RE-ELECTION OF HONORARY AUDITORS

the Hon. Treasurer, in proposing the re-election of Grant Thornton, West Walk, Leicester,
referred to the honour for the Society to present their Accounts over the name of these
distinguished Auditors. Their re-election was passed unanimously, and the President agreed to
write expressing thanks from the Society.

ANY OTHER BUSINESS

Mr M. Walpole, as Chairman of the Publications Committee, reported that the Committee had
appointed two new Watsonia Editors: Dr R. R. Mill and Dr E. C. Nelson. Both were present and
were welcomed. Dr R. J. Gornall had retired and Dr B. S. Rushton would now be Receiving Editor.
Thanking Dr Gornall on behalf of the Society he observed that the high standards of Watsonia had
increased during the ten years of Dr Gornall’s editorship. He was thanked for competently coping
with the very heavy workload, and for his skill at conforming to prescribed budgets. Thanks were
extended also to all the Society’s Editors, not least to Mr R. G. Ellis (B.S. B.I. News) and Mr D. H.
Kent (B.S.B.I. Abstracts).

The Report of the B.S.B.I. Monitoring Scheme by Dr T. C. G. Rich was complete in two large
format volumes, of which copies of Vol. II (the maps), sufficient for distribution to v.c. Recorders,
were now available, and a matching number of copies of Vol. I were in production. Because of their
large size, publication of further copies would be costly and, at the request of Council, the Hon.
Secretary asked how many of the members present (who were not v.c. Recorders) would be
interested in purchasing copies if these were to be published at approximately £20. Almost all those
members signalled by a show of hands as wanting to purchase a copy if published. Possible ways to
reduce the cost, e.g. smaller paper size for Vol. I, or ashortened text as an Introduction to the maps,
were discussed. Dr Rich noted that he had submitted three papers to Watsonia on the results of the
Scheme. Dr Rushton pointed out that if these papers were accepted by and published in Watsonia,
they could not be published elsewhere. A lengthy and lively discussion followed and the matter was
referred to Council with some urgency, in view of the time that had elapsed since the end of the field
work and the completion of the Report.

Following some general announcements, the President then thanked Dr J. R. Edmondson and his
assistants for the excellent local arrangements and the meeting closed at 12.23.

Mary BriGGs

PAPERS READ AT THE ANNUAL GENERAL MEETING

Four papers were presented on the theme of “Botanists and Books’’. Michael Walpole first spoke on
British Floras and the book collector, explaining his personal reasons for collecting local Floras and
sharing some anecdotes on the sometimes difficult relationships with antiquarian booksellers. He
found it particularly difficult to confine his collecting to a severely restricted field, given the large
number of publications which have botanical data in their contents.

Charles Nelson then gave a talk entitled ‘“Out of olde bokes . . . cometh al this newe science”:
musings on bibliography and botanists. Aptly complementing the previous speaker, he chose a
series of examples to illustrate his theme that it was not only the printed contents of the books which
were of significance to posterity, but also the jottings, scribblings and more deliberate mutilations of

70 REPORT

books (such as bookplates — several exuberant examples were shown) which provide rich pickings
for botanical historians.

After the tea break, Tim Rich presented a paper Towards an Atlas of the British Flora 2000, the
date being millennial rather than a firm prediction. Based on his experience during the recently
completed Monitoring Scheme project, he outlined various methods for the efficient capture of
distributional data and made a strong plea for the continuation of involvement in field recording
which would lead to the publication of a new Atlas.

The final speaker, Bob Barnard of H.M.S.O. Publications, gave a professional publisher’s
account of The new technology of publishing botany books, showing examples from various recent
H.M.S.O. publications to demonstrate new methods of typesetting, illustrating and distributing
books. Reference was made to CD-ROM technology as well as to more conventional printed
products, with helpful guidance on the various ways in which publishers price their products.

THE NATURAL HISTORY BOOK FAIR

This was the first occasion on which a Book Fair had been arranged as a commercial venture at the
Maritime Museum. It was organized by the Botany Department of Liverpool Museum to take place
in conjunction with the Annual General Meeting. The following booksellers participated:

B.S.B.I. Publications (Oundle)

Broadleaf Books (Cardiff)

Chantrey Books (Sheffield)

John Price (Marlborough)

Parry Books (Liverpool)

Subbuteo Natural History Books (Treuddyn)

Summerfield Books (Brough)

Tim Oldham, of Wyseby House Books, had to withdraw at a late stage because of illness.

It was generally agreed that this event enhanced the value of a conference on “Botanists and
Books” as well as providing tempting purchasing opportunities for bibliophiles. The agreeable
atmosphere of the Maritime Museum’s historic warehouse galleries was also appreciated.

EXHIBITION: SOME NORTH-WEST BOTANISTS AND THEIR BOOKS

A small display was mounted by the Botany Department of Liverpool Museum containing short
biographical sketches of some of the principal authors of botanical books in N.W. England together
with examples of the books themselves as well as items from the archives. One of the highlights of
the exhibition was a copy of Thomas Johnson’s 1633 edition of Gerard’s Herball.

The following botanists with North-west connections were featured: William Bean (1817-1864);
Joseph Dickinson (1805-1865); John Gerard (1545-1612); Conrad Theodore Green (1863-1940);
Leopold Hartley Grindon (1818-1904); Thomas Batt Hall (1814-1886); William Roscoe (1753-
1831); William Gladstone Travis (1877-1958); John Warren, Baron de Tabley (1835-1895); James
Alfred Wheldon (1862-1924) and Albert Wilson (1862-1949).

Also on show were samples of the manuscript of de Tabley’s Flora of Cheshire, together with
examples of specimens from his herbarium, on loan from the National Botanic Gardens, Glasnevin,
Dublin. The organizers are indebted to the authorities at Glasnevin, as well as to Mr Neville
Carrick, the City of Liverpool’s Librarian and Mr Eric Greenwood, Keeper of the Liverpool
Museum, for the loan of items for this exhibition.

A leaflet entitled Some North-west Botanists and their Books was prepared by Lorna Cunliffe-
Lister, a student from the Department of Museum Studies, University of Leicester, to accompany
the exhibition.

REPORT Til

FIELD EXCURSION HELD IN CONJUNCTION WITH THE A.G.M.
BODNANT GARDEN AND THE GREAT ORME, NORTH WALES. 5 MAY 1991

A coach party of 42 people together with about 20 travelling by car made the journey to Bodnant
Garden, near Conwy, on a day which started cloudy but developed into a gloriously sunny
afternoon. The garden at Bodnant, which was given to the National Trust by the late Lord
Aberconway, is set in a precipitous valley close to the estuary of the River Conway. It is managed
partly as a formal garden and partly as a ‘wilderness’ garden with steeply sloping rocky banks and a
network of crazily angled paths. The facilities have been enhanced by a garden centre (under
refurbishment) and a new restaurant and coffee shop which, despite the Bank Holiday crush, was
able to offer an efficient luncheon service.

After lunch the coach departed for Llandudno, hoping to take its passengers to a rendezvous with
the car-borne botanists at the foot of the Great Orme. Alas, Bank Holiday crowds attending a
“Victorian Extravaganza” in Llandudno, coupled with the closure (due to a landslip) of the coastal
road which encircles the peninsula, conspired to delay the coach to the point where the two parties
set off separately for the limestone massif of the Great Orme. Led by three local botanists, Wendy
McCarthy, Ted Phenna and Geoff Battershall, 17 members set off on an ‘alternative A.G.M.
excursion’. Numerous specialities were seen including Aster linosyris, Hypochoeris maculata,
Helianthemum canum, Hornungia petraea, Orchis morio and Cotoneaster integerrimus. The coach
party, having finally reached the ungrazed area below the cablecar station, were able to scramble up
the slopes above the foot of the Happy Valley gardens in order to see some of the characteristic
early-flowering limestone plants, including Scilla verna, and Brassica oleracea at its northern limit of
distribution in Wales.

Thanks are due to Tim Rich for contributing to this report, to all the speakers and exhibitors at the
Book Fair, to the staff of the Conference Office at the Merseyside Maritime Museum for their
efficient handling of the front-of-house arrangements, and to the Trustees of the National Museums
and Galleries on Merseyside for their hospitality.

J. R. EDMONDSON

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DAE

B.S.B.I. Conference Reports

BRITISH FLOWERING PLANTS AND MODERN SYSTEMATIC METHODS
Ed. A. J. Wilmott, 1948, 104 pages, 18 plates. £5.25.

. THE STUDY OF THE DISTRIBUTION OF BRITISH PLANTS.

Ed. J. E. Lousley, 1951. 128 pages, illustrations and maps.

. THE CHANGING FLORA OF BRITAIN

Ed. J. E. Lousley, 1953. 203 pages, 9 plates, 25 text figs.
SPECIES STUDIES IN THE BRITISH FLORA

Ed. J. E. Lousley, 1955. 189 pages, 2 plates and 23 text figs. £5.25.
PROGRESS IN THE STUDY OF THE BRITISH FLORA

Ed. J. E. Lousley, 1957. 128 pages, 4 plates and 9 text figs. £5.25.

. A DARWIN CENTENARY

Ed. P. J. Wanstall, 1961. 140 pages, 7 plates, 12 text figs.

. LOCAL FLORAS

Ed. P. J. Wanstall, 1963. 118 pages, 1 plate (map), 9 text figs.

. THE CONSERVATION OF THE BRITISH FLORA

Ed. E. Milne — Redhead, 1963. 90 pages.

. REPRODUCTIVE BIOLOGY AND TAXONOMY OF VASCULAR PLANTS

Ed. J. G. Hawkes, 1966. 182 pages, 1 plate, 9 text figs.

. MODERN METHODS IN PLANT TAXONOMY

Ed. V. H. Heywood, 1968. 312 pages, numerous text figs.
Held in association with the Linnean Society of London.

. THE FLORA OF A CHANGING BRITAIN

Ed. F. H. Perring, 1970. 158 pages, 21 text figs. £3.50 (1973 reprint).

. TAXONOMY, PHYTOGEOGRAPHY AND EVOLUTION

Ed. D. H. Valentine, 1972. 431 pages, numerous text figs and tables. Held in association with
The Linnean Society of London and the International Organisation of Plant Biosystematists.

. PLANTS WILD AND CULTIVATED

Ed. P. S. Green, 1973. 232 pages, 8 plates and 24 text figs. £3.20.

. THE OAK: ITS HISTORY AND NATURAL HISTORY

Ed. M. G. Morris & F. H. Perring, 1974. 376 pages, illustrations. £8.25

. EUROPEAN FLORISTIC AND TAXONOMIC STUDIES

Ed. S. M. Walters, with the assistance of C. J. King, 1975. 144 pages and 4 plates. Held in
association with the Linnean Society of London. £3.80.

THE POLLINATION OF FLOWERS BY INSECTS

Ed. A. J. Richards, 1978. 213 pages and 31 plates. Held in association with the Linnean
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Contents

F. & GROENHOF
Ty

ological

AN ok ::
cage?
Sak

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,
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Elected at the Annual General Meeting, 4th May 1991
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Editors of Watsonia

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Plant Records, C. D. Preston

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“Receiving editor, to whom all MSS should be sent (see inside back cover).

; @% @ rae eH is Bt : eu M
Watsonia, 19, 73-95 (1992) : (HATSRAL BISTORD)

- 2 SEP 1992

Recording bias in botanical surveys OCAAY CIRHARY

er SUSI SR EIS BEE RII |

T. C. G. RICH* and E. R. WOODRUFF

Biological Records Centre, Monks Wood Experimental Station, Abbots Ripton, Huntingdon,
Cambs.; PEI7 2LS

ABSTRACT

Recording bias in botanical surveys arises primarily from the recording behaviour of individual botanists,
sometimes coupled with the survey techniques and the types of plants being recorded. Recording bias is
probably widespread in botanical surveys; it does not invalidate the records, but requires that care is taken with
interpretation. Some generalizations are made to help assess recording bias, and are illustrated using examples
found during the B.S.B.I. Monitoring Scheme.

INTRODUCTION

In any botanical survey, there is an inevitable degree of recording bias (Hope-Simpson 1940; Sykes
et al. 1983; Nilsson & Nilsson 1983; Kirby et al. 1986; West & Hatton 1990, etc.). Efforts are
therefore usually made to minimise bias by adopting a controlled, systematic, repeatable method
(Greig-Smith 1964).

In the recording techniques adopted for most national or county plant atlases (e.g. Perring &
Walters 1962; Hall 1980), recorders individually select the areas within squares to visit, and then
record the species present to fill the time available, or until the list is felt to be comprehensive. Such
an unstructured recording technique might be expected to introduce considerable local bias to the
data collected, and especially if the surveys are to be repeated. For instance, only 52% of the records
collected by two independent parties of botanists six weeks apart in the same tetrads (2-km squares)
were common to both surveys (Rich & Woodruff 1990, 1992). Efforts are often made to achieve
even coverage (e.g. Dony 1963), but failure to achieve this is only one of many sources from which
bias can arise. Documentation of what was done and where and when and by whom can help with
interpretation of the data, but such details are rarely collected.

Bias originates primarily from differences in the recording behaviour and the ability of individual
botanists, but may also arise from the survey techniques or the types of plants being recorded. The
extent of bias in the data collected is rarely known, and few cases are described in detail. Perring &
Walters (1962) presented provisional distribution maps of about 40 taxa and explained why the data
were thought to be inadequate. Nilsson & Nilsson (1983) found that sampling error accounted for
two-thirds of the apparent species turnover rates on islands in Sweden. Preston & Eversham (1992)
describe selected examples of botanical and zoological recording bias.

The purpose of this paper is to draw attention to the problems of recording bias by using some
examples encountered during the B.S.B.I. Monitoring Scheme (Rich & Woodruff 1990, 1992). The
B.S.B.I. Monitoring Scheme was a 10-km square sample survey of Britain and Ireland during 1987
and 1988 to assess the current status of the flora. Over i600 botanists collected 985,000 records in
425 out of the 429 sample 10-km squares, representing 2660 taxa. Many of the examples are drawn
from a comparison of these data with those collected for the Adlas of the British florat (Perring &
Walters 1962).

Nomenclature follows Clapham et al. (1987).

* Present address: 24 Lombardy Drive, Peterborough, PE1 3TF
+ hereafter referred to as the Aflas.

74 T. C. G. RICH AND E. R. WOODRUFF

RECORDING BIAS IN BOTANICAL SURVEYS

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76 T. C. G. RICH AND E. R. WOODRUFF
EXAMPLES OF RECORDING BIAS

NOTE ON PRESENTATION OF THE MAPS

The maps show only data from the 10-km squares sampled for the B.S.B.I. Monitoring Scheme (one
in every nine; see Rich & Woodruff 1992). The symbols are enlarged so that they are clear to read
when the maps are reduced and do not indicate the actual areas covered. The symbols used are as
follows:

¢ Recorded only for the Monitoring Scheme (1987-1988).

@ Recorded for both the Atlas and the Monitoring Scheme.

Thus, a predominance of open circles may suggest a decline in relative frequency between the
surveys, and a predominance of pluses, an increase.

THE IMPORTANCE OF ASSESSING BIAS

The importance of assessing bias can be seen from the apparent similarities between the following
selected pairs of maps. Unless the extent of recording bias is known, any conclusions drawn from the
data may be misleading.

Rubus vestitus and Lagarosiphon major (Figs 1a, b): The increase in records for the former
species reflects an increase in recording of critical taxa coupled with incomplete historical data; for
the latter it reflects a real increase in the frequency of the plant.

Agrostemma githago and Polygonum nodosum (Figs 1c, d): The former species has decreased
markedly during the last 30 years; the latter has undergone a taxonomic revision and is no longer
recognised by most recorders.

BIASES RELATED TO THE QUALITY AND QUANTITY OF RECORDING

It is often said that the distribution of plants reflects the distribution of botanists, and this is
particularly true for the more critical taxa and for small areas. Fig. 2 shows how the apparent
distribution of Carex hostiana X viridula correlates with areas recorded by A. O. Chater and J.
Harron who know this obscure hybrid well. It is, however, probably widespread in the north and
west where its parents grow together. Another botanist, M. Porter, recorded critical taxa in great
detail in Brecon for the Monitoring Scheme, resulting in apparently highly localised concentrations
of records of species of Rubus, Hieracium, Taraxacum and Euphrasia (Rich & Woodruff 1990).

Taxonomic awareness and recording fashions may bias results on a wider scale and for commoner
taxa. Many critical groups have been more widely recorded for the Monitoring Scheme than for the
Atlas (e.g. Hieracium, Fig. 3a), but some were more widely recorded for the Adlas (e.g. Rhinanthus,
Fig. 3b), resulting in artificial changes in frequency. Compared to a general average of 16% more
records for the Monitoring Scheme than the Adlas, there are 24% more records for five selected
genera (Carex, Polygonum, Populus, Rumex and Salix) covered by the B.S.B.I. Handbook series
(Jermy et al. 1982; Kent & Lousley 1981; Meikle 1984). Trist & Sell (1988) drew attention to the
occurrence of two subspp. of Molinia caerulea in the British Isles; there were four records of the
subspp. in 1987 and 33 in 1988. These increases in records are no doubt due to increased awareness
of the taxa concerned. Fig. 4 shows 10-km squares where above average percentages of critical taxa
were recorded for the Atlas and Monitoring Scheme surveys; the squares correlate well with areas
known to have been well-recorded.

There are differences in opinion between recorders over which introduced species or garden
escapes to record. In recent years it has also become more acceptable to record all introductions as
they have become more widespread in the wild. Consequently, exaggerated rates of increase may be
observed for species which were present but often ignored during recording for the Aflas (Fig. 5).
However, it is also still more acceptable to record some introductions than others — taxa are more
likely to be recorded if they are included in national or local Floras or if listed on the record cards.
Crops such as Wheat (Triticum aestivum) and Barley (Hordeum vulgare), which are not listed on
cards, are poorly recorded on roadsides compared to others, such as Oil-seed Rape (Brassica napus)
(Fig. 6), which are listed on the cards and are widely recorded.

Bias also arises from differences in taxonomic opinion and also from common errors; many
recorders simply follow the major floras. There is little agreement on the current taxonomic status of
the subspp. of Juncus bulbosus, resulting in confusion between the records (Fig. 7a, b). Viola canina

RECORDING BIAS IN BOTANICAL SURVEYS

A. O. Chater, JH = J.

Figure 2. Correlation between (a) the distribution of Carex hostiana X viridula and (b) the areas recorded by two botanists (AC

Harron).

T. C. G. RICH AND E. R. WOODRUFF

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RECORDING BIAS IN BOTANICAL SURVEYS

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Ficure 4. 10-km squares with an above average % of critical taxa recorded (number of critical taxa/total number of taxa recorded per square): (a) Atlas survey; (b)

Monitoring Scheme survey. Britain and Ireland have been assessed separately due to inherent differences in the flora. Open circles show 10-km squares not

recorded for both surveys.

T. C. G. RICH AND E. R. WOODRUFF

Ficure 5. Examples of increased recording of garden escapes resulting in exaggerated rates of increase: (a) Lunaria annua; (b) Ribes sanguineum.

RECORDING BIAS IN BOTANICAL SURVEYS

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82 T. C. G. RICH AND E. R. WOODRUFF

RECORDING BIAS IN BOTANICAL SURVEYS

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84 T. C. G. RICH AND E. R. WOODRUFF

may have been over-recorded inland for the Atlas (Fig. 7c), possibly because in Bentham & Hooker,
the Flora that most British botanists at that time had been raised on, V. riviniana was included in V.
canina (F. H. Perring, pers. comm.). Hyacinthoides hispanica has been over-recorded in error for
the much more common but less well known Hyacinthoides hispanica x H. non-scripta (Page 1987;
Fig. 7d); the latter is not included in Clapham et al. (1987).

Other plants may simply be overlooked if recorders are unaware of their presence in an area.
Festuca altissima is an uncommon plant of rocky woods and ravines and was almost certainly under-
recorded for the Atlas (Fig. 8a). Chenopodium ficifolium, a weed similar in appearance to the
common C. album, may also have been overlooked in some areas (Fig. 8b).

Some habitats may be recorded better or more poorly than others due to differences in
accessibility. Arctic-alpines such as Luzula arcuata and Juncus castaneus in Scotland, and
Polygonum viviparum in Ireland, were under-recorded for the Monitoring Scheme partly due to the
remoteness of the localities and partly due to inclement weather. Conversely, car-parks and
churchyards have been well-recorded as they are easily accessible.

Increases in numbers of records may result simply from increased recording effort for the
Monitoring Scheme (Rich & Woodruff 1992). Numerous examples could be cited, but a clearer
demonstration is probably from outside the Scheme itself where the effects can be seen in a wider
context. The spread of Cardaria draba has been documented by Scurfield (1962). Fig. 9 shows the
cumulative number of 10-km squares from which C. draba has been recorded. The enormous
increase in the 1950s coincides with the Adlas field work and reflects a simple increase in recording
effort rather than a dramatic spread of the plant. Similar patterns can also be seen in Epilobium
ciliatum, Veronica filiformis and Impatiens glandulifera. The Monitoring Scheme results suggest
little increase in C. draba since 1960.

BIASES RELATED TO RECORDING METHODS

Constraints imposed by the recording methods may result in some systematic biases in addition to
those introduced by the botanists. Bias introduced by changes in the areas recorded, the
repeatability of surveys, concentration on the selected A, J and W tetrads, and by the time span of
recording have been briefly discussed by Rich & Woodruff (1990, 1992). Examples of how changes
in coverage and in the areas recorded may result in apparent increases of plants are shown in Fig. 10.
Plantago maritima was recorded for the Monitoring Scheme in 22 out of the 26 coastal 10-km
squares not recorded for the Atlas; if these records are ignored there is no significant change in
frequency. Similarly, apparent increases in Trifolium repens around the coast are due to the new
squares being recorded, and apparent losses in Ireland are squares not re-recorded for the
Monitoring Scheme. Examples of species under-recorded in the Dublin square due to concentration
on the selected tetrads are Oenanthe aquatica and Myriophyllum spicatum.

Exaggerated rates of decline of casual species may result from different time spans of surveys.
Casuals or fugitives (Preston & Eversham 1992) are usually short-lived, non-persistent species
which are unpredictable in occurrence (they are often accidentally introduced by man). As records
accumulate with time, a longer survey period ts likely to yield more 10-km square records than a.
shorter period. Table 1 shows the number of sample 10-km squares recorded for three arable weed
species; in all cases there are considerably more records for 1930-1960 than 1987-1988 and it might
be concluded that the species have declined by 75% or more. If the numbers of records per year are -
calculated, all taxa appear to have become more frequent. Neither conclusion is strictly valid
because the numbers of records also need to be corrected to take into account the amount of
recording effort which generated them. There 1s little doubt that these species have declined, but at
a lower rate than suggested by a simple comparison of numbers.

The taxa listed on the cards may introduce bias by prompting records for selected taxa. The
records for Malus sylvestris sensu lato, M. sylvestris sensu stricto and M. domestica are highly
correlated with the taxa listed on the five regional record cards (Fig. 11). Similar effects were found
in Asplenium trichomanes, Juniperus communis and Veronica serpyllifolia. The commonest
segregate or infraspecific taxon of an aggregate or species may also be under-recorded simply
because it is the assumed taxon; there were only two records for Hedera helix var. helix for the
Monitoring Scheme compared to 34 for var. hibernica even though the former is the more common
taxon (McAllister & Rutherford 1990).

RECORDING BIAS IN BOTANICAL SURVEYS

Ficure 8. Examples of species which may have been overlooked for the Afélas: (a) Festuca altissima; (b) Chenopodium ficifolium.

86 T. C. G. RICH AND E. R. WOODRUFF

900

600

300

Cumulative number of 10-km
squares recorded

1800 1850 1900 1950

Date

FiGurE 9. Cumulative increase in the number of 10-km squares recorded for Cardaria draba with time.

BIASES RELATED TO THE TYPES OF PLANTS BEING RECORDED
Some biases in recording may depend on the type of plants being recorded in nian to the
behaviour of the botanists.

The apparency, or ease with which a plant is seen may dictate how consistently it is recorded.
Chamerion angustifolium has large, purple flowers at eye-level and is unlikely to be missed. Leersia
oryzoides is a large grass of ditches and watersides which rarely flowers; it is very difficult to see
amongst Phalaris arundinacea with which it usually grows and closely resembles vegetatively, and is
easily overlooked. Small, fine-leaved or widely dispersed taxa are often more poorly recorded than
large, broad-leaved or clumped species (Sykes et al. 1983; Clymo 1980). Plants abundant in an area
are more likely to be found than those less frequent, simply due to the higher probability of a
recorder finding them.

The seasonality of appearance of plants is well-known. In the genus Scilla, for example, S. verna is
most conspicuous in spring and early summer, whilst S$. autumnalis is seen mainly in late summer
and autumn. The seasonality of recording by botanists is also well-known, most activity taking place
during the summer. Species which are most conspicuous at the beginning or end of a season are
likely to be less consistently recorded than those most conspicuous in the middle.

Seasonal bias may arise from variations in the occurrence of the plant (e.g, spring annuals,
woodland herbs), variations in the apparency, or from difficulties in identifying particular taxa at
certain times of year (e.g. Nasturtium spp. are most reliably identified from ripe seeds which are
unavailable early in the season (Rich 1987)). Fig. 12 shows examples of seasonal bias based on
records collected during 1987 and 1988, and includes records of both flowering and vegetative
plants. Fig. 12a shows the relative numbers of records collected in each month; most records are
collected between May and September, reflecting seasonality of both plants and recorders.

RECORDING BIAS IN BOTANICAL SURVEYS

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T. C. G. RICH AND E. R. WOODRUFF

RECORDING BIAS IN BOTANICAL SURVEYS

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90 T. C. G. RICH AND E. R. WOODRUFF

TABLE 1. NUMBER OF 10-KM SQUARES RECORDED AND NUMBER OF RECORDS PER YEAR
FOR THREE SPECIES OF ARABLE WEED FOR 1930-1960 AND 1987-1988 BASED ON RECORDS
FROM THE B.S.B.I. MONITORING SCHEME SAMPLE SQUARES ONLY

Number of 10-km Number of

squares recorded records per year
Species 1930-60 1987-88 1930-60 1987-88
Agrostemma githago 29 i 0-94 3°5
Ranunculus arvensis 55 14 1-77 7
Scandix pecten-veneris 71 6 J3 3

Trifolium repens is a virtually ubiquitous species present all year, and its seasonal recording pattern
would therefore be expected to be similar to that of all records; Fig. 12b shows that it 1s.

Histograms for species showing seasonal variations in occurrence are shown in Figs 12c-e. Adoxa
moschatellina is a perennial herb of woodlands and waysides with a very short period of growth from
about March to June, after which it withers rapidly and disappears. Hyacinthoides non-scripta is also
primarily a plant of the spring, but the fruiting stalks persist and it is consequently recorded until late
summer. Spiranthes spiralis is a perennial herb of calcareous grasslands which flowers in late August
and September but whose leaves are usually absent during the summer (Wells 1967). In these cases
the seasonality of occurrence of the plants is matched by the records.

Two examples of changes in apparency are shown in Figs 12f & 12g. Arum maculatum is a
perennial herb of woodlands, hedgerows and waysides, etc. throughout Britain and Ireland. It is
conspicuous in spring when the leaves and inflorescences appear, but becomes less obvious in
summer when the leaves die back, the fruiting heads are small and green, and other vegetation
grows up around them. In August, the fruits begin to ripen and turn red, and the plants once again
become conspicuous. These changes in apparency are reflected by the bimodal nature of the
records; note that the plant is present in the summer but relatively under-recorded. The seasonality
of records of Viscum album, an evergreen parasitic herb usually of deciduous trees and shrubs, is not
quite as might be predicted. Records increase to May and then decrease, presumably related to the
appearance of leaves on the trees. A rise in records might be predicted again in October when leaves
are shed, but there is a surprising peak in August instead; the cause of this peak is not known. The
small increase at Christmas may not be coincidence.

Fig. 12h shows the seasonality of records of Salicornia europaea sensu stricto. Salicornia is a
critical genus (e.g. Rich & Rich 1988) whose species can only be reliably distinguished in autumn
when in fruit, and Fig. 12h reflects this. Salicornia records not determined to species show a much
broader spread of records as expected. Other similar examples of taxa which can only be identified
at certain times of year include Ruppia, Taraxacum and Hieracium.

Such seasonal biases may influence assessments of change with time and might be reflected in the
distribution maps. Fig. 13 shows 10-km squares which were not recorded before July or after June
for the Monitoring Scheme in 1987 and 1988. Autumn and spring species might be expected to be
under-recorded in these squares.

Perring & Walters (1962) noted that some species had died down before observers had arrived to
record them for the Atlas, and thus appear to be rarer on the maps than they actually are. This effect
was particularly marked in S.W. Ireland due to the early flowering season and the remoteness of the
south west from the main centres of botanical activity. This is shown by the Monitoring Scheme
results for one vernal species, Anemone nemorosa (Fig. 14a). The six new records for the
Monitoring Scheme in S.W. Ireland are a direct result of more work earlier in the season; five of
these new records are for squares visited early in the year (compare Fig. 14a with Fig. 13). No doubt
the species also occurs in some of the squares only visited later. Similar results are shown for other
vernal taxa such as Ranunculus ficaria. An increase in Chrysosplenium oppositifolium in Ireland
(Fig. 14b), another species most conspicuous early in the year but present all season, may also be
explained by this phenomenon.

RECORDING BIAS IN BOTANICAL SURVEYS

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T. C. G. RICH AND E. R. WOODRUFF

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jou soienbs (q) ‘Ajn¢ 910JaqQ pop1099I JOU sareNbs (Le) :gQ6] PUL LQG] Ul WOYS SULIOPUOJ] IY} JO} SP1O9I1 JY} 0} SeIQ JeUOSBIS B YIIM SoJENbs WY-0] “E] AINA

RECORDING BIAS IN BOTANICAL SURVEYS

‘wniyofyisoddo wniuajdsosksyD (q) ‘vsosowau auowauy (e)

i 2d

:SRIG [LUOSEIS 0} ONP purer] “A'S Ul popiodal-1opunN A]qeqoid exe ‘pl aNd

94 T. C. G. RICH AND E. R. WOODRUFF

DISCUSSION

It is clear from the examples above that recording bias is widespread in data collected for the Atlas
and for the Monitoring Scheme. Approximately one third of the taxa analysed for the Monitoring
Scheme were found to have unacceptable degrees of recording bias which could not be corrected.
More than one form of bias may also be present — all examples presented here include a bias of
greater recording effort (Rich & Woodruff 1990, 1992) for the Monitoring Scheme. Presumably
biases similar to those presented here occur in most other botanical surveys, though they are rarely
pointed out.

The extent of recording bias indicates that care should be taken with interpreting sets of records.
Assessing recording bias is difficult and requires intimate knowledge of the taxa concerned, how
they are recorded now and how they were recorded in the past, their habitats, general distribution
and frequencies, variations in the quality and quantity of recording, etc. Although each case has to
be judged on its merits, a few generalizations can be made:

1. Critical, infraspecific, hybrid and the more obscure taxa will generally show larger amounts of
recording bias related to individual recorders.
2. Aliens, casuals, garden escapes, forestry trees, crops and deliberately planted taxa may be less
consistently recorded than native species.
3. Areas briefly covered by few botanists will be less consistently recorded than areas well-covered
by many botanists.
. Localized areas may show considerable bias related to the activities of individual recorders.
. Some habitats with difficult or limited access (e.g. mountains, cliffs, water) will be relatively
poorly recorded. Others with easy access (e.g. car-parks, churchyards) may be well-recorded.
6. Large, obvious or clumped taxa will be more consistently recorded than small, inconspicuous or
widely dispersed ones. Abundant species will be more consistently recorded than rarer species
but national rarities tend to be well documented. :

7. Species characteristic of the beginning and end of the field season will be less consistently
recorded than those in mid season.

8. The number of records will primarily be dictated by recording effort put into collecting them.

9. Methods of survey may introduce systematic bias.

nN &

As records accumulate, the influence of recording bias will diminish, provided that adequate
quality control is exercised. Experience is required to judge whether the bias is large enough to
affect the interpretation of the records. It is easier to spot bias by comparing two similar surveys than
to assess it in isolation.

The widespread occurrence of recording bias suggests that information about recorders and their
behaviour should be collected and analysed as carefully as information about the organisms
themselves. Unrecognised bias in a sample of records will result in an incorrect interpretation of the
data. Those who collect, compile and present the data should therefore also provide interpretation
to guide those unfamiliar with the problems.

ACKNOWLEDGMENTS

The work would not have been possible without the 1600 volunteers, largely members of the
B.S.B.I., who contributed to the Monitoring Scheme in 1987 and 1988. Thanks are especially due to
the B.S.B.I. vice-county Recorders and other nominated botanists who helped to co-ordinate the
recording.

We are grateful to the staff at B.R.C., especially Chris Preston and Paul Harding, for help during
the Monitoring Scheme. Arthur Chater, Eimear Nic Lughadha and John Hellawell have provided
many useful comments on the manuscript.

This work was funded by the Nature Conservancy Council (nominated officer John Hellawell),
and the Department of the Environment, Northern Ireland.

RECORDING BIAS IN BOTANICAL SURVEYS 95
REFERENCES

CLAPHAM, A. R., TuTin, T. G. & Moore, D. M. (1987). Flora of the British Isles, 3rd ed. Cambridge.

Ciymo, R. S. (1980). Preliminary survey of the peat-bog Hummel Knowe Moss using various numerical
methods. Vegetatio 42: 129-148.

Dony, J. G. (1963). The expectation of plant records from prescribed areas. Watsonia 5: 377-385.

GrEIG-SMITH, P. (1964). Quantitative plant ecology, 2nd ed. London.

HA t, P. C. (1980). Sussex plant atlas. Brighton.

Hope-Simpson, J. F. (1940). On the errors in the ordinary use of subjective frequency estimates in grassland. J.
Ecol. 28: 193-209.

Jermy, A. C., CHATER, A. O. & Davin, R. W. (1982). Sedges of the British Isles, 2nd ed. B.S.B.I. Handbook No.
1. London.

Kent, D. H. & Lous.ey, J. E. (1981). Docks and Knotweeds of the British Isles. B.S.B.1. Handbook No. 3.
London.

Kirpy, K. J., Bines, T., Burn, A., MAcKintosu, J., PiTkIN, P. & Smiru, I. (1986). Seasonal and observer
differences in vascular plant records from British woodlands. J. Ecol. 74: 123-131.

MCcALLIsTER, H. A. & RUTHERFORD, A. (1990). Hedera helix L. and H. hibernica (Kirchner) Bean (Araliaceae)

_ in the British Isles. Watsonia 18: 7-15.

MEIKLE, R. D. (1984). Willows and Poplars of Great Britain and Ireland. B.S.B.1. Handbook No. 4. London.

Nitsson, S. G. & Nirsson, I. N. (1983). Are estimated species turnover rates on islands largely sampling errors?
Amer. Nat. 121: 595-597.

PaGcE, K. W. (1987). Hybrid Bluebells. B.S.B.I. News 47: 9.

PERRING, F. H. & WALTERS, S. M., eds. (1962). Atlas of the British flora. Cambridge.

Preston, C. D. & EversHaM, B. (1992). Recording bias and species mobility as factors affecting the
interpretation of distribution maps (In prep.).

Ricu, T. C. G. (1987). What-a-cress? or, How to do Nasturtium R.Br. B.S.B.I. News 46: 18-19.

Ricu, T. C. G. & Ricu, M. D. B., eds. (1988). Plant Crib. London.

Ricu, T. C. G. &Wooprurr, E. R. (1990). B.S.B.I. Monitoring Scheme 1987-1988. Unpublished report to the
Nature Conservancy Council.

Ricu, T. C. G. & Wooprurr, E. R. (1992). A sample survey of Britain and Ireland to assess trends in the flora
since 1962: The B.S.B.I. Monitoring Scheme 1987-1988. Peterborough. (In prep.)

ScuRFIELD, G. (1962). Biological flora of the British Isles. Cardaria draba (L.) Desv. J. Ecol. 50: 489-499.

Sykes, J. M., Horritt, A. D. & Mountrorp, M. D. (1983). Use of visual cover estimates as quantitative
estimators of some British woodland taxa. J. Ecol. 71: 437-450.

Trist, P. J. O. & SELL, P. D. (1988). Two subspecies of Molinia caerulea (L.) Moench present in the British
Isles. Watsonia 17: 153-157.

WELLS, T. C. E. (1967). Changes in a population of Spiranthes spiralis (L.) Chevall. at Knocking Hoe National
Nature Reserve, Bedfordshire 1962-1965. J. Ecol. 55: 83-99.

West, N. E. & Hatton, T. J. (1990). Relative influence of observer error and plot randomization on detection of
vegetation change. Coenoses 5: 45—49.

(Accepted November 1991)

Watsonia, 19, 97-105 (1992) 97

A method for predicting the probability of species occurrence
using data from systematic surveys

M. G. LE DUC, M. O. HILL and T. H. SPARKS

Institute of Terrestrial Ecology, Monks Wood Experimental Station, Abbots Ripton, Huntingdon,
Cambs., PE17 2LS

ABSTRACT

Presence data for species in 2-km squares, recorded systematically during the B.S.B.I. Monitoring Scheme,
were smoothed to derive probability response surfaces for Euonymus europaeus L., Hyacinthoides non-scripta
(L.) Chouard ex Rothm., Trientalis europaea L. and Veronica montana L. Logistic regression was used to
predict species frequencies from the response surfaces together with information on species occurrence in 10-km
squares. Predicted frequencies were compared with those reported in some recent county floras. Agreement was
generally good, but county differences in recording intensity were apparent.

INTRODUCTION

Accurate information on the spatial distribution of plants is now needed more than ever as human
impacts on the environment intensify. Agricultural expansion and intensification (Green 1989),
atmospheric pollutants (e.g. nitrogen compounds — see Tamm 1991) and climate change (Huntley et
al. 1989) — thought to be a consequence of the increasing release of ‘greenhouse’ gases — are all seen
to result in habitat change and species loss. Some gains are also to be expected as governments try to
reduce agricultural surpluses by extensification and habitat creation, for example planting new
woods on farms (Insley 1988). ;
Currently, and perhaps foreseeably, it is not possible to predict the presence or absence of a
species from a knowledge of environmental factors and autecological characteristics alone.
Prediction is dependent on good floristic survey data. Plant distribution maps with 10-km square
resolution (Perring & Walters 1976), and local floras with tetrad (2-km square) resolution, are
examples of such data for Britain. Dony (1963) described how floristic surveys can be used to predict
the numbers of species occurring in tetrads. Hill (1991) demonstrated a method for using
environmental data to estimate the probability of finding bird and plant species in 10-km squares.
He concluded that the quality of the estimates varied with habitat preference, and that those species
with strong edaphic requirements (e.g. Helianthemum nummularium (L.) Miller) were only poorly
predicted in a broad-scale analysis.
For many species, the frequency of occurrence in tetrads provides a better indication of local
-abundance than a map of distribution at the 10-km square scale. However, a complete survey of
vascular plants in Britain and Ireland at the tetrad scale would hardly be feasible, even if it were
desirable. Fortunately, a systematic survey of a selected subset of tetrads not only is feasible but was
_accomplished by the B.S.B.I. Monitoring Scheme (Rich & Woodruff 1990). Data from this survey
'can be used to estimate the probability of finding species in tetrads that were not surveyed, and
‘hence give an indication of local frequency.
_ The main purpose of this paper is to develop and compare methods for estimating such
| probabilities, using data from the Monitoring Scheme and other systematic surveys. In addition, we
: show how probability estimates can be used to generate species frequency maps at the national
scale.

| MATERIALS AND METHODS
|

The B.S.B.I. Monitoring Scheme (funded by the Nature Conservancy Council and the Department
of the Environment, Northern Ireland) was a survey carried out in 1987 and 1988 and administered

98 M. G. LE DUC, M. O. HILL AND T. H. SPARKS

TABLE 1. SAMPLING STATISTICS FOR THE B.S.B.I. MONITORING SCHEME
The British subset used in this work excludes data from Ireland and the Channel Islands.

Actual Number in
Sample number British
Sample units size surveyed subset
10-km squares 429 425 298
Tetrads (A, J & W only) 1114 1080 796
Mean number of tetrads per 10-km square 2-60 2:54 2-67

through B.S.B.I. News (see Ellis 1986; Rich 1986, 1987, 1988, 1989). For the survey, one in nine of
the 10-km squares were systematically selected from the British and Irish National Grids. Within
selected 10-km squares, presence records for plant species were recorded in each of three
systematically positioned tetrads (designated A, J and W). Some tetrads did not contain land, so
that, on average, slightly fewer than three tetrads were sampled per 10-km square (Table 1). The
Monitoring Scheme data are held by the Biological Records Centre (B.R.C.) at the Institute of
Terrestrial Ecology (I.T.E.), Monks Wood. They are in ORACLE database format on a VAX
computer cluster running under VMS (Rich & Woodruff 1990).

Records of species presence or absence in tetrads were smoothed to a response surface whose z-
axis value is the probability of finding that species in the local tetrad. Each smoothed value is a
weighted average of the neighbouring values, with weights specified by the bivariate Gaussian
function with a root-mean-square deviation 30 km (Fig. 1). This smoothing radius was chosen
because 30 km is the spacing of the Monitoring Scheme 10-km squares. A smaller radius would
result in a response surface that showed marked local variation, reflecting frequencies in individual
10-km squares.

The smoothed value is

where wy = exp(—(x,” + y,”)/r’), p; = estimated probability of finding the i" species in the target
tetrad, w, = weight assigned to the k" tetrad in the sample area, a, = value (1 or 0) specifying —
presence or absence of the i" species in the k" tetrad, r = smoothing radius (30-km), and x, and y,
are the easting and northing distances of the k"" tetrad from the target tetrad. A smoothing radius of
30 km ensures that 98% of the weight comes from within a 60 km radius. Note that the summation is
taken over tetrads surveyed for the Monitoring Scheme. A tetrad near the coast is given a smoothed
value by averaging over nearby tetrads inland. This average is taken over a smaller number of points
than for a non-coastal position, but is not otherwise affected by proximity to the sea.
Since presence and absence data are not normally distributed, the method of logistic regression
analysis (cf. Jongman et al. 1987) was used to estimate species frequency in 10-km squares. Each
Monitoring Scheme 10-km square was allocated a species frequency value which was calculated as
the ratio of the number of occupied tetrads to the number of recorded (maximum three) tetrads.
These values were regressed against the mean of the expected probabilities, estimated from the
response surface, averaging probabilities over all the tetrads (25 maximum) within that square. Two
models were considered: firstly, a model using only the spatially smoothed probability as
independent variable (Model 1 below); secondly, a model (Model 2) using the spatially smoothed
probability together with 10-km presence and absence data. For this purpose, 10-km data were
obtained from the records held by B.R.C. at I.T.E., Monks Wood. These data comprise validated
plant records from a variety of sources and were the records used to plot the Atlas of the British flora
(Perring & Walters 1976). |
The regression models, fitted by means of generalized linear modelling using the GENSTAT

computer package, were
loge (5) = a; + dip; Modell

PREDICTING THE PROBABILITY OF SPECIES OCCURRENCE 99

log. 2s] == al ate bip; =F c;B; Model 2

where q; = probability of finding the i species in a given tetrad of a Monitoring Scheme 10-km
square, p; = mean estimated probability of occurrence smoothed over the tetrads in the 10-km
square, B; = presence or absence (one or zero) of the i” species in the 10-km square, and a;, b; & ¢;
are constants.

The accuracy of the smoothed probability surface was further investigated using a validation set of
data from independent surveys obtained from a selection of those English county Floras meeting
three criteria. Firstly, publication had to be relatively recent; secondly, records had to be available
in atlas form for ease of data extraction; thirdly, mapping had to be at tetrad or 1-km square
resolution. Those selected were for Bedfordshire (Dony 1976), Devon (Ivimey-Cook 1984),
Durham (Graham 1988), north-east Essex (Tarpey & Heath 1990), Hertfordshire (Dony 1967),
Kent (Philp 1982), Leicestershire (Primavesi & Evans 1988) and Sussex (Hall 1980). None of the
available atlases from Wales or Scotland met the criteria (McCosh 1988). Only those 10-km squares
falling wholly within the county (or vice-county) boundaries were considered. For each species and
each 10-km square a table of presences out of the number of tetrads per 10-km square (25) was
produced. For the north-east Essex Flora the published data are for 1-km squares and were
summarized for each tetrad prior to processing.

Data from the county atlases were compared with both point estimates from simple Gaussian
smoothing and predicted values from each of the logistic regression models. The basis for the
comparison was the average number of presences in tetrads per 10-km square, county by county.
Analysis of variance was used to test the significance of differences. Accuracy of predictions was
measured by the root-mean-square difference between predicted and observed values.

To illustrate the technique we have selected four species, namely Euonymus europaeus L.,
Hyacinthoides non-scripta (L.) Chouard ex Rothm., Trientalis europaea L. and Veronica montana L.
E. europaeus is a southern species of calcareous soils. T. europaea is a boreal species having a
requirement for cooler northern winters. The other two species are generally distributed in older
woodlands, but H. non-scripta is much the commoner of the two. Tetrad presences and absences
(obtained from the B.S.B.I. Monitoring Scheme database) for each species have been plotted in
Fig. 2. Version 6 of the UNIRAS computer package (I.U.C.C. Information Services Group 1989)
was used for this and subsequent distribution maps and figures. Orkney and Shetland have been
omitted. For them, as for the Isle of Man (which was included, but which had only three tetrads), a
larger smoothing radius than 30 km might be desirable.

RESULTS

The response surfaces obtained by Gaussian smoothing are illustrated in Fig. 3. Regression
coefficients and significance levels for Models 1 and 2 are shown in Table 2. Highly significant results
can be expected because the independent regression variables were derived from the observed
values (dependent variables) by smoothing. Both Models 1 and 2 contain the derived variable p;.

The comparisons between county atlas records and the estimated values from Gaussian-
smoothed and regression models are shown in Table 3. The Gaussian-smoothed values were
obtained by summing p; for each tetrad in the 10-km square; predicted values from Models 1 and 2
were obtained by inserting appropriate p; values into the regression equations to obtain values of qj.
Although many of the estimated values were close to those expected from the county Floras there
were some substantial differences (Table 3).

The mean deviation (bias) was smallest for the prediction method using Model 2, but the bias of
all three methods was small and not statistically significant (Table 4). The root-mean-square error
for Model 2 was less than for Model 1 and approached that of the Gaussian-smoothed probabilities.
The analysis of variance shows no effect due to species but a highly significant county effect. The
bulk of the county effect can be attributed to underestimation by the three methods of tetrad
frequencies in Kent and possible over-estimation in Bedfordshire.

100 M. G. LE DUC, M. O. HILL AND T. H. SPARKS

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FicurE 1. Gaussian smoothing of occurrence in tetrads (2-km squares). At any point in Britain the probability of
a species being found in that tetrad is estimated as a weighted mean local frequency. Weights are defined by a
Gaussian function with root-mean-square deviation 30 km. The diagram shows the weight function projected on
to a 10-km square grid with the A, J and W tetrads for the one-in-nine sample indicated.

PREDICTING THE PROBABILITY OF SPECIES OCCURRENCE 101

TABLE 2. LOGISTIC REGRESSION OF OBSERVED AGAINST PREDICTED FREQUENCIES IN
10-KM SQUARES OF THE B.S.B.I. MONITORING SCHEME
Coefficients a;, b; and c; are defined in the text for Models 1 and 2. Degrees of freedom were (1, 296) for Model 1
and (2, 295) for Model 2.

Deviance
Species a; b; C explained (%) Significance
Model 1
Euonymus europaeus —4-27 8-50 i 74-3 p<0-001
Hyacinthoides non-scripta —3-86 7-40 oe 69-2 p<0-001
Trientalis europaea = S25) Meza — 83-3 p<0-001
Veronica montana —o]) 9-29 — 62-7 p<0-001
Model 2
Euonymus europaeus —)-35 6°35 7:34 80-8 p<0-001
Hyacinthoides non-scripta —10-28 6-86 6-80 71-9 p<0-001
Trientalis europaea — 10-20 7-68 6-73 87-9 p<0-001
Veronica montana — 10-81 7-65 7-67 Ta p<0-001

TABLE 3. OBSERVED (1) AND PREDICTED (2-4) NUMBERS OF TETRADS OCCUPIED BY
SPECIES PER 10-KM SQUARE IN SELECTED COUNTIES

1 — average number of tetrads occupied according to the county atlases; 2 — expected value using the Gaussian-

smoothed Monitoring Scheme data; 3 and 4 — expected values using regression models 1 and 2 respectively.

Beds. Devon Durham Essex Herts. Kent Leics. Sussex Total

n 5 49 1) 1) 5 22 10 24 137
Euonymus europaeus
1 11-4 9-8 0-0 13-0 15:2 19-7 0-4 15-4 11-0
2 15-7 14-1 0-0 15-1 19-6 8-0 0-5 11-4 10-4
3 18-4 15-4 0-4 17:5 22:9 4:8 0-4 11-3 10-7
4 18-0 11-4 0-3 13-0 21-9 7-0 0-7 11-3 9-4
Hyacinthoides non-scripta
1 14-8 20-1 13-1 US)eo) 20-2 23:2 14-4 23°8 19-6
2 20-1 18-8 17-0 19-4 22:8 132 8:3 18-4 17-1
3 21-8 20-4 18-8 21-6 23:6 12:8 5-8 20-4 18-2
4 21:8 20-4 16-7 21-4 23-4 13-2 57) 20-4 18-0
Trientalis europaea
1 0 0 0 0 0 0 0 0 0
2 0 0 0-2 0 0 0 0 0 0
3 0-2 0-1 0-1 0-1 0-2 0-1 0-1 0-1 0-1
4 0 0 0-1 0 0 0 0 0 0
Veronica montana

1 0-8 12-0 6-9 5-0 11-0 14-2 3-6 16-0 11-1
2 4-] 13-2 11-4 G2 10-0 6-1 3-1 10:5 9-9
3 2-4 17-0 14-5 6-0 11-5 5-9 1-5 12-4 11-7
4 2-7 15-2 11-8 6-7 12-1 6-7 1-4 11-9 10-9

n = number of 10-km squares.
DISCUSSION

Smoothed distribution maps (Fig. 3) demonstrate the potential of the Monitoring Scheme data for
depicting probabilities of occurrence in tetrads. Similar smoothed maps could be used in future to
compare survey and re-survey results given a common survey protocol.

The ability of all the methods, including simple Gaussian smoothing and regression, to predict the

102 M. G. LE DUC, M. O. HILL AND T. H. SPARKS

Hyacinthoides non-scripta

Trientalis europaea Veronica montana

FiGure 2. Species occurrence in tetrads (2-km squares) recorded during the B.S.B.I. Monitoring Scheme.

PREDICTING THE PROBABILITY OF SPECIES OCCURRENCE 103

Euonymus europaeus Hyacinthoides non-scripta

MMB asove 08

ERC‘ - o8
0.4- 06

0.2- 0.4

[_]BELOW 02
Location probability

Trientalis europaea Veronica montana

_ Ficure 3. Probabilities of species occurrence in tetrads (2-km squares), estimated by smoothing the data in Fig. 2
with a Gaussian function.

104 M. G. LE DUC, M. O. HILL AND T. H. SPARKS

TABLE 4. ANALYSIS OF THE DIFFERENCES BETWEEN TETRAD FREQUENCIES FOR 10-KM

SQUARES, COMPARING FREQUENCIES OF EUONYMUS EUROPAEUS, HYACINTHOIDES NON-

SCRIPTA AND VERONICA MONTANA PREDICTED FROM THE MONITORING SCHEME WITH
THOSE OBSERVED IN COUNTY FLORAS

Models | and 2 are defined in the text. Method effect refers to tests of the null hypothesis that the mean deviation

iS zero.
Mean Method Species County Kent vs
Observed mean deviation effect effect effect others
county density (bias) RMSE tia Fy 14 Fo 14 Fi 14
Gaussian smoothed —0-50 4-91 —1-05 ns 0-25 ns USE SS 56:0a tee
Regression Model 1 0-32 6-10 0-46 ns 0-10 ns S200 > 39-eae
Regression Model 2 0-19 5-34 —0-36 ns 0-00 ns BOF Ee ea Ores

RMSE = root-mean-square error; ns = not significant; *** = p<0-001.

B.S.B.I. Monitoring Scheme data was generally quite good. The mean deviation (bias) was smallest
for the prediction method using Model 2, the root-mean-square error indicating its advantage over
Model 1. However the error was least for simple Gaussian smoothing.

There was a notable and statistically very significant difference between counties (Table 4). In
terms of effort per tetrad, Kent was more intensively surveyed for the county Flora than for the
Monitoring Scheme, whilst Bedfordshire was less so. In any survey the uniformity of sampling effort
is of great importance. The B.S.B.I. Monitoring Scheme was carefully controlled with this objective
(Rich & Woodruff 1990), but differences must inevitably have occurred. Variation also exists
between the county Floras, some being over-sampled in comparison with the Monitoring Scheme,
whilst others were relatively under-sampled.

For validation we have selected county Floras with a high and fairly uniform sampling coverage.
Even though the per-tetrad effort may sometimes have been less than that achieved by the
Monitoring Scheme, overall they will all have had more intensive sampling. Thus the resolution of
the response surfaces produced from the Monitoring Scheme will be poorer than those which could’
be obtained from the county Floras. In general we would expect those species with a fairly general
but patchy distribution, such as those requiring habitats in old woods, to be less easy to predict than
those species with distributions depending on some more widespread factor of the physical
environment such as climate or soil type. This seems to be the case when comparing the deviances
explained for E. europaeus and T. europaea on the one hand, with H. non-scripta and V. montana —
on the other (Table 2). It is also supported by the closer agreement between overall county atlas
data and the Gaussian-smoothed response surface (rows 1 and 2 in Table 3) for E. europaeus than
for H. non-scripta and V. montana.

The ability to predict species presence or absence using regression methods also seems to be
somewhat species-specific (Table 2). Those whose distribution is strongly restricted by specific
environmental factors such as climate (FE. europaeus and T. europaea) are seen to be better
predicted than the others. Predictions were substantially improved by including information on 10-
km square occurrence (Model 2). It is interesting that the coefficients a;, b;, c; in Model 2 were so
close in value that a single regression would have sufficed for all four species.

One of the main advantages of the logistic regression approach is that it can readily be extended to
include other information (Le Duc et al. 1992). Such information might include, for instance, soil
type (Avery 1973) and local climate (Bendelow & Hartnup 1980). Perhaps more important for many
widespread species would be inclusion of additional habitat information such as the presence of
woods, rivers, or a coastline. Such information is now becoming available in, for instance, the I.T.E.
land classification database (Bunce et al. 1981). The more accurately the present frequency of a
species can be estimated the better we shall be able to detect change in the future.

CONCLUSIONS

In Great Britain, sufficiently good survey data are now available to derive reliable national
estimates of the probability of species occurrence in tetrads. Such estimates can be validated using

|
;
)
|
|
|

{

PREDICTING THE PROBABILITY OF SPECIES OCCURRENCE 105

independent data from county Floras. Using Gaussian-smoothed data from the Monitoring
Scheme, combined with additional information about each tetrad, regression models can be
developed which would improve the accuracy of estimates. These estimates can be used in future to
detect the effects of major disturbances such as climate change or large-scale shifts in land use.

ACKNOWLEDGMENTS

This work was partly supported by a grant from the Forestry Research Co-ordination Committee.
We thank Chris Preston for his assistance in selecting the county atlases, and Tim Rich for
suggestions on use of the Monitoring Scheme database.

REFERENCES

Avery, B. W. (1973). Soil classification in the Soil Survey of England and Wales. J. Soil Sci. 24: 324-338.

BENDELOw, V. C. & Hartnup, R. (1980). Climatic classification of England and Wales. Soil Survey Technical
Monograph No. 15. Harpenden.

Bunce, R. G. H., Barr, C. J. & Wuirtaker, H. A. (1981). Land classes in Great Britain: Preliminary
descriptions for users of the Merlewood method of land classification. Merlewood Research and Develop-
ment Paper No. 86. Grange-over-Sands.

Dony, J. G. (1963). The expectation of plant records from prescribed areas. Watsonia 5: 377-385.

Dony, J. G. (1967). Flora of Hertfordshire. Hitchin.

Dony, J. G. (1976). Bedfordshire plant atlas. Luton.

ELuis, G. (1986). The new mapping scheme. B.S.B.I. News 43: 7-9.

GRAHAM, G. G. (1988). The flora and vegetation of County Durham. Durham.

GREEN, B. H. (1989). Agricultural impacts on the rural environment. J. Appld Ecol. 26: 793-802.

Hatt, P. C. (1980). Sussex plant atlas. Brighton.

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HunrtLeEY, B., BARTLEIN, P. J. & PRenticE, I. C. (1989). Climatic control of the distribution and abundance of
beech (Fagus L.) in Europe and North America. J. Biogeog. 16: 551-560.

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(Accepted February 1992)

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bettas

Watsonia, 19, 107-112 (1992) 107

The distinction between the Festuca ovina L. and Festuca
rubra L. aggregates in the British Isles

C. A. STACE, A.-K. K. A. AL-BERMANI and M. J. WILKINSON*

Department of Botany, University of Leicester, Leicester, LE] 7RH

ABSTRACT

The Festuca rubra L. and F. ovina L. aggregates (Poaceae) are defined and reliable means of distinguishing them
in the British Isles are given. The best character concerns the young tiller leaf-sheaths, which are tubular in the
former and split and overlapping in the latter aggregate. Characters that are often used to separate the two
aggregates but are unreliable and therefore frequently cause misidentification are also listed.

INTRODUCTION

Festuca ovina L. and F. rubra L. fall into section Ovince Fries (= sect. Festuca) according to
Hackel’s (1882) classification of the genus in Europe. Apart from these two species, which were
each subdivided into a complex hierarchy of subspecies, varieties and subvarieties, Hackel included
only nine other species, none of them British. He divided the section into two groups:
Intravaginales, including F. ovina and eight other species; and Extravaginales vel Mixtae, with F.
rubra and F. porcii Hackel.

Markgraf-Dannenberg (1980) abandoned Hackel’s infrageneric classification, placing the 170
European species that she recognised into a number of intormal, unnamed groups. However, it is
clear that 129 of these species would have been included in section Ovinae by Hackel. The increase
from eleven to 129 over a period of 100 years is partly due to the description of newly discovered
taxa, and partly due to a much narrower species concept that gradually developed.

All the British taxa now recognised were placed by Hackel (1882) into either F. ovina or F. rubra,
which are for convenience here referred to as the F. ovina and F. rubra aggregates. These taxa are
listed under the names used by Hackel, Markgraf-Dannenberg and us (Wilkinson & Stace 1991, Al-
Bermani 1991) in Table 1.

Both the aggregates are very variable and also very important from both ecological and economic
points of view. It is vital that plants of such importance are identified correctly, so it is particularly
unfortunate that not only have the segregates of both aggregates been very widely misunderstood,
but the two aggregates have been and are still frequentiy confused, particularly by British botanists.
Virtually all the current British Floras, identification manuals and flower-guides that we have
examined contain errors that would prevent accurate determination of at least some plants.
Although most taxa of F. ovina agg. are easily separated from most taxa of F. rubra agg., certain
taxa are not so easily placed. For example, within F. ovina agg., F. lemanii Bast. and F. brevipila
Tracey are usually much more robust than, for exampie, F. ovina and are often misidentified as F.
rubra. Conversely, F. rubra subsp. commutata Gaudin usually lacks rhizomes and is sometimes
identified as a robust variant of F. ovina agg. In addition, pseudoviviparous variants of F. rubra are
quite frequent and usually misdetermined as F. vivipara (L.) Sm., which belongs to the F. ovina agg.
Although characters such as spikelet, lemma or awn lengths, leaf thickness and flatness and degree
of adaxial ridging, culm height, degree of tuftedness and presence of rhizomes are important
diagnostically in the section as a whole, they are of very little value on their own in distinguishing
between the two aggregates, being in fact the most usual causes of confusion. Moreover, habitat
preferences are of very limited value in separating these aggregates. For accurate determination it is

*Present address: Scottish Crop Research Institute, Invergowrie, Dundee, DD2 5DA.

C. A. STACE, A.-K. K. A. AL-BERMANI AND M. J. WILKINSON

108

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DISTINCTION BETWEEN FESTUCA OVINA L. AND F. RUBRA L. 109

important that all these misleading characters be abandoned and that attention be focused on the
characters described in the next section.

The purpose of this paper is to provide infallible means of distinguishing between the F. ovina and
F. rubra aggregates in the British Isles. Distinctions between the various segregates as listed in Table
1 have been detailed by Wilkinson & Stace (1991) for the F. ovina aggregate and will be detailed by
Al-Bermani & Stace (in prep.) for the F. rubra aggregate. The data presented in the present paper
have been gathered from many thousands of specimens, both living and preserved, over the past 20
years. Several hundred clones are grown and frequently studied in the University of Leicester
Botanic Garden.

The characters discussed here would require some modification if they were to be used
successfully in some other parts of Europe.

DIAGNOSTIC CHARACTERS

The first two characters below provide an infallible distinction between the two aggregates; the
others may be useful guides but should not be relied upon.

TILLER LEAF-SHEATHS — FUSED OR OVERLAPPING

In almost all grass leaves there is a clear distinction between the sheath, which encircles the stem
and/or developing leaves above, and the blade, which arises from the top of the sheath. In some
cases this overlapping structure persists to the base of the sheath, i.e. to the stem-node below, but
often it progresses only part of the way down to the node, becoming a fused tube further down. In
other cases the sheath has no free overlapping edges, but is a fused tube from the node below right
up to the mouth of the sheath. This is particularly characteristic of the genera Glyceria and Melica.

In the Festuca rubra aggregate the leaf-sheaths are tubular right up to or almost to the mouth,
whereas in the Festuca ovina aggregate they have free overlapping edges for at least the upper 40%
(and usually the upper 75%) of their length (Fig. 1). Only in very rare cases in the latter aggregate
are the leaf-sheaths as much as 50% tubular.

This difference is very clear-cut, but careful examination (preferably with a lens) must be made in
order to avoid mistakes. Fresh material is much more easily observed than dried or pressed
material. Moreover the tubular sheaths of F. rubra agg. are easily split on handling. It is essential
that this observation is made on sterile innovation shoots (tillers), not on flowering shoots (culms),
and that only the younger sheaths are examined. Older sheaths, including all those on the culms,
become naturally split to the base. The older leaves should be successively stripped back from a
tiller, leaving the uppermost sheath that was partly exposed as the one to examine.

This is the single best character to distinguish the two aggregates.

TILLERS — EXTRAVAGINAL OR INTRAVAGINAL
Tillers arise from lower nodes of culms or of other tillers (actually from the axil between the stem
and the base of the leaf-sheath) in one of two ways.

In intravaginal branching (Fig. 1) the new shoot grows up more or less parallel with its parent stem
and remains enclosed for some distance within the leaf-sheath in whose axil it arose. Higher up, the
new shoot diverges from its parent, and later on the parental leaf-sheath often decays away, but the
parallel growth of the old and new stems at the very base usually persists as evidence of intravaginal
branching. In the Festuca ovina aggregate all branching is of this sort.

In extravaginal branching (Fig. 1) the new shoot does not grow up parallel with the parent stem
but grows out more or less at right angles, breaking through the base of the parent leaf-sheath (cf.
lateral branches of Equisetum stems). Members of the Festuca rubra aggregate always exhibit
extravaginal branching, but this varies in relative frequency from being the only type of branching to
being much less common than intravaginal branching (hence Hackel’s group Extravaginales vel
Mixtae). With copious fresh material the presence of extravaginal branches is therefore diagnostic
of F. rubra agg. (and their absence diagnostic of F. ovina agg.), but with herbarium material, which
is often very poorly collected, it is not safe to use this character except in a positive way.

110 C. A. STACE, A.-K. K. A. AL-BERMANI AND M. J. WILKINSON

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FicurE 1. Morphological characters of Festuca rubra agg. and F. ovina agg. A, extravaginal tillers. B,
intravaginal tillers. C, fused tiller leaf-sheath as in F. rubra agg. D, overlapping tiller leaf-sheath as in F. ovina
agg. E, vestigial auricles as in F. rubra agg. F, distinct auricles as in F. ovina agg.

DISTINCTION BETWEEN FESTUCA OVINA L. AND FESTUCA RUBRA L. 111

Ficure 2. Patterns of sclerenchyma distribution (in black) in tiller leaf sections of Festuca rubra agg. (A—G) and
F. ovina agg. (H-L).

GROWTH HABIT — TUFTED OR RHIZOMATOUS

Rhizomes always develop from extravaginal branches, but both intravaginal and extravaginal
branches may remain close to the parent stem, resulting in a densely tufted habit. Hence the
presence of rhizomes is diagnostic of F. rubra agg., but their absence is not diagnostic of F. ovina
agg. Some rhizomes exist even on plants of F. rubra agg. that are very densely tufted, except in F.
heterophylla Lam. and some specimens of F. rubra subsp. commutata. Mat-forming plants are
almost always rhizomatous to some degree.

112 C. A. STACE, A.-K. K. A. AL-BERMANI AND M. J. WILKINSON

AURICLES — CONSPICUOUS OR INCONSPICUOUS

Auricles are developed in many grasses at the top of the leaf-sheath, on either side of its point of
junction with the leaf-blade and ligule. In some species, such as Festuca pratensis Hudson, these are
very conspicuous and developed as pointed projections. In the F. rubra aggregate auricles are
virtually absent (Fig. 1), but in the F. ovina aggregate they are present as distinct rounded
extensions (Fig. 1). This difference is clear-cut and unequivocal when the two types are viewed
together but it is a comparative rather than absolute character and should be used only after
experience with the ranges exhibited by both aggregates.

LEAF ANATOMY — PATTERN OF SCLERENCHYMA
The pattern of sclerenchyma distribution as seen in transverse section of tiller leaves is a very
important character in the genus Festuca. Knowledge of the range of patterns found in the F. rubra
and F. ovina aggregates (Fig. 2) can be used to distinguish between them.

Sclerenchyma bundles are usually found in the subepidermal position opposite each of the veins
(including the midrib) on the abaxial side, and in the subepidermal position at the leaf margins (e.g.
Fig. 2B, G, J). Sometimes smaller subepidermal bundles are found opposite the veins on the adaxial
side (e.g. Fig. 2A, E, F), and sometimes a girder of sclerenchyma connects the abaxial bundle with
its adjacent vein (e.g. Fig. 2E, F). In some cases the abaxial sclerenchyma bundles extend laterally
to form a band of sclerenchyma, in extreme cases forming a continuous zone from leaf-margin to
leaf-margin (e.g. Fig. 2F, H).

Both the continuous and the discrete patterns of sclerenchyma occur in both the F. rubra and F.
ovina aggregates, but in the case of each pattern the two aggregates can, with practice, be
distinguished. The continuous/discontinuous band is almost always accompanied by some girders
connecting with the veins and/or by some small adaxial sclerenchyma bundles in the case of F. rubra
agg., but never in the case of F. ovina agg. In addition the sclerenchyma at the leaf-margins and
abaxially to the midrib is often much thicker than elsewhere in F. ovina agg. (e.g. Fig. 2J, K), but
not in F. rubra agg. The discrete abaxial sclerenchyma bundles are usually of approximately equal
size and are sometimes accompanied by small adaxial bundles in F. rubra agg. (e.g. Fig. 2B),
whereas in F. ovina agg. adaxial bundles are always absent and the marginal and midrib abaxial
bundles are usually conspicuously larger (e.g. Fig. 2J) than the others (or there are no others, e.g.
Fig. 2L) in F. ovina agg. In addition, when the sclerenchyma is in discrete bundles, the leaf outline is
much more angular in F. rubra agg. (e.g. Fig. 2B, C, G) than in F. ovina agg. (e.g. Fig. 2J, L).

SUMMARY

Only two characters can be used with complete success in distinguishing the Festuca ovina and F.
rubra aggregates in the British Isles. Other characters vary in usefulness from being helpful after
considerable experience to being highly misleading. The two diagnostic characters are summarized
in the following couplet:

Sheaths of young tiller-leaves fused into tube almost up to top; some or all tillers extravaginal ...

F. rubra agg.
Sheaths of young tiller-leaves with at least the upper 40% with free, overlapping margins; all tillers
INUTEAV AGIAN: 6 hic ian co ea sje sera ate cals ere via Gi cancers Slstatia ec mn Ge ate sco te cine earn F. ovina agg.

REFERENCES

AL-BERMANI, A.-K. K. A. (1991). Taxonomic, cytogenetic and breeding relationships of Festuca rubra sensu
lato. Ph.D. Thesis, University of Leicester.

HaAckEL, E. (1882). Monographia Festucarum Europaearum. Kassel & Berlin.

HAcKEL, E. (1885). Festuca ‘“‘glauca’’, in BENNETT, A. Report of the distributor for 1884. Rep. botl Exch. Club
Br. Isles 1: 101-119.

MARKGRAF-DANNENBERG, I. (1980). Festuca L., in Tutin, T. G. et al., eds. Flora Europaea 5: 125-153.
Cambridge.

WILKINSON, M. J. & Stace, C. A. (1991). A new taxonomic treatment of the Festuca ovina L. aggregate
(Poaceae) in the British Isles. Bot. J. Linn. Soc. 106: 347-397.

(Accepted January 1992)

Watsonia, 19, 113-120 (1992) 118

Host range and specificity of Orobanche minor Sm. on
Crymlyn Burrows

C. R. HIPKIN

School of Biological Sciences, University of Wales, Swansea, SA2 8PP

ABSTRACT

A survey of hosts reported for the broomrape Orobanche minor Sm. (Orobanchaceae) in Britain and Ireland,
indicates a very wide range for this species with most hosts recorded from the Leguminosae and the Compositae.
Results from an investigation of the host range of a large coastal population of O. minor Sm. var. minor (sensu
Rumsey & Jury) on Crymlyn Burrows, West Glamorgan, confirm 15 host species by observation of direct root
contact between host and parasite. Some individual broomrapes were noted to parasitise more than one (up to
three) host, each a different species. Seeds of O. minor var. minor taken from plants parasitising Eryngium
maritimum germinated to produce plants that were able to parasitise Hypochoeris radicata, Trifolium hybridum
and T. pratense.

INTRODUCTION

Of the 45 species of Orobanche L. recognised in Europe by Webb & Chater (1972), seven were
assigned to the Orobanche minor group and two of these, O. minor Sm. and O. loricata Reichenb.,
are in the British flora (Clapham et al. 1987). Both these species have predominantly south-eastern
distributions in Britain, with O. minor extending discontinuously west and north-west, more or less
along the coast. Taxonomically, the O. minor group is rather problematic, a situation which is
exacerbated by the great intraspecific variation exhibited by O. minor itself. In a recent account of
Orobanche L. in Britain and Ireland, Rumsey & Jury (1991) divide O. minor into four varieties,
(var. minor, compositarum, flava and maritima), and describe a number of features which aid their
identification including a guide to their host ranges.

Correct identification of the host can be helpful in the identification of several species of
Orobanche. Unfortunately, however, although host specificity within the O. minor group may be of
some significance taxonomically, it is confusing for at least two reasons. Firstly, some taxa appear to
have rather wide, overlapping host ranges. Secondly, it is usually difficult to identify the host
unequivocally in the field (Chater 1986) and it is likely that several hosts reported in the literature
have been recorded in error. Nevertheless, some taxa may parasitise a well defined, narrow
spectrum of hosts, e.g. O. minor var. maritima (Rumsey & Jury 1991), and the concept of
physiological races with very specific host requirements (Musselman & Parker 1982) has often been
suggested. Comments in county Floras indicate that coastal populations have frequently caused

_ difficulties and this appears to be, in part, a consequence of an incomplete understanding of the host

vem Se
ee

ranges of O. minor and O. maritima Pugsley (Pugsley 1940) in particular. Recent exchanges in the
literature (e.g. Hambler 1991; Rumsey 1991) underline the controversial nature of this problem.

. In this account, the host range of a large population of O. minor growing in mobile and partially
fixed sand dune communities on Crymlyn Burrows in West Glamorgan (v.c. 41) is described. These
studies exemplify the problems associated with coastal populations of O. minor and confirm a
number of hosts, unequivocally.

MATERIALS AND METHODS

A large sand dune population of Orobanche minor was studied on Crymlyn Burrows (GR SS/

_ 710.930). In recent years the size of this broomrape population has usually numbered more than

114

C. R. HIPKIN

TABLE 1. HOSTS FOR OROBANCHE MINOR RECORDED IN BRITAIN AND IRELAND

Host

CARYOPHYLLACEAE
Honkenya peploides

GERANIACEAE
Geranium columbinum
G. rotundifolium
Erodium cicutarium
Pelargonium spp.

LEGUMINOSAE
Clianthus puniceus
Vicia faba
Ononis repens

Medicago arabica
M. lupulina

Trifolium arvense
T. campestre
T. hybridum

T. medium
T. pratense

T. repens

Lotus corniculatus
Onobrychis viciifolia
ARALIACEAE
Hedera hibernica
UMBELLIFERAE
Eryngium maritimum

Anthriscus sylvestris
Pastinaca sativa

Reference

Rumsey & Jury (1991)

White (1912)

Petch & Swann (1968)
Lousley (1976)
Murphy (1972)

Jones (1985)

Margetts & David (1981)

Jones (1985)

Savidge et al. (1963)

White (1912)

Jones (1985)

Rumsey & Jury (1991)

McClintock (1975)

Bowen (1968)

Jones (1985)

Trimen & Thiselton-Dyer
(1869)

McClintock (1975)

Jones (1985)

Grose (1957)

Jones (1985)

Savidge et al. (1963)

Ravenshaw (1860)

Salmon (1863)

Archer Briggs (1880)

Grose (1957)

Savidge et al. (1963)

Bowen (1968)

Messenger (1971)

McClintock (1975)

Kent (1975)

White (1912)

Murray (1896)

Grose (1957)

Jones (1985)

Wolley-Dod (1970)*
White (1912)*
McClintock (1975)**
Bevis et al. (1978)**
Davey (1909)*
Margetts & David (1981)**
Allen (1984)

Jones (1985)

Rumsey & Jury (1991)
Grose (1957)

Grose (1957)

Holland et al. (1986)
Jones (1985)

Host

Daucus carota
(including subsp.
gummifer)

Garden spp.

(e.g. carrot, caraway)
BUDDLEJACEAE

Buddleja davidii
POLEMONIACEAE

Polemonium caeruleum
CONVOLVULACEAE

Calystegia soldanella
SOLANACEAE

Nicotiana cultivar

SCROPHULARIACEAE

Parahebe catarrhactae
LABIATAE

Salvia verbenaca’

Glechoma hederacea

PLANTAGINACEAE
Plantago coronopus

P. major
P. maritima
CAMPANULACEAE
Campanula
portenschlagiana
C. poscharskyana
DIPSACACEAE
Dipsacus fullonum
COMPOSITAE
Brachyglottis
cv. ‘Sunshine’

Olearia macrodonta
Achillea millefolium
Tripleurospermum
inodorum
Matricaria spp.
Carduus nutans
Cirsium arvense
C. vulgare

Reference

Ravenshaw (1860)*
Grose (1957)

White (1912)*
McClintock (1975)*
Davey (1909)*

Margetts & David (1981)§
Holland et al. (1986)
Jones (1985)

Lousley (1976)

Jones (1985)

Patrick & Hollick (1975)
Rumsey & Jury (1991)
Jones (1985)

Margetts & David (1981)

White (1912)
Murray (1896)**
Grose (1957)
White (1912)
Jones (1985)

Ravenshaw (1860)
McClintock (1975)
Davey (1909)*
Jones (1985)

Grose (1957)
McClintock (1975)§

Murphy (1972)
Jones (1985)

Jones (1985)

Bowen (1968)

Swann (1975)*

Lousley (1976)

Margetts & David (1981)
Jones (1985)

Rumsey & Jury (1991)*
Jones (1985)

Grose (1957)

Rumsey & Jury (1991)*
Kent (1975)

Rumsey & Jury (1991)*
Grose (1957)

Jones (1985)

HOST RANGE AND SPECIFICITY OF OROBANCHE MINOR SM. 115

TABLE 1. cont.

Host Reference Host Reference
COMPOSITAE cont'd
Hypochoeris radicata Bowen (1968)* Petch & Swann (1968)*
Petch & Swann (1968)* Jones (1985)
McClintock (1975) Holland et al. (1986)
Jones (1985) Rumsey & Jury (1991)*
Rumsey & Jury (1991)* ~—C. vesicaria Dony (1953)
Leontodon autumnalis McClintock (1975)
Picris echioides Dony (1953) GRAMINEAE
Crepis capillaris Linton (1919) Elymus farctus Rumsey & Jury (1991)
Grose (1957) Ammophila arenaria Murray (1896)
Grose (1957)* Murphy (1972)

* = recorded as var. compositarum; * = recorded as O. amethystea; ** = recorded as O. amethystea but queried
by author(s) as O. minor; * = recorded as O. maritima; ** = recorded as O. maritima but queried by author(s) as
O. minor.

5,000 individuals. This presented opportunities to sample a relatively small number of specimens
with minimal disturbance to a thriving population. Plants rooted in sand were excavated carefully
(usually with ease and minimal disturbance to the rooting systems of host and parasite) and
examined for host-parasite root connections. Hosts were traced via their root systems and
identified. Often, it was possible to replace plants with little disturbance after substantiating host
root contact.

Plants were also cultivated in containers using as hosts Trifolium hybridum (cv. ‘Alsyke’, Elsoms
Seeds Ltd, Spalding, England), T. pratense and Hypochoeris radicata (both grown from local
stock). Seeds of O. minor were collected from individuals parasitising Eryngium maritimum on
Crymlyn Burrows. Host plants were established in pots before inoculation with broomrape seeds
which were applied in suspension directly on to the host roots.

REPORTED HOST RANGE OF OROBANCHE MINOR SM.

A cursory glance at the literature would reveal that O. minor has been recorded most frequently as a
parasite of Trifolium spp., often as a weed in clover fields. It became well known, however, that
other (often exotic) species were also parasitised by this broomrape and enthusiastic observations
have led to a number of reports of largely unsubstantiated hosts. Table 1 lists a representative
selection of hosts that have been reported over the last 130 years, and includes reports referring to
all varieties of O. minor as well as O. amethystea Thuill. which has been recorded in error for O.
minor in Britain (Clapham et al. 1987; Rumsey & Jury 1991). It is not a definitive list and,
undoubtedly, many species that have been recorded as hosts for O. minor are missing from it.

HISTORY AND IDENTIFICATION OF OROBANCHE MINOR SM. ON CRYMLYN BURROWS

The accurate records of Rev. J. Lightfoot describing his visit in 1773 to “‘Breton Sands’’, in the
Vicinity of Crymlyn Burrows (Carter 1954), do not mention any broomrapes. One of the first records
of O. minor in the region was made by A. M. Barnard at Swansea in 1853 and nearly a hundred years
later, in 1943, J. A. Webb, a very active, local botanist, recorded it at nearby Aberavon West
Burrows (Hyde & Wade 1957) where it still grows today. During this period, this species would have
been regarded as a local rarity and its presence would have attracted the attention of active

| recorders. It is possible that O. minor was present in small numbers on Crymlyn Burrows at that

time but, if so, it remained unrecorded until 1970 when a small population of about 100 individuals

| was noted. By 1979, the population had increased markedly to approximately 6000 plants (author’s
+ own records). It appears that O. minor is either a relatively recent arrival at Crymlyn Burrows
| (within the last 50 years) or was present in small numbers until the 1970s but had been overlooked by
_ anumber of recorders. Similar, but smaller, populations of O. minor now occur elsewhere in West

116 C. R. HIPKIN

Glamorgan, e.g. on the sand dune systems of Swansea Bay, South Gower and Margam Burrows and
in Mid Glamorgan, on Kenfig Burrows.

Several casual recorders have visited Crymlyn Burrows in the last ten years and named the
Orobanche population there tentatively as O. maritima Pugsley. However, the morphological
features of this population conform strongly with O. minor var. minor as described by Rumsey &
Jury (1991).

Thus:

1. Plants varied greatly in size, up to 60 cm in height, with the largest individuals usually found in
association with Eryngium maritimum and then often in large clumps. Plants associated with other
hosts were rarely as vigorous.

2. The bases of stems were not obviously bulbous.

3. The sizes and densities of inflorescences were variable. For example, inflorescence lengths
varied from 5 cm to 50 cm with overall flower densities varying between 1-4 and 3-2 flowers/cm.
Some individuals had inflorescences which were lax below and dense above and some were
relatively lax throughout. Other plants exhibited inflorescences which were relatively dense
throughout their length.

4. Bracts were 14-18 mm, similar in length to the flowers but often slightly Splenic in upper
flowers.

5. Calyces were 11-13 mm and unequally bifid.

6. Corollas were 15-18 mm, suffused purple, glandular pubescent, the lower lip with subequal
crisped lobes.

7. Flower diameters were always greater than 5 mm.

8. Filaments were hairy below and inserted 2 mm above the corolla base.

9. Stigma lobes were purple and united at base. |

A smali number of pigment-less individuals were noted in the Crymlyn Burrows populations
during the study; similar albino forms have been noted by Rumsey & Jury (1991).

It should also be noted that Daucus carota subsp. gummifer is not a host for O. minor on Crymlyn
Burrows.

HOST RANGE OF OROBANCHE MINOR SM. ON CRYMLYN BURROWS

O. minor var. minor has been observed to parasitise a number of hosts on Crymlyn Burrows. So far,
15 species of host have been identified and these are indicated in Table 2. Each host was —
substantiated by direct observation of contact between the host root and the distinct, swollen
haustorial region of the parasite. Where indicated, root specimens were taken, sectioned and
examined under the light microscope. When this was done, microscopic examination confirmed
physical interaction between host and parasite vascular systems.

The most commonly recorded host was Eryngium maritimum. On Crymlyn Burrows, this species
is very common in open mobile sand communities behind the foredunes, where there are often high
densities of broomrape spikes (up to 25 spikes/m*). Other plants recorded with broomrapes 1 in this
type of community are listed in Table 2(a). Many broomrape spikes were noted in the vicinity of
Marram Grass (Ammophila arenaria) in this community. Indeed, several plants were noted growing
in the middle of marram tufts. All such plants were investigated; none made any apparent contact
with marram roots, but did make contact with the extensive root systems of FE. maritimum.
Parasitism of hosts other than E. maritimum appears to be relatively rare in this community but a
single broomrape individual was observed to parasitise Pastinaca sativa.

More species were parasitised by broomrapes in partially fixed sand communities where E.
maritimum is rare but legumes and composites are conspicuous. Typical species in these
communities are listed in Table 2(b). The most commonly recorded hosts here were Ononis repens,
Lotus corniculatus and Hypochoeris radicata. In addition, the composites Leontodon taraxacoides,
Crepis capillaris, C. vesicaria, Senecio jacobaea, Erigeron acer and Tragopogon pratensis were also
recorded as hosts. There was only one substantiated record of root contact with the legume Anthyllis
vulneraria, although this species dominates large areas of partially fixed-dune on Crymlyn Burrows.
Other minor hosts recorded were the crucifers Arabis hirsuta and Coincya monensis subsp.
recurvata, and the plantain, Plantago lanceolata.

HOST RANGE AND SPECIFICITY OF OROBANCHE MINOR SM.

117

TABLE 2. HOSTS FOR OROBANCHE MINOR VAR. MINOR IN MOBILE AND PARTIALLY FIXED
SAND COMMUNITIES ON CRYMLYN BURROWS

(a) Mobile sand communities

+ root contact with O. minor observed.

(b) Partially fixed sand communities

CRUCIFERAE CRUCIFERAE
Coincya monenis subsp. recurvata Arabis hirsuta ats
Matthiola sinuata Coincya monenis subsp. recurvata ats
CRASSULACEAE VIOLACEAE
Sedum acre Viola canina —
ONAGRACEAE V. tricolor subsp. curtisii =
Oenothera cambrica CARYOPHYLLACEAE
UMBELLIFERAE Cerastium diffusum —
Eryngium maritimum* C. fontanum ~
Pastinaca sativa Arenaria serpyllifolia —
EUPHORBIACEAE GERANIACEAE
Euphorbia paralias Geranium molle -
CONVOLVULACEAE LEGUMINOSAE
Calystegia soldanella Ononis repens* +
COMPOSITAE Trifolium arvense —
Senecio jacobaea Lotus corniculatus +
S. squalidus Anthyllis vulneraria +
Hypochoeris radicata UMBELLIFERAE
Leontodon taraxacoides Pastinaca sativa —
Crepis capillaris GENTIANACEAE
CYPERACEAE Centaurium erythraea ==
Carex arenaria PLANTAGINACEAE
GRAMINEAE Plantago lanceolata +
— Vulpia fasciculata COMPOSITAE
Ammophila arenaria Senecio jacobaea a
Phleum arenarium S. squalidus —
| S. vulgaris =
Erigeron acer +
Carlina vulgaris =
Hypochoeris radicata oP
) Leontodon taraxacoides +
Tragopogon pratensis +
Crepis capillaris ste
) C. vesicaria A

_ * Haustoria-root connections of some plants were sectioned and examined under the light microscope.

Taraxacum officinale agg.
GRAMINEAE

Festuca rubra

Aira praecox

Ammophila arenaria

Phleum arenarium

In transitional areas between mobile and partially fixed sand communities, Eryngium maritimum
often grows in close proximity to Ononis repens, Lotus corniculatus and Hypochoeris radicata. In
i those places, broomrapes were observed making root contact with each of these species.

_ On four separate occasions, single specimens of O. minor were observed making root contact
“with two separate hosts. The following dual-host combinations were noted: (a) Hypochoeris
-radicata and Ononis repens (in partially fixed sand); (b) Eryngium maritimum and Lotus
corniculatus (transitional areas); (c) Crepis capillaris and Ononis repens (in partially fixed sand); (d)
Crepis capillaris and Plantago lanceolata (partially fixed sand).

On one occasion a triple-host combination of Arabis hirsuta, Hypochoeris radicata and Lotus
corniculatus was noted in a partially fixed sand community.

_ Minor hosts such as Arabis hirsuta and Erigeron acer often appeared greatly weakened by the

118 C. R. HIPKIN

parasite and supported smaller broomrape specimens with lax inflorescences. In contrast, common
hosts such as Eryngium maritimum, Ononis repens and Lotus corniculatus often showed normal
vigour and the roots of leguminous hosts that were investigated always bore abundant nodules. E.
maritimum often supported large broomrape specimens with dense inflorescences in clusters of 20
or more individual spikes.

Seeds of O. minor var. minor taken from specimens parasitising Eryngium maritimum in mobile |

sand communities were sown into separate pots containing Hypochoeris radicata (grown from seed
taken from an individual in a typical partially fixed-dune community), Trifolium hybridum (cv.
‘Alsyke’) or T. pratense (transplanted from a local meadow) as potential hosts. All three species
were able to act as hosts and supported specimens of O. minor that flowered and set seed.

O. minor on Kenfig Burrows (GR SS/796.812) appeared to parasitise Eryngium maritimum and
Ononis repens, mostly, whereas a small and short-lived population which occurred on a roadside
verge in Swansea (GR SS/652.912) appeared to be parasitising Trifolium pratense and T. repens.
However, none of these plants was excavated and investigated for root contact.

DISCUSSION

Positive verification of host species for O. minor in the field is difficult, particularly when specimens
are growing in compacted soil, due to the fragile connections between parasite and host roots.

Moreover, in some areas, O. minor is a rare or local species and potentially destructive sampling |
should not be attempted. Under these circumstances the observer has to rely on a judgement based |
on the presence and proximity of suitable hosts. Most records of host species, such as those in the |

survey shown in Table 1, are probably based on such observations. However, significant patterns
emerge from this survey and a number of relevant points may be deduced from it as follows:

1. Dicotyledonous perennials predominate as hosts for O. minor, whereas woody perennials are
rarely reported.

2. Certain families such as Leguminosae and Compositae appear to contain several species which
are suitable hosts for O. minor and there are many reports in the literature of parasitism on
members of these families.

3. Other families also appear to be well represented in the survey such as Umbelliferae,
Plantaginaceae, Geraniaceae and Labiatae.

4. Reports of the parasite on grasses (Murray 1896; Murphy 1972; Rumsey & Jury 1991) are
infrequent and of interest. Parasitism of grasses (or other Monocotyledons) is rare in the genus
Orobanche although it is a feature of other genera such as Aeginetia and Striga (Stewart & Press
1990).

The study described here verified a number of the hosts given in Table 1 and underlines the
importance of certain families such as Umbelliferae, Leguminosae and Compositae (see Table 2).
Nevertheless, although Leguminosae was well represented in the study area, it was notable that
certain species such as Anthyllis vulneraria, Lathyrus pratensis, L. tuberosus, Medicago sativa subsp.
falcata, M. lupulina, Melilotus alba, M. officinalis, Trifolium arvense, T. fragiferum, Vicia cracca
and V. sativa were rarely or never parasitised. In contrast, a much higher proportion of the
composites represented were parasitised. Interestingly, two crucifers, Arabis hirsuta and Coincya
monensis subsp. recurvata, were recorded as hosts although members of Cruciferae have rarely
been reported as hosts before.

One novel feature that emerged from this study was the occasional observation of O. minor var.
minor individuals parasitising more than one host, each one a representative of a separate family.
Presumably, this happens during establishment on a primary host when a haustorium also makes
contact with the root of another compatible, secondary host in the near vicinity.

—— eee ——

Parasitism of more than one host at the same time may occur commonly with other populations of |

O. minor var. minor, a consequence of its very wide host spectrum. However, there is evidence that

O. minor has developed host specific strains (Musselman & Parker 1982) and, clearly, populations ;

of these would be limited in this respect.
From the outset, it seemed possible, but intuitively unlikely, that the Crymlyn Burrows
population was composed of more than one physiological strain, each with a limited set of hosts. For

example, it could be argued that the mobile dune population which appeared to parasitise Eryngium |

HOST RANGE AND SPECIFICITY OF OROBANCHE MINOR SM. 119

maritimum almost exclusively, was physiologically distinct from the partially fixed dune population
which appeared to parasitise a wider spectrum of hosts. However, O. minor var. minor also occurs
in transitional areas between well-defined mobile and partially fixed sand communities, where E.
maritimum and other hosts are parasitised. Furthermore, it was shown that seed taken from a
specimen parasitising E. maritimum in a mobile dune community germinated to produce vigorous
specimens in pots containing Hypochoeris radicata, Trifolium pratense or T. hybridum. This
evidence, together with observations of individuals parasitising multiple, unrelated hosts (including
the combination of E. maritimum and Lotus corniculatus), does not support the argument that there
are a number of host specific strains of O. minor var. minor on Crymlyn Burrows.

The broomrape population of Crymlyn Burrows has become a striking feature of this actively
accreting coastal dune system in recent years. However, although it is likely that a small population
of O. minor was established there about 50 years ago, it seems probable that only in the last 20 years
or so has it spread so conspicuously into mobile and partially fixed dune areas. Indeed, the
establishment and spread of this population on Crymlyn Burrows resemble colonisations there by a
number of non-native colonists such as Coincya monensis subsp. recurvata, Hirschfeldia incana,
Senecio squalidus and Conyza canadensis. It is notable, therefore, that Rumsey & Jury (1991)
suggest that O. minor var. minor may not be native in the British Isles.

ACKNOWLEDGMENTS

I am grateful to Mr Ian Tew for his invaluable help in the histological preparation of specimens and
to Mrs Hilary Hipkin for her assistance in the field. Thanks also to Quentin Kay for allowing me
access to his collection of county Floras.

REFERENCES

ALLEN, D. E. (1984). Flora of the Isle of Man. Douglas.
ARCHER Briccs, T. R. (1880). Flora of Plymouth. London.
Bevis, J., KETTLE, R. & SHEPHARD, B. (1978). Flora of the Isle of Wight. London.
Bowen, H. J. M. (1968). The Flora of Berkshire. Oxford.
Carter, P. W. (1954). Some account of the history of botanical exploration in Wales. Trans. Cardiff Nat. Soc.
82: 5-31.
CHATER, A. O. (1986). Orobanche hosts — a cautionary tale. B.S.B.I. News 42: 10.
CLAPHAM, A. R., Tutin, T. G. & Moore, D. M. (1987). Flora of the British Isles, 3rd ed. Cambridge.
Davey, F. H. (1909). Flora of Cornwall. Penryn.
Dony, J. G. (1953). Flora of Bedfordshire. Luton.
Grose, D. (1957). Flora of Wiltshire. Devizes.
HamBLer, D. J. (1991). Orobanche maritima: an overlooked feature? B.S.B.I. News 58: 18-19.
HOLianp, S. C., Cappick, H. M. & DupDLey-Smitu, D. S. (1986). Supplement to the Flora of Gloucestershire.
Bristol.
Hype, H. A. & WapE, A. E. (1957). Welsh flowering plants. Cardiff.
Jones, M. ee Orobanche hosts. B.S.B.1. News 41: 29.
Kent, D. H. (1975). The historical Flora of Middlesex. London.
Linton, E. F. (1919). Flora of Bournemouth. Bournemouth.
_Lous.ey, J. E. (1976). Flora of Surrey. London.
| Marcetts, L. J. & Davin, R. W. (1981). A review of the Cornish Flora 1980. Redruth.
/McCuintock, D. (1975). The wild flowers of Guernsey. London.
/ MESSENGER, K. G. (1971). Flora of Rutland. Leicester.
‘Murpny, D. P. (1972). Orobanche hosts at Glasnevin. Ir. Nat. J. 17: 279.
Murray, R. P. (1896). Flora of Somerset. Taunton.
MUSSELMAN, L. J. & PARKER, C. (1982). Preliminary host ranges of some strains of economically important
broomrapes (Orobanche). Econ. Bot. 36: 270-273.
| Patrick, S. & Hoiiick, K. M. (1975). Supplement to the Flora of Derbyshire. Derby.
Petcn, C. P. & Swann, E. L. (1968). Flora of Norfolk. Norwich.
Pussiey, H. W. (1940). Notes on Orobanche L. J. Bot., Lond. 78: 105-116.
)Ravensuaw, T. F. (1860). Flowering plants and ferns of Devonshire. London.
\Rumsey, F. J. (1991). Orobanche maritima revisited. B.S.B.I. News 59: 9-10.

en ae ae

120 C. R. HIPKIN

Rumsey, F. J. & Jury, S. L. (1991). An account of Orobanche L. in Britain and Ireland. Watsonia 18: 257-295.

SALMON, J. D. (1863). Flora of Surrey. London.

SaviDGE, J. P., HEywoop, V. H. & Gorpon, V., eds. (1963). Travis’s Flora of South Lancashire. Liverpool.

STEWART, G. R. & Press, M. C. (1990). The physiology and biochemistry of parasitic angiosperms. Ann. Rev.
Plant Physiol. 41: 127-151.

Swann, E. L. (1975). Supplement to the Flora of Norfolk. Norwich.

TRIMEN, H. & THISELTON-DyErR, W. T. (1869). Flora of Middlesex. London.

Wess, D. A. & Cuater, A. O. (1972). Orobanche L., in Tutin, T. G. et al., eds. Flora Europaea 3: 286-293.
Cambridge.

Waite, J. W. (1912). Flora of Bristol. Bristol.

Wo Ley-Dop, A. H. (1970). Flora of Sussex. Bristol.

(Accepted October 1991)

Watsonia, 19, 121-126 (1992) 121

The current distribution and abundance of Orchis ustulata L.
(Orchidaceae) in the British Isles — an updated summary

MJ.“ Y- FOLEY

87 Ribchester Road, Clayton-le-Dale, Blackburn, Lancs., BB1 9HT

ABSTRACT

Recent British records of Orchis ustulata L. (Orchidaceae) are described and the current range of the species is
shown both as a distribution map and in tabular form. This updates information presented by the author in
earlier work.

INTRODUCTION

The status and distribution of Orchis ustulata L. in the British Isles has been previously described on
a regional basis (Foley 1987, 1990). Further exploratory work and additional knowledge gained as a
result of publication of this information suggests that an updated summary is now appropriate. The
distribution maps which were originally included were to a differing degree of precision, and the
opportunity is now taken to combine all the available information on geographical distribution into
a single map plotted on a 10-km square basis.

DISCUSSION

The additional records shown below are relatively small in number and do not significantly affect the
overall distribution pattern of O. ustulata as previously described. They often represent small
isolated populations containing just a few plants at, or close to, localities previously thought likely to
be extinct. Such examples include Langdon Bay (E. Kent, v.c. 15), Waddingham (N. Lincs., v.c. 54)
and Knaresborough (Mid-W. Yorks., v.c. 64), at each of which a welcome reappearance of the plant
has occurred. Other new records are for small populations from areas in which there is apparently
no previous known occurrence — Scarcliffe (Derbys., v.c. 57), Sandale (N.E. Yorks., v.c. 62), and
some in S. Wilts. (v.c. 8). Just occasionally, population strength has been previously underesti-
mated, in some cases significantly so, as at Martin Down (S. Hants., v.c. 11) which in fact is now
confirmed to hold two important colonies. One of these is of especial interest in that it is in an area
ploughed in 1957, the only instance so far noted where recolonisation has occurred so quickly.
During 1987 a significant find was made in N.W. Yorks. (v.c. 65) by the author and a colleague
where several hundred flowering plants occur in association with O. morio L. on traditionally
managed, lightly grazed pasture. This is undoubtedly the best surviving population in northern
England. Except by the tenant farmer, by whom its significance and extent were not appreciated,

_ this strong population has apparently been previously overlooked.

Two other unrecorded populations of significance were identified at Great Cheverell Hill (S.

| Wilts., v.c. 8) by G. Goodfellow, and others have been located, also in v.c. 8, on Ministry of
_ Defence land during lulls in artillery firing. The good flowering season of 1989 illustrated that in

Ae Re A NS

Britain, there are still some very significant populations of O. ustulata and this was particularly
apparent at Parsonage Down (S. Wilts., v.c. 8), where one estimate suggests that the population is
in excess of 30,000 flowering plants. This is perhaps the most important surviving single population
of O. ustulata in north-western Europe.

The late-flowering (July-August) form of O. ustulata, often recorded from southern England, has
recently been described as var. aestivalis by Kimpel (1988) and subsequently raised to subspecific

122 M. J. Y. FOLEY

a he

rank as subsp. aestivalis by Kimpel & Mrkvicka (1990). These latter authors record details of its
morphometry and deviation from the type, based on observations made on Central European
populations where white-flowered plants are also noted for this newly described taxon.

The additional British records described below, together with those previously contained in Foley
(1987, 1990) (the latter modified to include new information where appropriate), have been
combined and mapped on a 10-km square basis as shown in Fig. 1. The species is considered to be

PLOTTED ON
UTM GRID

Ficure |. The past and present distribution of Orchis ustulata in the British Isles. Plotted in 10-km squares: @
still surviving, < considered extinct.

ir AL

DISTRIBUTION AND ABUNDANCE OF ORCHIS USTULATA L. 123

extant if it has been confirmed to be present in or after 1987, or if (and especially in the case of small
colonies where an annual reappearance in flower can be unreliable) it has been seen shortly before
this date and there has been no known change to the habitat likely to cause its loss. Additionally, all
54 10-km squares of the British Isles in which O. ustulata still occurs are listed in Table 1, and this
also includes details of the respective population strengths. The 174 10-km squares for which the
plant now appears to be extinct are given in Table 2, although it is possible that in some cases the
plant may still reappear in small numbers.

Further records for O. ustulata from any new or existing localities are still welcomed, and
especially so are details of any summer-flowering populations.

RECORDS

The following records should be added to, or where appropriate replace, those given earlier (Foley
1987, 1990). Population strengths (in parentheses) use the code employed in the earlier papers, as
follows: 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.

N. Wilts., v.c. 7: SU/2.7, three new localities to the north of Aldbourne (A, A, A) (G. Goodfellow
pers. comm.).

S. Wilts., v.c. 8: ST/8.3, Whitesheet Hill, a few plants over a large area (A); ST/9.5, Great Cheverell
Hill (see Foley 1990), three additional sites (A, B, E); SU/0.2, Hoopside (A); Middleton Down
(R. Laurence pers. comm.) (A); Knighton Down, c. 50 plants in 1989 (D); Throope Down, on
rank grassland, possibly lost (PX); SU/0.3, Steeple Langford — Cow Down, very good numbers in
1987 when grazing relaxed (R. Laurence pers. comm.) (E) — this should replace “‘(U)” in Foley
(1990); Hadden Hill, about 30 plants in 1989 (C); SU/0.4, Parsonage Down, recent estimates
including the very good season of 1989 suggest a population of tens of thousands (one estimate
30,000+ plants), perhaps the best site in north-western Europe — this extends over an area
approaching 95 ha and is now considered to be one continuous large colony of plants; SU/0.5, Slay
Down, English Nature [formerly N.C.C.] information of a colony in centre of M.O.D. impact
area — strong colony, no details, perhaps (D) in 1988; SU/1.2, Homington Down, 40 plants at one
site in 1989, status should therefore be (A, C) (see Foley 1990); SU/1.3, Cockey Down,
discovered by P. Mobsby in 1988 (A); SU/1.4, Alton Down, two small colonies recorded by
English Nature (A, A); SU/1.5, Chisenbury Warren, small colony on M.O.D. land in 1984 (A);
Giant’s Grave, Pewsey Hill, up to 40 plants in 1985 on an ancient monument (C); Milton
Lambourne, 20 plants in 1989 (B); SU/2.2, Witherington Down, 20 plants in 1988 — status now
(B); SU/2.5, Haxton Down, reported here in 1986 by D. Soden on M.O.D. land (A). (Except
where shown, all details for v.c. 8 are ex G. Goodfellow pers. comm.)

S. Hants., v.c. 11: SU/0.1, Martin Down, two populations, one of which is in old arable, last
ploughed in 1957 (E), and on old grassland (E). Also on Bokerley Dyke (P. Toynton pers.
comm.) (D). There were substantial populations in 1991 with two plants of the white-flowered
form occurring on the Dyke.

E. Sussex, v.c. 14: TQ/4.0, Mount Caburn, Glynde at 44.08 — correct (U) to (F), there were 3000+
plants here in 1991 (F. Rose pers. comm.); TQ/5.0, Charleston Bottom, correct (A) to (D);
Lullington Heath, correct (A) to (D), plants of late-flowering variant at both sites (D. C. Lang

. pers. comm.).

E. Kent, v.c. 15: TR/3.4, Langdon Bay, a single flowering plant was seen in 1981 (F. Horsman pers.

- comm.) (A).

Surrey, v.c. 17: TQ/2.4, three specimens in KDL collected in 1885 (X).

W. Suffolk, v.c. 26: TL/9.5, etc., Shelland, and TM/0.4, near Hadleigh (see Foley 1990) — there is
some doubt about these records.

N. Lines., v.c. 54: SK/9.9, a single plant appeared in 1991 at the Waddingham site (see Foley 1987)
after an absence of ten years (I. Weston pers. comm.).

Derbys., v.c. 57: SK/2.5, eight additional very small populations (see Foley 1990), located as
follows: one locality south of White Edge (L. Storer pers. comm.); four additional sites to the

124 M. J. Y. FOLEY

TABLE 1. LOCATION AND RESPECTIVE STRENGTHS OF EXTANT POPULATIONS OF ORCHIS
USTULATA IN THE BRITISH ISLES BY 10-KM SQUARE

Grid reference of Number of Population strength
10-km square populations (for code — see text)

ST/8.1
ST/8.3
ST/9.5
SD/9.8
SD/9.9
SZ/4.8
SU/0.1
SU/0.2
SU/0.3
SU/0.4
SU/0.5
SU/0.6
SU/1.2
SU/1.3
SU/1.4
SU/1.5
SU/1.6
SU/2.2
SU/2.4
SU/2.5
SU/2.7
SU/2.8
SU/3.3
SU/3.5
SU/3.6
SU/4.5
SU/5.5
SU/5.8
SP/1.3
SK/2.5 1
SK/5.7
SK/9.4
SK/9.9
SE/0.8
SE/1.8
SE/2.8
SE/3.5
SE/7.1
SE/8.8
SE/9.8
NZ/4.3
TV/5.9
TQ/2.1
TQ/3.0
TO/4.0
TO/4.1
TO/5.0
TOQ/8.6
les
TL/3.3
TR/0.4
TR/2.4
TR/3.4
*G.R. withheld

°
H
ry

wOwW>
go
tT

>
Q
Q

,A,A,A,A,A,A,A,A,A,A,B,D,E

>
>
>

oo
es)
0

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w
w
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tn

,A,A,A,B

OR ODO OB RR ND RR OD BR RR BR RR BR RRB Re nN Re eK KN BR WN PR BRR WN W BRK NY BRR

Co a le A a a SO a oO ee a lt lt ol a lt Ot a a a a a a a

* details withheld for a sensitive locality in v.c. 65 (N.W. Yorks.).

DISTRIBUTION AND ABUNDANCE OF ORCHIS USTULATA L. 125

TABLE 2. 10-KM SQUARES FOR WHICH ORCHIS USTULATA IS NOW CONSIDERED TO BE
EXTINCT IN THE BRITISH ISLES

SX/5.4 NY/4.5 SK/5.5 TQ/2.0

SX/8.3 NY/5.3 SK/5.9 TQ/2.4
SX/9.5 NY/5.4 SK/6.4 TQ/2.5
SX/9.6 NY/6.1 SK/7.4 FOR
SY/8.8 N77 1 SK/7.5 TOQ/5.5
SY/8.9 SZ/0.8 SK/9.0 TOQ/S5.7
ST/3.6 S229 SK/9.1 TQ/6.7
ST/4.5 SZ/5.7 SK/92 TQ/7.6
ST/4.7 SZ/5..8 SK/9.5 TL/0.1
Si/5.7 SU/1.7 SE/2a7. TL/0.2
ST/7.6 SU/1.8 SE/3.4 TL/0.3
Si/7.7 SU/2.3 SE/3.6 TL/0.8
ST/8.0 SU/3.2 SE/3 7 TE/O:9
ST/8.5 SU/3.8 SE/4.3 TE/2.3
ST/8.6 SU/3.9 SE/4.4 TL/3.4
ST/9.1 SU/4.2 SE/4.8 TL/4.4
S/9°3 SU/4.3 SE/5.0 TL/4.5
ST/9.4 SU/4.8 SES: TL/S.4
ST/9.6 SU/5.4 SE/5.4 SPIES YS
ST/9.9 SU/6.2 SE/5:5 TL/S.6
SO/4.7 SU/6.7 SE/6.2 TL/6.6
SO/5.1 SU/7.1 SE/6.3 Wes
SO/5.7 SU/7.8 SE/6.4 TL/7.6
SO/5.8 SU/9.4 SE/6-7 TL/8.3
SO/5.9 SP/0.0 SE/6.8 TL/9.5
SO/6.0 SP/0.2 SE/7.4 TF/0.0
SO/7.4 SP/0.3 SE/7.6 TF/0.1
SO/7.6 SP/1.0 SEIT TF/0.3
SO/7.8 SP/T:1 SE/7.8 TE/OS
SO/8.0 SP/1.2 SES TF/6.0
$O/8.8 SP/3.0 SE/9.4 TA/0.1
$O/9.0 SP/4.0 SE/9.9 TA/0.7
SO/9.1 SP/4.1 NZ/1.0 TA/0.8
SO/9.2 SP/6.0 NZ/2.1 TA/1.0
S$O/9.3 SP/8.0 ; NZ/2.5 TA/1.4
$J/5.0 SEO NZ/3.1 TA/1.7
SD/4.7 SK/0.4 NZ/3.3 TR/0.6
SD/5.6 SK/0.7 NZ/3.6 TR/1.3
SD/6.9 SK/2.6 NZ/3.7 TR/1.5
SD/8.6 SK/4.5 NZ/4.5 TR/2.5
mY /1 3 SK/4.9 NZ/4.6 TM/0.4
NY/3.4 SK/5.0 NZ/9.0 TM/3.8
NY/3.5 SK/5.2 TQ/0.9

NY/4.3 SK/5.4 TQ/1.4

north-west of Brassington (L. Storer & B. G. Tattersall pers. comm.); two additional sites to the

_ south-west of Longcliffe (east of Beardsley Plantation) (L. Storer pers. comm.); near Pinder’s
Rock in 1987, recorded by M. Buckley — all these probably (A); SK/5.7, Scarcliffe, lightly grazed
Magnesian limestone meadow, discovered in 1986 by R. A. Frost, three plants (A).

S. E. Yorks., v.c. 61: SE/6.4, Fulford Ings, collected by H. Britten on 13 May 1928, herb. P. Burnett
(R. Gulliver pers. comm.) (X); SE/9.4, Arras (not TA/0.4, Arram (Foley 1987) — an error)
(Robinson 1902) (X).

N. E. Yorks., v.c. 62: SE/5.5, Clifton Ings, still extant in 1926 when collected by H. E. Britten, herb.
P. Burnett (R. Gulliver pers. comm.) (X); SE/6.7, Hovingham, specimen collected in 1871 by
“M.T.”, herb. P. Burnett (R. Gulliver pers. comm.) (X); Wiganthorpe Park and also Terrington
Carr and adjoining area, recorded by Yorkshire Naturalists’ Union in 1898 (X); SE/7.6 (or 7.7),

126 M. J. Y. FOLEY

Castle Howard district, old record “‘in dry stoney meadows” (X); St Ann’s meadow, Castle
Howard Park (X); SE/7.8, Gundale, possibly Gundale Slack (U); SE/8.8, Sandale, one plant in
1989 (I. Lawrence pers. comm.) (A).

Mid-W. Yorks., v.c. 64: SE/3.5, Knaresborough, east of Birkham Wood, the last sighting was in
1967 after which the site was ploughed, but a single flowering plant was re-found by J. Barker in
1988 nearby. Regrettably this was immediately dug up by an unknown person (A).

N.W. Yorks., v.c. 65: SD/9.8, Seata, Aysgarth, limestone pasture (A); SE/0.8, east of Aysgarth,
two new small populations (A, A); SE/1.8, near Middleham, riverside pasture (A). (Note: In
1987 the author and S. Priest discovered a new locality in v.c. 65 on traditionally managed
pasture. This is almost certainly the strongest surviving population in northern England (E). No
details are given here as this is a very sensitive site, but relevant details are known to English
Nature.)

Durham, v.c. 66: NZ/2.5, Urpeth in 1913, J. W. Heslop-Harrison (Graham 1988) (X).

Cumberland, v.c. 70: NY/5.4, Armathwaite, on west bank of the Eden north of the bridge. Last
recorded in 1944 by F. Simpson (X).

ACKNOWLEDGMENTS

Contributions of information on new or overlooked localities have been received from many
people, all of whom I would now like to thank. Especially this includes G. Goodfellow, D. C. Lang,
R. Laurence, D. Millward, M. Sanderson, B. G. Tattersall and P. Toynton, for their detailed help.

REFERENCES

Fo.ey, M. J. Y. (1987). The current distribution and abundance of Orchis ustulata L. in northern England.
Watsonia 16: 409-415.

Fotey, M. J. Y. (1990). The current distribution and abundance of Orchis ustulata L. in southern England.
Watsonia 18: 37-48.

GranaM, G. G. (1988). The flora and vegetation of County Durham. Durham.

KUmpeL, H. (1988). Uber eine spatblihende Orchis ustulata-Sippe. Haussknechtia (Jena) 4: 23-24.

KUmpeL, H. & Mrxkvicka, A. C. (1990). Untersuchungen zur Abtrennung der Orchis ustulata L. subsp. aestivalis
(Kiimpel) Kimpel & Mrkvicka. Mitt. Bl. Arbeitskr. Heim. Orch. Baden-Wiirtt. 22: 306-324.

Rosinson, J. F. (1902). The flora of East Riding of Yorkshire. London & Hull.

(Accepted January 1992)

|
|

Watsonia, 19, 127-129 (1992) 127

Urtica galeopsifolia Wierzb. ex Opiz (Urticaceae) in Wicken
Fen (E. England)

D. V. GELTMAN

Herbarium, Komarov Botanical Institute, Prof. Popov St 2, St Petersburg 197376,
Russian Federation

ABSTRACT

A plant resembling the common nettle, Urtica dioica L., but almost lacking stinging hairs, occurs in Wicken Fen,
Cambridgeshire. It is here considered as a separate species U. galeopsifolia Wierzb. ex Opiz, which differs from
the former by several morphological characters and also by ploidy level. Some suggestions about the possible
hybrid origin of U. dioica are put forward.

INTRODUCTION

Many British botanists know the strange almost stingless form of the common nettle (Urtica dioica
L.), which occurs within the National Trust’s Wicken Fen nature reserve in Cambridgeshire,
GR TL/55.70. These unusual plants grow in damp mesotrophic fen communities, in particular, at
the margins of shrubby areas with other shade-tolerant herbs. Similar plants occur nearby, as at
Chippenham Fen (Perring et al. 1964).

The variation of morphological characters of U. dioica, with special reference to this particular
population, was intensively investigated by Pollard & Briggs (1982, 1984a). They explained that the
difference of this plant from typical ones arose through a process of gene flow. An exact taxonomic
recognition had not been made, but it was mentioned that, probably, the “‘Wicken nettle’ belongs to
var. angustifolia Wimm. & Grab. or var. subinermis Uechtr. The Flora of Cambridgeshire (Perring
et al. 1964) treats these plants as f. angustifolia (Wimm. & Grab.) Moss.

While studying the taxonomy of Urtica L. in the former U.S.S.R., I paid attention to the
information published by Pollard & Briggs (1982). My assumption was that the ‘““Wicken form of
common nettle” seemed to belong to a separate species, U. galeopsifolia Wierzb. ex Opiz, which I
have recognised for Eastern and Central Europe (Geltman 1986, 1992) or to some intermediate
form between this species and U. dioica.

URTICA DIOICA AND U. GALEOPSIFOLIA

U. galeopsifolia was described from Hungary by F. M. Opiz, Naturalientausch 9: 107 (1825),
according to Domin (1943), and it differs from U. dioica mainly by characters of the leaf blades: they
almost completely lack stinging hairs, but always possess a more or less dense indumentum
consisting of simple (non-stinging) hairs. It is also differentiated by the location of its inflorescence;
the lowest flowering branches of U. galeopsifolia appear on the level of the 13th—22nd node, and in
U. dioica on the level of the 7th-14th node. Probably, this feature is linked to the period of
flowering: U. galeopsifolia starts to flower later than U. dioica, approximately in mid-July (Geltman
1986).

Not infrequently U. galeopsifolia has somewhat longer and comparatively narrow leaves, but this
character is unsatisfactory, as typical U. dioica with numerous stinging hairs may sometimes have
such a leaf shape. It is necessary to stress that neither U. galeopsifolia nor U. dioica have such
narrow leaves as the Asiatic species U. angustifolia Fisch. ex Hornem., which does not occur in
Europe.

Unlike U. dioica, which is mostly tetraploid with 2n = 52 or, probably, 2n = 48 (there are only a

128 D. V. GELTMAN

few records of 2n = 26), U. galeopsifolia is presumably diploid (2n = 26) and only one specimen was
determined as tetraploid (Geltman 1984).

U. galeopsifolia prefers a quite distinct type of habitat: damp woodlands, especially with Alnus
glutinosa (L.) Gaertner, river banks and valleys, and eutrophic fens. I found a very clear example of
the ecological separation of the two species in question in Central Ukraine, Cherkassy region.
U. galeopsifolia occurred in Phragmites fen and Alnus glutinosa woodland, situated in deep valleys,
bordered with hills, and U. dioica is found in more or less dry (at least, not swampy) Carpinus
betulus L. woodlands on the hills, especially near roads and cattle paths.

In the former U.S.S.R., U. galeopsifolia is distributed in the European part to the south of
latitude 60°N, and also in the southern regions of Siberia eastwards to the river Angara; some
localities are also known from Caucasia. It should be widely distributed in Atlantic, Central and
East Europe, but before I had visited some British herbaria, I had seen quite reliable specimens of
this species only from Hungary, Czechoslovakia and the Netherlands (Geltman 1986).

In August 1991, I was fortunate to be able to visit Wicken Fen and to observe the local nettle
population. In my view, the almost stingless plants which occur in the fen proper do not differ from
U. galeopsifolia, as it occurs in Russia. So, U. galeopsifolia is a new species record for the British
flora.

U. galeopsifolia, nevertheless, is not a ‘completely good’ species, especially in terms of the species
concept adopted in Flora Europaea (Geltman 1992). Sometimes transitional forms to U. dioica may
be found, such as forms without stinging hairs but with scattered simple ones (I usually refer such
plants to U. dioica). So, probably, some botanists may prefer to treat this taxon as a subspecies; the
corresponding combination U. dioica subsp. galeopsifolia (Wierzb. ex Opiz) Chrtek does exist
(Chrtek 1981). But I consider that species rank is more appropriate for this taxon for the following
reasons. It is correct to apply the rank of subspecies to allopatric taxa of widely distributed species.
But in the case of U. dioica and related species we have, obviously, quite a different situation: U.
dioica is, probably, a species of hybrid origin (Geltman 1990). The first of its ancestors might be U.
galeopsifolia (or a species closely related to it) and the second, U. sondenii (Simm.) Avrorin ex
Geltman, which occurs mainly in the taiga zone of West and Central Siberia and Northern Europe
also. Detailed distribution characteristics with map are given in Geltman (1986). This species has
completely glabrous leaf blades with neither stinging nor simple hairs; some scattered stinging and
simple hairs are located on the nodes of stems and on the inflorescence axis. It grows in meadows
and Salix-carrs in river valleys, near streams, in damp forests, etc. Like U. galeopsifolia, it is a
diploid (2n = 26).

U. galeopsifolia was likely to be formed at least by the Pliocene period and might have penetrated _
to the British Isles at the time of existence of bridges with the continent. It survived the Pleistocene
glaciations not far from the southern limits of the ice front. At the same time U. sondenii seems also
to exist in the periglacial area, because the first migration of Siberian taiga species to Europe was
possible at the end of Miocene, the next ones in the late Pliocene and in the interval between Dnepr
and Moscow glaciations (Minyaev 1965)*. So, U. galeopsifolia and U. sondenii were likely to have"
opportunities for hybridisation. Probably, there were no ecological barriers, because the ecological
niches of both species are similar.

As a result of hybridisation of diploid U. galeopsifolia and U. sondenii, a tetraploid ‘primary’ U.
dioica could have been formed. These plants may have been very polymorphic in leaf shape and
indumentum and had no stinging hairs on the leaf blades. Such ‘primary’ forms spread in Europe
after the last glaciation and may also have appeared in Wicken Fen. Then, derived from these
plants, the ‘secondary’, typical U. dioica with stinging hairs on leaf blades developed. The main
selective factor might be a pressure from animals, including domestic ones. As Pollard & Briggs
(1984b, p. 507) have shown, “grazing by large mammals could act as a strong selective force for
higher stinging hair densities in nettles’. So, the origin of typical U. dioica was, apparently,
connected with human activity. ‘““Response to the modification of the habitat by human settlement
seems to be indicated by the nettle as early as the time of Mesolithic culture” (Godwin 1975, p. 242).

Intermediate forms between typical U. dioica with numerous stinging hairs and U. galeopsifolia
are, in fact, mostly primary hybrid forms of U. dioica, which have survived to the present time.

*The interval between the Dnepr and Moscow glaciations seems to correspond with the Hoxnian stage
(interval between Anglian and Wolstonian ice periods) in Britain.

URTICA GALEOPSIFOLIA WIERZB. EX OPIZ IN WICKEN FEN 129

Because there are no classical allopatric races in this case, I prefer to treat U. dioica, U. galeopsifolia
and U. sondenii as separate species in the framework of an U. dioica aggregate or ‘complex’.

After the most recent glaciation, U. galeopsifolia in Britain survived in damp territories, but in
historic times it has become almost extinct due to drainage and now seems to be preserved only in
protected wetland areas (like Wicken Fen). Forms intermediate between U. galeopsifolia and
typical U. dioica may be found in various seminatural situations.

It is necessary also to mention U. pubescens Ledeb., described from the delta of the river Volga
(Ledebour 1833). Sometimes this taxon (more frequently at the rank of the variety U. dioica var.
pubescens (Ledeb.) Trautv.) has been recorded from various regions of Europe. In my opinion, this
taxon is a separate species, closely related to U. galeopsifolia, but differing from it in some minor
characters (Geltman 1986). U. pubescens occurs in special wetland territories, mainly in the Volga
delta, and recently has been found in lower Dnepr. It may be a relict, which survived in such
localities from Tethyan times.

ACKNOWLEDGMENTS

I would like to thank Dr S. M. Walters and Dr J. R. Edmondson for their assistance at the time of
my visit to Wicken Fen and also for commenting on the manuscript. My trip to Britain (in which
programme a visit to Wicken Fen was included) became possible due to sponsorship from the Royal
Society. I am also grateful to the Missouri Botanical Garden, in whose library I worked preparing
this publication.

REFERENCES

CurTEK, J. (1981). Poznamky ke kvetene slovenska. Cas. Narod. Muz., Pad. Prir. 150 (3-4): 209-216.

Domin, K. (1943). Filipa Maximiliana Opize “‘Naturalientausch”’ (1823-1830). Praha.

GELTMAN, D. V. (1984). Cytotaxonomical studies of the species of the genus Urtica (Urticaceae) in the flora of
the U.S.S.R. Bot. Zhurn. 60: 1524-1530. [In Russian]

GELTMAN, D. V. (1986). Systematic and ecological-geographic characteristics of the species from the affinity of
Urtica dioica (Urticaceae) in the flora of the U.S.S.R. Bot. Zhurn. 71: 1480-1489. [In Russian]

GELTMAN, D. V. (1990). Some problems of phylogeny of the species of the subsection Urtica of the genus Urtica
(Urticaceae). Bot. Zhurn. 75: 840-845. [In Russian]

GELTMAN, D. V. (1992). Urtica L., in Tutin, T. G. et al., eds. Flora Europaea, 2nd ed. 1 (In press).

Gopwin, H. (1975). History of the British flora, 2nd ed. Cambridge.

LEDEBOUR, C. F. (1833). Flora altaica, 4. Berlin.

Minyaev, N. A. (1965). Sibirskie taezhnye elementy vo flore severo-zapada evropeyskoy chasti S.S.S.R. [Siberian
taiga elements in the flora of the north-west of the European part of the U.S.S.R.] in Arealy Rasteniy flory
S.S.S.R., pp. 50-92. Leningrad.

PERRING, F. H., SELL, P. D. & WA rTeERs, S. M. (1964). A flora of Cambridgeshire. Cambridge.

PoLLarD, A. J. & Briccs, D. (1982). Genecological studies of Urtica dioica L. I. The nature of intraspecific
variation in U. dioica. New Phytol. 92: 453-470.

PoLiarD, A. J. & Briccs, D. (1984a). Genecological studies of Urtica dioica L. I. Patterns of variation at
Wicken Fen, Cambridgeshire, England. New Phytol. 96: 483-499.

PoLLarD, A. J. & Briccs, D. (1984b). Genecological studies of Urtica dioica L. III. Stinging hairs and plant-
herbivore interactions. New Phytol. 97: 507-522.

(Accepted January 1992)

Watsonia, 19, 131-140 (1992) 131

Short Notes

RUBUS BLOXAMII (BAB.) LEES (ROSACEAE) WITH RED STYLES

Rubus bloxamii, a widespread and locally abundant bramble of central southern England and the
central Midlands, normally has styles that are yellowish-green. Towards the south-western limit of
its English range, however, at the southern end of the border between N. Somerset, v.c. 6, and S.
Wilts., v.c. 8, and abundantly along the Blackdowns astride the border between S. Devon, v.c. 3,
and W. Somerset, v.c. 5, A. Newton and L. J. Margetts have respectively come across populations
in which the styles are bright red (but which otherwise do not differ from the normal examples of the
species in Britain in any significant respect).

On the opposite side of the Channel, in various parts of Normandy and (according to Sudre 1911)
just to the south in dép. Sarthe, there is a common bramble which has long been known as R.
multifidus Boulay & Malbranche. One place it occurs in particular profusion is the Forét de St-
Sauveur, 30 km south of Cherbourg, and specimens I collected there in 1991 have proved to match
the red-styled R. bloxamii of England. A. Newton and H. Vannerom share my impression that
except in this one character R. multifidus is not distinguishable from R. bloxamii and should
accordingly be treated as conspecific, a conclusion reached earlier by Friderichsen (MS note by
Rogers 1897 on BM sheet), Rogers (1900) and, ultimately, Watson (1958). The name nevertheless
seems worth retaining at varietal level in view of the interesting difference in range, and the
necessary new combination is accordingly now made:

Rubus bloxamii (Bab.) Lees var. multifidus (Boulay & Malbranche ex Corbiére) D. E. Allen, comb.
et stat. nov.

R. multifidus Boulay & Malbranche MS (Assoc. rub. exsicc.); Corbiere, Nouv. Fl. Normandie 208

(1894); R. menkei subsp. multifidus (Boulay & Malbranche) Boulay in Rouy & Camus, Fl. Fr. 6: 104

(1900); R. menkei microgene multifidus (Boulay & Malbranche) Sudre, Rubi Eur., 160 (1911).

Corbiére cites no fewer than ten numbers distributed by the Association rubologique as belonging
to R. multifidus. The lectotype must clearly be chosen from the earliest of these (no. 36: Forét de la
Londe, dép. Seine-Inférieure, 1873, A. Malbranche) and I designate as that the example of that
number which I have examined in Paris (P). ;

It is worth adding that R. bloxamii var. bloxamii also occurs in Normandy, but appears to be much
the rarer of the two there. In the North Cotentin (dép. Manche) I have so far seen it in only two
places, in both cases just a bush or two. H. Vannerom informs me (in litt. 1992) that two of
Letendre’s specimens of ‘R. multifidus’ from dép. Seine-Inférieure, Association rubologique nos.
656 and 772, have yellowish styles too. In the Channel Isles, on the other hand, where R. bloxamii is
locally common in Guernsey though scarce in Jersey, the populations consist of var. bloxamii
exclusively. |

REFERENCES

Rocers, W. M. (1900). Handbook of British Rubi. London.
Supre, H. (1911). Rubi Europae (19). Paris.

_ Watson, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge.

D. E. ALLEN
Lesney Cottage, Middle Road, Winchester, Hants., SO22 5EJ

RUBUS CORBIERI BOULAY (ROSACEAE) IN THE BRITISH ISLES

In 1897 the then leading British authority on Rubus, W. Moyle Rogers, paid a first (and only) visit to
the Channel Isles and on two of them encountered quantities of an unfamiliar bramble. While he

132 SHORT NOTES

found it ‘‘rather frequent” on Guernsey, on Sark it was “remarkably abundant” (Rogers & Rogers
1898), so much so that, in the words of his companion on that day, it ‘““seemed to monopolise nearly
the whole place” (Derrick 1898). And as subsequent batologists have found, these words are indeed
no exaggeration: this large, coarse, white-flowered plant is one of the most dominant features of the
island’s landscape. |

Rogers promptly sent a specimen to Focke, who considered it “very near” R. schlechtendalii
Weihe ex Link (Rogers & Rogers 1898), an opinion Focke was later to refine to ““maybe a form of
schlechtendalii. Not typical”, when G. C. Druce sent him a specimen he had brought back from Sark
in 1906 (now in OXF). Although Rogers initially placed the plant under that name (Derrick 1898,
Rogers 1899), within a year he had decided it was best regarded instead as a strong and highly
glandular form of R. macrophylloides Genev. (Rogers & Rogers 1898). It was not that species,
however, as Riddelsdell was eventually to note on one of Rogers’ sheets (herb. Barton &
Riddelsdell 8743: BM) after study of Genevier’s type and description. Nor was it R. adscitus Genev.,
as Watson first supposed on re-examining Druce’s specimen in 1931 (a determination he
subsequently abandoned for R. boraeanus Genev.) — although it is certainly closely related to that
species, as A. Newton remarked of a further Sark specimen that was submitted to him by Lady
Anne Brewis in 1973 (now in STP).

It has always seemed likely that the bramble would prove to be on the neighbouring French
mainland; but as the Rubus flora of that area has not been much studied, it might well be one of the
many species there as yet undescribed. Fortunately, however, the latter has not proved to be the
case.

In 1987, on a first reconnaissance of the Cherbourg area, I came across a bramble growing in
plenty round the north foot of Le Roule, the high crag which overlooks that city, that proved to
match a panicle in BM collected by Corbiére in 1889 from what are today the outskirts of Cherbourg
and later distributed under the name Boulay bestowed on the bramble in his honour, R. corbieri.
Corbiére tended to interpret Rubus species over-broadly, and his determinations cannot be
accepted uncritically; but the very distinctiveness of this particular plant taken together with his
published description of R. corbieri (Corbiére 1894) leave no room for doubt in this instance.
Moreover, there is a further Cherbourg specimen of his so named in P (Association rubologique no.
894) that clearly belongs to the same entity.

Subsequently I discovered that this Cherbourg plant was identical with a specimen in my
herbarium that I had collected in Guernsey in 1978, in a deserted garden in the middle of St Peter
Port. Because R. corbieri is rather variable, however, it was some time before I began to suspect that
this might also be the bramble which occurs more widely in the north of that island and in such ~
profusion on Sark. Fuller exploration of the north of the Cotentin Peninsula in 1991, bringing at the
same time wider acquaintance with the variation exhibited by the species in the field, showed
conclusively that that suspicion was well-founded. R. corbieri proved to occur conspicuously all
along the coast to the west of Cherbourg (though apparently absent to the east), rising to abundance
in the vicinity of Cap de la Hague, the headland facing Alderney, the northernmost of the Channel
Isles. The comparable abundance of the species on Sark and its presence in much smaller quantity
on Guernsey, which is situated farther out to the west, thus represents a natural prolongation, and
gradual tailing-off, of its Cotentin range. I did not have an opportunity of following the coastline
south from Cap de la Hague, but the discovery of a patch of R. corbieri just outside Carteret, some
40 km in that direction, suggests that the species may similarly prove to extend down to there. Most
unusually for a Rubus species, it would appear to prefer the proximity of the sea, the coarseness of
its foliage presumably rendering it well fitted to the rigours of that environment. Even so it
noticeably avoids the most exposed situations, tending to occur on banks offering some shelter from
the wind.

Unexpectedly, R. corbieri also turns out to occur in England. In the autumn of 1990, while going
through OXF in search of Channel Isles Rubus material, I came across an unmistakable specimen of
it collected on Corfe Common in south-east Dorset, v.c. 9, by L. Cumming in 1916 and distributed
through both of the exchange clubs that year as a white-flowered form of R. boraeanus (as
determined by Rogers). The sheet bears a later determination by Watson, confirming that name. In
one of the published notes on the gathering (Rogers 1918) the plant was described as locally
abundant in the locality in question, which encouraged the belief that it might still be present there;
and a visit to the Common the following July quickly revealed that not only was that the case, but

SHORT NOTES 133

also that the population remains very extensive. Later the same day the species was encountered ina
second locality in quite another part of Purbeck, along a lane crossing Studland Heath (GR SZ/
01.84 and SZ/02.84). E. F. Linton has also since been found to have collected it in 1892 in a locality
nearly midway from Corfe to Swanage, the specimen (BM) having been dismissed by Rogers and R.
P. Murray as R. bloxamii X vestitus. In CGE, too, there turns out to be a gathering of it (no. 64/368)
by B. A. Miles as recently as 1964 from the south-west part of Rempstone Heath (GR SZ/983.840),
in this case with no name hazarded. Possibly, therefore, the species may prove widespread in the
south-east corner of v.c. 9. Although that has been relatively well worked by batologists in the past,
the superficial resemblance of R. corbieri to the common R. bloxamii (Bab.) Lees could well have
led to its being passed over as the latter.

Work on the Rubus flora of Hampshire, v.cc. 11 and 12, and the Isle of Wight, v.c. 10, in recent
years has been sufficiently intensive to render it unlikely that so obtrusive a species has escaped
notice in those neighbouring counties. Floristically, in any case, the affinity of their Rubus floras
with that of the French coastal areas directly opposite appears to be but slight. At least the north end
of the Cotentin Peninsula has a Rubus flora with a markedly more western facies than theirs, which
makes the presence of R. corbieri in Dorset more congruous and Devon, v.cc. 3 and 4, the likeliest
further English county in which the species may occur.

The descriptions of R. corbieri by Corbiére (1894) and by Boulay (1900) — the latter of whom
reduced the species to a ‘Forme’ of R. multifidus Boulay & Malbranche (a common Normandy
bramble) and that in turn to a subspecies of R. menkei Weihe -— are. sufficiently detailed and
accessible to render providing a further one unnecessary. It is nevertheless worth emphasising that
the species, while unambiguously a member of Series Vestiti (Focke) Focke, varies considerably in
the quantity of stalked glands present as well as in the size of the rachis prickles. In addition, the
flowers tend to be pinkish on first opening or on plants growing in shade.

Sudre in his Batotheca Europaea, fasc. 10, no. 487, relegated R. corbieri to the synonymy listed on
the printed label of the bramble he distributed as R. fuscus subsp. retrodentatus (Mueller & Lef.)
Sudre. The latter, however, a specimen of Questier’s from dép. Valois, is quite a different plant and
it would seem that Sudre had a mistaken idea of the Cherbourg species.

Representative material from Dorset, the Channel Isles and the North Cotentin has been
deposited in BM.

REFERENCES

Bou ay, N. (1900). Rubus, in Rouy, G. & Camus, E.-G. Flore de France, vol. 6. Asniéres & Paris.

CorsikErE, L. (1894). Nouvelle Flore de Normandie. Caen.

Derrick, G. T. (1898). Additions to the Sark flora. Guernsey Soc. Nat. Sci. Rep. & Trans. 1897: 166-169.

Rocers, W. M. (1899). The Rubi of Guernsey and Sark. Guernsey Soc. Nat. Sci. Rep. & Trans. 1898: 296-297.

Rocers, W. M. (1918). Rep. Watson bot. Exch. Club 3: 14.

Rocers, W. M. & Rocers, F. A. (1898). On the Rubi and Rosae of the Channel Islands. J. Bot., Lond. 36: 85-
90.

D. E. ALLEN
Lesney Cottage, Middle Road, Winchester, Hants., SO22 5EJ

A NEW SPIRAL VARIANT OF JUNCUS EFFUSUS L. (SUNCACEAE)

The Spiral Rush of gardens, Juncus effusus L. cv. ‘Spiralis’, was discovered as a single plant in
Northern Ireland by David Bishop, Curator of the Botanic Garden, Belfast in 1869. It was brought
into cultivation and has continued as a curiosity in gardens since then. It differs from the ordinary J.

_ effusus in that the stems spiral upwards, often completing six to eight turns. James McNab, Curator

——oe se

at that time of the Royal Botanic Garden, Edinburgh and a most intelligent experimenter, obtained
plants of the Irish discovery and sowed seeds from them. He reported that the seedlings all showed

_ the spiral character (McNab 1873). There seems to be no other record of this variety in the wild.

In the autumn of 1988, I noticed on rough croft grazing above Big Sand, Gairloch, W. Ross (v.c.
105), areas of Juncus which at first sight had every appearance of having been flattened by animals
lying on the plants. Some of the colonies were 1.3-1.9 m across. Closer examination showed that

134 SHORT NOTES

many of the stems were gently spiral and that all of them were only suberect in contrast to the strictly
erect habit of normal J. effusus growing nearby. Plants were brought into cultivation. Seed from the
‘spiral’ plants at Big Sand germinated easily and all the seedlings were spiral and suberect. The Irish
‘Spiralis’ differs from the Scottish in that the stems in the Irish are quite erect and are much more
obviously spiral six to eight turns as against two to three.

There are many other populations in N.W. Scotland between Plockton and Dundonnell: on stfe
village green in Plockton, at Flowerdale, Gairloch, at Red Point, Gairloch, at Cove, Poolewe, on
the Inverewe peninsula, Poolewe, along the roadside by Loch Tollie between Gairloch and
Poolewe, at Slaggan west of Mellon Udrigle and by the roadside at Dundonnell. It has not been
found in the eastern or central Highlands, although once known it can be spotted from a moving car.
It usually occurs mixed with normal plants and the contrast between the two is quite striking.

There seem to be no previous records of this spiral rush on mainland Britain but there are
specimens in E from the Northern Isles: Orkney, 1906, M. Spence; Ramsdale, Orphir, 1927, H. H.
Johnston; Flotta, Orkney, 1932, J. Sinclair and all are similar to the West Ross plants rather than the
Irish. I have also seen a photograph of an Orkney specimen collected by Linton in LIV. However, in
1991, it was evident that the variety had a wide and certainly long established distribution in the
Outer Hebrides where I found it on the Island of Killegray (Harris) on the Shiant and Monach Isles
just west of Loch Boisdale (Uist) and on Mingulay (Barra).

In view of its considerable occurrence as wild populations over a considerable area of N.W.
Scotland it seems appropriate to treat it as a distinct variety of Juncus effusus:

Juncus effusus L. var. suberectus D. M. Henderson, var. nov.
Hovotypus: Big Sand, Gairloch, W. Ross, v.c. 105, on rough croft land with var. effusus, 6
December 1988, D. M. Henderson (E).

A varietate typica culmis leniter spiralibus suberectis differt.

REFERENCES
McNags, J. (1873). Remarks on Juncus effusus spiralis and varieties of ferns. Trans. Proc. bot. Soc. Edinb. 12:
502-504.

D. M. HENDERSON
Inverewe House, Poolewe, Wester Ross, IV22 2LQ -

CONTRIBUTIONS TO A CYTOLOGICAL CATALOGUE OF THE BRITISH AND IRISH
FLORA, 2

The only recent, published summary of chromosome counts made on native material of British and
Irish vascular plants is contained in Clapham et al. (1987). The source of many of these counts,

however, is unlocalised material or untraceable. The present series of notes is intended to |

contribute to a properly documented cytological account of our flora (Wentworth ef al. 1991). We |
present here chromosome counts of 102 flowering plant species from 122 populations. Only one
plant from each population was studied, except where noted. All counts were made from squashes |
of root-tips, except where noted; supernumerary chromosomes are designated by the suffix ‘S’.
Voucher specimens have been placed in LTR.

Achillea millefolium L., 2n = 54: Caerns., v.c. 49, E. of Sarn, SH/24.32; Westmorland, v.c. 69, near
Ulpha, c.5 km N.E. of Lindale, SD/44.81.

Agrimonia eupatoria L., 2n = 28: W. Lancs., v.c. 60, Silverdale, Gait Barrows N.N.R., SD/48.77;
W. Lancs., v.c. 60, Warton Crag, R.S.P.B. nature reserve, SD/49.72. 4

Agrimonia procera Wallr., 2n = 56: W. Lancs., v.c. 60, Warton Crag, R.S.P.B. nature reserve, SD/ |
49.72. .

Ajuga reptans L., 2n = 32: W. Lancs., v.c. 60, near Forton, SD/48.53.

Allium vineale L. var. vineale, 2n = 32 + 0-2S: W. Lancs., v.c. 60, near Carnforth, SD/49.71.

SHORT NOTES 135

Anagallis tenella (L.) L., 2n = 22: Caerns., v.c. 49, Lleyn peninsula, W. end, SH/2.3.

Angelica sylvestris L., 2n = 22: W. Lancs., v.c. 60, near Lancaster, SD/47.59.

Anthriscus sylvestris (L.) Hoffm., 2n = 16: Leics., v.c. 55, Rutland, 4 km S. of Oakham, near
Brooke, SK/85.06.

Apium graveolens L., 2n = 22: W. Lancs., v.c. 60, Potts Corner, SD/41.57.

Apium nodiflorum (L.) Lag., 2n = 22: Dorset, v.c. 9, near Sydling St Nicholas, ST/63.00.

Arum maculatum L., 2n = 56: Leics., v.c. 55, Swithland Wood, SK/53.12.

Atriplex portulacoides L., 2n = 36: W. Lancs., v.c. 60, Potts Corner, SD/41.57.

Barbarea vulgaris R.Br., 2n = 16: Caerns., v.c. 49, Tudweiliog, 1 km to W. on Tyd’dyn road, SH/
22.36.

Blackstonia perfoliata (L.) Hudson, 2n = 40: Co. Durham, v.c. 66, coastal cliffs between Blackhall
and Crimdon, NZ/47.38.

Centaurea scabiosa L., 2n = 20 + 3-8S: Co. Durham, v.c. 66, coastal cliffs between Blackhall and
Crimdon, NZ/47.38.

Chelidonium majus L., 2n = 12: Caerns., v.c. 49, Llaniestyn, SH/26.33.

Chrysanthemum segetum L., 2n = 18: Caerns., v.c. 49, N. of Botwnnog, overlooking Cellar Farm,
SH/26.32.

Chrysosplenium oppositifolium L., 2n = 42: Derbys., v.c. 57, Dovedale, SK/14.53.

Cichorium intybus L., 2n = 18: Co. Durham, v.c. 66, coastal cliffs between Blackhall and Crimdon,
NZ/47.38.

Cirsium vulgare (Savi) Ten., 2n = 68: Caerns., v.c. 49, Botwnnog, above Cellar Farm, SH/26.32.

Clematis vitalba L., 2n = 16: Surrey, v.c. 17, Silent Pool, near Shere, TQ/06.48.

Conyza canadensis (L.) Crong., 2n = 18: Leics., v.c. 55, Leicester University Botanic Garden
glasshouse (spontaneous weed), SK/61.01.

Corylus avellana L., 2n = c. 22: Leics., v.c. 55, Swithland Wood, SK/53.12.

Deschampsia setacea (Hudson) Hackel, 2n = 14: Caerns., v.c. 49, Cilan, near Abersoch, SH/29.24.

Digitalis purpurea L., 2n = 56: W. Lancs., v.c. 60, Lancaster University grounds, SD/48.57.

Euonymus europaeus L., 2n = 32: Westmorland, v.c. 69, Brigsteer Park, 2-5 kmS. of Brigsteer, SD/
48.87.

Eupatorium cannabinum L., 2n = 20: Dorset, v.c. 9, Hartland Moor N.N.R., c. 4 km S.E. of
Wareham, SY/96.85; W. Norfolk, v.c. 28, Foulden Common, TL/76.99; W. Lancs., v.c. 60,
near Carnforth, SD/49.71.

Filipendula ulmaria (L.) Maxim., 2n = 14: Leics., v.c. 55, Swithland Woods, SK/53.12; W. Lancs.,
v.c. 60, Lancaster University grounds, SD/48.57.

Galeopsis bifida Boenn., 2n = 32: Caerns., v.c. 49, E. of Sarn, SH/24.32; Caerns., v.c. 49, N. of
Botwnnog, overlooking Cellar Farm, SH/26.32.

Galium cruciata (L.) Scop., 2n = 22: W. Lancs., v.c. 60, Potts Corner, SD/41.57.

Galium mollugo L., 2n = 44: W. Cornwall, v.c. 1, Goonhilly Earth Station, SW/73.21; E. Cornwall,
v.c. 2, Rame, near church, SX/42.49.

Galium odoratum (L.) Scop., 2n = 44: Westmorland, v.c. 69, Brigsteer Park, 2-5kmS. of Brigsteer,
SD/48.87.

Geranium dissectum L., 2n = 22: W. Lancs., v.c. 60, near Lancaster, SD/46.62.

Geranium lucidum L., 2n = 40: Westmorland, v.c. 69, near Meathop Moss, c.3kmN.E. of Lindale,
SD/44.81.

Geranium pratense L., 2n = 28: Derbys., v.c. 57, Milldale, SK/14.55; Co. Durham, v.c. 66,2 km N.
of Barnard Castle, NY/05.18.

Geranium sanguineum L., 2n = 84: Co. Durham, v.c. 66, coastal cliffs between Blackhall and
Crimdon, NZ/47.38.

Geum urbanum L., 2n = 42: Derbys., v.c. 57, Milldale, SK/14.55.

Glaux maritima L., 2n = 30: W. Lancs., v.c. 60, W. of Cockerham, near Bank Houses, SD/42.53;
W. Lancs., v.c. 60, Potts Corner, SD/41.57.

Glechoma hederacea L., 2n = 36: Leics., v.c. 55, Rutland, Brooke, by the church, SK/85.06; W.
Lancs., v.c. 60, Silverdale, E. of Thrang End Farm, SD/49.77.

_ Heracleum sphondylium L., 2n = 22: W. Lancs., v.c. 60, Lancaster University grounds, SD/48.57.
_ Hippuris vulgaris L., 2n = 32: W. Lancs., v.c. 60, near Forton, by canal, SD/48.53.
_ Hydrocotyle vulgaris L., 2n = 96: Caerns., v.c. 49, Llanbedrog, valley W. of Penarwel, SH/32.32.

136 SHORT NOTES

Hypericum hirsutum L., 2n = 18: W. Lancs., v.c. 60, near Carnforth, SD/50.70; Westmorland, v.c.
69, Ravens Lodge, S.E. edge of Whitbarrow, base of Whitescar, SD/46.85.

Hypericum perforatum L., 2n = 32: Surrey, v.c. 17, Silent Pool, near Shere, TQ/06.48.

Lathyrus nissolia L., 2n = 14: W. Lancs., v.c. 60, near Lancaster University, SD/48.56.

Limonium britannicum Ingrouille subsp. celticum Ingrouille var. pharense Ingrouille, 2n = 35:
Caerns., v.c. 49, Bardsey Island, E. of Pen Cristin, Ogof y Gaseg, SH/12.21.

Lotus corniculatus L., 2n = 24: W. Lancs., v.c. 60, Gait Barrows N.N.R., near Silverdale, SD/
48.77.

Luzula pilosa (L.) Willd., 2n = 62: Co. Waterford, v.c. H6, Nier Valley, S/2.1.

Lycopus europaeus L., 2n = 22: Caerns., v.c. 49, Pwllheli, SH/37.34.

Lysimachia nemorum L., 2n = 16: Leics., v.c. 55, Swithland Wood, SK/53.12; W. Lancs., v.c. 60,
near Barkin Bridge, SD/60.63.

Lythrum portula (L.) D. A. Webb, 2n = 10: Caerns., v.c. 49, Rhos Botwnnog, SH/26.32.

Medicago sativa L. subsp. sativa, 2n = 32: Co. Durham, v.c. 66, coastal cliffs between Blackhall and
Crimdon, NZ/47.38.

Myosotis sylvatica Hoffm., 2n = 18: W. Lancs., v.c. 60, near Yealand Conyers, SD/50.74.

Oenanthe crocata L., 2n = 22: W. Lancs., v.c. 60, Lancaster University grounds, SD/48.57.

Oenanthe lachenalii C. C. Gmelin, 2n = 22: W. Lancs., v.c. 60, Potts Corner, SD/41.57.

Parietaria judaica L., 2n = 26: W. Cornwall, v.c. 1, Newbridge, near St Just, SW/42.31; W. Lancs.,
v.c. 60, near Silverdale, SD/45.74.

Paris quadrifolia L., 2n = 20: W. Lancs., v.c. 60, Gait Barrows N.N.R., near Silverdale, SD/
48.77.

Pedicularis sylvatica L. subsp. sylvatica, 2n = 16: W. Cornwall, v.c. 1, between Penzance and St
Just, SW/39.31. |

Phleum arenarium L., 2n = 14: Caerns., v.c. 49, Abersoch dunes, N. part of Porth Fawr, SH/31.27.

Pinguicula lusitanica L., 2n = 12: Dorset, v.c. 9, Slepe Heath, c. 2-5 kmS.E. of Wareham, SY/94.86
(count made on integumentary tissue).

Plantago lanceolata L., 2n = 12: W. Lancs., v.c. 60, Lancaster University grounds, SD/48.57.

Plantago media L., 2n = 24: Dorset, v.c. 9, Fontmell Down, c. 5 km S.S.E. of Shaftesbury, ST/
88.18; W. Lancs., v.c. 60, near Lancaster, by dismantled railway, SD/46.62; Westmorland, v.c.
69, Helsington Barrows, c. 3 km S.E. of Kendal, SD/48.89.

Primula elatior (L.) Hill, 2n = 22: Cambs., v.c. 29 (no further details known).

Primula veris L., 2n = 22: Co. Durham, v.c. 66, coastal cliffs between Blackhall and Crimdon, NZ/
47.38.

Prunella vulgaris L., 2n = 28: Co. Durham, v.c. 66, coastal cliffs between Blackhall and Crimdon,
NZ/48.38; W. Lancs., v.c. 60, Lancaster University grounds, SD/48.57. .

Pulicaria dysenterica (L.) Bernh., 2n = 18: Dorset, v.c. 9, Hartland Moor N.N.R., c.4kmS.E. of
Wareham, SY/96.85.

Quercus petraea (Mattuschka) Liebl., 2n = 24: Leics., v.c. 55, Swithland Wood, SK/53.12.

Quercus robur L., 2n = 24: E. Norfolk, v.c. 27, Badley Moor, NT/00.11.

Radiola linoides Roth, 2n = 18: Caerns., v.c. 49, Cwmistir, near Edeyrn, SH/25.38.

Ranunculus auricomus L., 2n = 32: W. Lancs., v.c. 60, near Yealand Conyers, SD/50.74.

Ranunculus bulbosus L., 2n = 16: W. Lancs., v.c. 60, Potts Corner, SD/41.57.

Ranunculus flammula L. subsp. flammula, 2n = 32: Westmorland, v.c. 69, Killington, SD/61.88.

Ranunculus omiophyllus Ten., 2n = 32: Caerns., v.c. 49, Garn Fadryn, SH/27.35.

Ranunculus sceleratus L., 2n = 32: W. Lancs., v.c. 60, Potts Corner, SD/41.57.

Rhamnus catharticus L., 2n = 24: W. Norfolk, v.c. 28, Foulden Common, TL/76.99.

Ribes nigrum L., 2n = 16: Leics., v.c. 55, Swithland Wood, SK/53.12.

Ribes uva-crispa L., 2n = 16: W. Lancs., v.c. 60, Silverdale, E. of Thrang End Farm, SD/49.77.

Rubus chamaemorus L., 2n = 56: Derbys., v.c. 57, Snake Pass summit, N. side of car park, SK/
08.92.

Salsola kali L., 2n = 36: N. Somerset, v.c. 6, near Berrow, ST/28.53.

Sambucus nigra L., 2n = 36: Leics., v.c. 55, Swithland Wood, SK/53.12.

Sanguisorba minor Scop. subsp. minor, 2n = 28: Westmorland, v.c. 69, Brigsteer Park, 2-5 km S. of |}
Brigsteer, SD/48.87.

Sanguisorba officinalis L., 2n = 56: W. Lancs., v.c. 60, near Aldcliffe, SD/46.60.

SHORT NOTES 137

Sanicula europaea L., 2n = 16: Leics., v.c. 55, Swithland Wood, SK/53.12; Westmorland, v.c. 69,
Brigsteer Park, 2-5 km S. of Brigsteer, SD/48.87.

Scabiosa columbaria L., 2n = 16: Westmorland, v.c. 69, Helsington Barrows, c. 3 km S.E. of
Kendal, SD/48.89.

Scrophularia nodosa L., 2n = 36: W. Lancs., v.c. 60, near Lancaster, SD/47.59.

Sedum telephium L., 2n = 24: W. Lancs., v.c. 60, Gait Barrows N.N.R., near Silverdale, SD/47.77.

Senecio viscosus L., 2n = 40: W. Lancs., v.c. 60, near Carnforth, SD/49.71.

_ Solanum dulcamara L., 2n = 24: W. Lancs., v.c. 60, near Aldcliffe, close to estuary of River Lune,

SD/45.60.

_ Spiranthes spiralis (L.) Chevall., 2n = 30: Caerns., v.c. 49, Abersoch Bay, below Marchros, on

Abersoch golf-links, SH/31.26.

| Stachys officinalis (L.) Trev., 2n = 16: W. Cornwall, v.c. 1, Chyenhal, SW/45.27.
- Stellaria holostea L., 2n = 26: Leics., v.c. 55, Swithland Wood, SK/53.12.

\
1 :

.
|
|

Tanacetum vulgare L., 2n = 18: W. Lancs., v.c. 60, River Lune estuary, near Lancaster, SD/45.62.

_ Taxus baccata L., 2n = 24: Leics., v.c. 55, Swithland Wood, SK/53.12.

Teucrium scorodonia L., 2n = 32: Cheviot, v.c. 68, Ingram, River Breamish, NU/01.16.
' Triglochin palustris L., 2n = 24: E. Norfolk, v.c. 27, Badley Moor, NT/00.11, (three plants
counted).
_Tussilago farfara L., 2n = 60: W. Lancs., v.c. 60, Lancaster University grounds, SD/48.57;
W. Lancs., v.c. 60, near Carnforth, SD/50.71.
) Typha latifolia L., 2n = 30: W. Lancs., v.c. 60, Lancaster University grounds, SD/48.57.
Valeriana dioica L., 2n = 16: Leics., v.c. 55, Swithland Wood, SK/53.12.
_Verbascum nigrum L., 2n = c. 30 + 4 (3-7)S: S. Hants., v.c. 11, near Winchester, SU/4.2.
Veronica serpyllifolia L. subsp. serpyllifolia, 2n = 14: Caerns., v.c. 49, Bardsey Is., Chapel garden,
feesH/12 22.
Vicia sepium L., 2n = 14: W. Lancs., v.c. 60, above Saltmire Bridge, SD/51.75; Westmorland, v.c.
69, Whitbarrow (S.E. edge), Raven’s Lodge, underneath White Scar, SD/46.85.
\Wahlenbergia hederacea (L.) Reichenb., 2n = 36: cultivated plant, originally from S. Devon, v.c. 3,
Dartmoor, Newbridge near Hone (W. of Bovey Tracy), SX/71.70.

ACKNOWLEDGMENTS

. |

from their reserves.

_ We should like to thank English Nature and the R.S.P.B. for permission to collect plant material

;
’
‘A :

REFERENCES

i CLAPHAM, A. R., Tutin, T. G. & Moore, D. M. (1987). Flora of the British Isles, 3rd ed. Cambridge.
_ Wentworth, J. E., BaiLey, J. P. & Gorna_t, R. J. (1991). Contributions to a cytological catalogue of the
British and Irish flora, 1. Watsonia 18: 415-417.

P. M. HoLiincswortH, R. J. GORNALL & J. P. BAILEY
Botany Department, The University, Leicester, LEI] 7RH

THE ROSA HERBARIUM OF A. H. WOLLEY-DOD

¢ As far as is known, until last year Wolley-Dod’s Rosa collection in BM had never been examined
_vully, and certainly had never in its entirety been examined critically, since Wolley-Dod himself. A
_ 2w of the specimens have been remounted, but most are on Wolley-Dod’s original sheets. Large

Jumbers are unmounted, many of them between old and yellow sheets of newspaper dating from

ie 1920s. The author of this note was asked to go through the collection and label the specimens in
je light of modern taxonomic opinion. He received the collection on loan early in 1990, and did

{
‘|
4
|

138 SHORT NOTES

most of the routine work himself, with the assistance of G. G. Graham where consultation or a
second opinion seemed desirable.

The collection contains 3091 specimens. Of these, over 100 were indeterminate, in some cases
because the specimens had deteriorated, but mainly because they were too immature or too scrappy
for determination. A few in the latter category carried a caustic comment from Wolley-Dod, and as

he had not attempted identification one wonders why he bothered to keep them. Pasted on to many

of the sheets are comments by Sudre, whose reliability Wolley-Dod came to distrust, and more
valuable comments by R. Keller, who wrote in a crabbed hand indiscriminately in English, German,
French or Latin.

Besides the large numbers of specimens collected by Wolley-Dod himself, he had acquired
considerable collections from others, many of them illustrious names in the history of the study of
Rosa. These, in alphabetical order, are as follows: C. Bailey, J. G. Baker, W. Barclay, E. B.
Bishop, C. E. Britton, R. & M. Corstorphine, E. M. Day, G. Claridge Druce, Rev. A. Ley, Rev. E.
F. Linton, Rev. W. R. Linton, Rev. E. S. Marshall, Rev. H. J. Riddelsdell, Rev. W. Moyle Rogers,
I. M. Roper and W. A. Shoolbred. There are also a few specimens from other well-known botanists
such as Rev. A. Bloxam and A. R. Horwood.

Study of the localities from which collections were made would give a fair indication of the
distribution of species from north to south. Otherwise, distribution maps compiled from the records
would tend rather to show the chosen hunting grounds of the main collectors. Wolley-Dod collected
mainly in Devon, Surrey, Derbyshire, Cheshire and the Lake District; Baker from around Thirsk in
Yorkshire; Druce from Oxfordshire; Ley from Gloucestershire, Herefordshire and parts of Wales:
Moyle Rogers from Devon and Gloucestershire. Other collectors also appear to have concentrated
mainly on these same regions, with a plethora of records from Surrey and scattered records from
elsewhere in England. Most of the collections in Scotland are from the Perthshire vice-counties,
with outlying records by E. S. Marshall from Westerness and Wester Ross, Mrs Corstorphine from
Angus and H. Halcro Johnston from Hoy and Mainland, Orkney. There are only about 30 records
from Ireland, mostly collected by C. H. Waddell in County Down. The Channel Islands are
represented by two specimens from Jersey.

There are a few specimens from Europe. The main interest of these lies in the only two specimens
of Rosa elliptica Tausch, both from France. These show that Wolley-Dod knew and recognised this
species, but presumably had not found it in Britain, which seems to corrobrate the assumption that
this is not a native British species (Graham & Primavesi 1990).

Allowing for the somewhat patchy distribution of records described above, there appears in
general to have been little change in frequency of Rosa species and their hybrids throughout the

country between the period during which the collection was made and the present day. Naturally, —

there are losses in some areas, principally where there has been expansion of the larger towns and
cities. There are, for instance, a number of records which at first sight appear astonishing nowadays
because the collections were made in what are now solidly built-up areas of Greater London. Only
one species shows a marked change in frequency. This is Rosa agrestis Savi. There are about 20
specimens of this species in Wolley-Dod’s collection, from scattered localities all over the southern
part of England. Nowadays this appears to be a very rare species in England, though it is still
frequent in parts of Ireland. What has caused this decline in frequency is not known. The other two
sweet briar species, R. rubiginosa L. and R. micrantha Borrer ex Smith, appear to have undergone
little or no change in frequency since Wolley-Dod’s time.

In his Revision of the British Roses, Wolley-Dod (1930-31) repeatedly refers to this herbarium,
but he did not re-label the specimens to conform with the nomenclature of that work. The names are
those of the multiplicity of species described in his earlier works (Wolley-Dod 1908, 1910). The
research which he undertook is reflected in additional comments attached to the sheets. As well as
those of Sudre and Keller already mentioned, there are Wolley-Dod’s own type-written or hand-
written comments, and cuttings from Botanical Exchange Club reports. One cannot help feeling
that in the light of modern opinion the work undertaken to elaborate all the Groups, varieties and
forms described in Revision of the British Roses was an unprofitable expenditure of time that could
have been more usefully employed. Indeed, Wolley-Dod himself later expressed doubt and
dissatisfaction, stating that rarely in the field did one find specimens which corresponded closely or
even remotely with the author’s description of a named variety (Wolley-Dod 1936). Keller was more
nearly on the right lines, as he frequently suggested the possibility of hybridity. It is a pity that

SHORT NOTES 139

Wolley-Dod did not pay more attention to Keller’s comments, but he would not recognize a Rosa
hybrid unless the hips showed complete or partial sterility. In the course of the recent examination
of this herbarium, we were able to label many specimens as definite hybrids between two species, a
large number where there was plainly slight introgression of another named species, some where
there was introgression of another indeterminate species, and a few which were such a mixture as to
be impossible to determine. As regards the latter category it is not worth spending time on them;
there are plenty of roses which can be determined and recorded. Many modern taxonomists allow
inclusion of the second and third categories above as permissible variants of a named species (Stace
1975; Webb 1951). There are still unsolved problems in Rosa taxonomy and nomenclature, but the
appalling difficulties which Wolley-Dod faced, and with his researches laid the foundations for later
work, have now largely been overcome.

REFERENCES

GraHaM, G. G. & PrimaveEs!, A. L. (1990). Notes on some Rosa taxa recorded as occurring in the British Isles.
Watsonia 18: 119-124.

Stace, C. A. (1975). Hybridization and the flora of the British Isies, p. 3. London.

Wess, D. A. (1951). Hybrid plants in Ireland. Jr. Nat. J. 10: 201-204.

~ Wo ttey-Dop, A. H. (1908). The subsection Eu-caninae of the genus Rosa. J. Bot., Lond. 46 (Suppl.): 1-110.

Wo tey-Dop, A. H. (1910). The British roses (excluding Eu-car:nae). J. Bot., Lond. 48 (Suppl.): 1-141.

Wo tey-Dop, A. H. (1930-31). A revision of the British roses. J. Bot., Lond. 68 (Suppl.): 1-16 & 69 (Suppl.):
17-111.

Wo LLeEy-Dop, A. H. (1936). Some rose notes. Rep. botl Soc. Exch. Club Br. Isl. 11: 68-81.

A. L. PRIMAVESI
Ratc‘iffe College, Syston, Leicester LE7 8SG

ADDITIONAL RECORDS OF SORBUS LANCASTRIENSIS E. F. WARBURG
(ROSACEAE)

Following documentation of the known sites of Sorbus lancastriensis E. F. Warburg (Rich &

Baecker 1986), a number of additional populations have come to our attention, including two on

Silurian rock, two about Grange-over-Sands, and one wes: of the Leven Estuary. This note lists the

additional sites (we have visited all new localities in June 1991), and updates our previous list. Full

details have again been lodged with the vice-county recorders, English Nature, the local Naturalists’

Trusts and the Biological Records Centre, Monks Wood.

WW Lanics:. v.c. 60:

Gait Barrows N.N.R. (SD/4.7). Three plants have now been found, with Sorbus aria, confirming
the Ratcliffe (1977) record queried by Rich & Baecker (1986).

Wesimorland, v.c. 69:

Farleton Knott, one plant on top of cliff (SD/5.7), and twe plants on cliff face, about 300 m north of
the above plant (SD/5.8). These confirm the Ratcliffe (1977) record not found in 1982.

Furness, v.c. 69b:

Birkrigg Common (SD/2.7). “One small tree in limestone pavement on summit’, F. L. Woodman
& K. A. Gunning, 27 May 1982 (LANC). This is the first known site west of the Leven Estuary; we
have been unable to re-find the plant.

Kirkhead (SD/3.7). About 30-40 plants in scrubby woodland on the west side, with a few Sorbus
rupicola. This is probably the same locality as found by C. Bailey in 1881 on ‘exposed limestone
ridges near Wray’s Holme Tower” (CGE, BM).

Old Park Wood (SD/3.7). We under-estimated the population size of this site; there are over 100
plants mixed with a smaller number of S. rupicola plants.

Grange-over-Sands (SD/4.7). One plant by B5277 at west edge of golf course.

Witherslack and Yewbarrow (SD/4.8). Isolated plants occur on roadsides, in hedges and on
limestone rocks in at least six separate localities around the villages. One plant, to the N.E. of
Witherslack, is in a shady hedge on Silurian baserock, a very atypical habitat.

140 SHORT NOTES

Poolbank (SD/4.8). One small tree on woodland edge by roadside, first found by G. Halliday in
1974 (LANC). This is a second site on the Silurian slate; the soil pH is about 6-6-5, equivalent to
the pH of many of the limestone soils on which it occurs.

Latterbarrow Nature Reserve (SD/4.8). At least four plants on the reserve, and about ten on the
S.W. edge in woodland above the old A590. This is almost certainly the same locality as plants
collected in 1937 by E. F. Warburg near The Derby Arms (BM).

Currently, there are thus approximately 2000 plants known from about 35 sites in eight 10-km
squares, with one extinct population.
ACKNOWLEDGMENTS

We would like to thank the Keepers of BM, K and LANC for access to specimens, and Eimear Nic
Lughadha for help in the field.

REFERENCES

RATCLIFFE, D., ed. (1977). A nature conservation review, vol. 2. Cambridge.
Ricu, T. C. G. & BAEcKER, M. (1986). The distribution of Sorbus lancastriensis E. F. Warburg. Watsonia 16: 83—
85.

T. C. G. Rico & M. BAECKER
24 Lombardy Drive, Peterborough, PE1 3TF

Watsonia, 19, 141-154 (1992) 141

Plant Records

Records for publication must be submitted to the appropriate Vice-county Recorder (see B.S$.B.J. Year Book
for 1992), 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: Ist 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), from which
the species’ numbers are taken. Taxonomy and nomenclature follow the New Flora of the British Isles by C. A.
Stace (1991), except for the genera Rubus and Hieracium. The Ordnance Survey national grid reference follows
the habitat and locality; the 100-km square is given in alphabetical form at the start of the grid reference rather
than as numerical co-ordinates. 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 are field records if no other source is
stated.

Records from the following vice-counties are included in the text below: 1, 2, 4,9, 11-15, 17, 22, 25, 26, 28, 29,
35, 36, 38, 39, 41-50, 52, 58, 59, 61-65, 67-71, 73, 75, 77, 79-81, 88, 93, 98, 99, 102, 111.

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.

+b x

1/com. DIPHASIASTRUM COMPLANATUM (L.) Holub *67,S. Northumb.: With D. alpinum under
Calluna, William’s Cleugh, NY/639.991. G. A. & M. Swan, 1988, herb. G.A.S., det. A. C. Jermy as
morphotype decipiens.

4/9 X 5. EQUISETUM ARVENSE L. X E. FLUVIATILE L. *46, Cards.: Disused railway verge,
Pendinas, SN/581.801. A. O. Chater, 1991, NMW.

+13/cor. BLECHNUM CORDATUM (Desv.) Hieron. *98, Main Argyll: Rock exposures, Craignish
Castle, NM/773.015. A. McG. Stirling & B. H. Thompson, 1991, BM, det. A. C. Jermy. Present
since late 1970s.

15/8. ASPLENIUM SEPTENTRIONALE (L.) Hoffm. *43, Rads.: S.-facing rock face, Yatt Wood,
Dolyhir, SO/24.58. D. C. Boyce, D. R. Drewett & R. G. Woods, 1991.

21/6 X 7. DRYOPTERIS CARTHUSIANA (Villars) H. P. Fuchs xX D. pitatata (Hoffm.) A.
Gray *80, Roxburghs.: Birch wood, Din Moss, Hoselaw, NT/805.316. R. W. M. Corner, 1991,
BM, det. A. C. Jermy.

21/8. DRYOPTERIS AEMULA (Aiton) Kuntze *93, N. Aberdeen: Wooded den, Tore of Troup,
NJ/832.613. D. Welch, 1991, ABD.

25/cam. X int. POLYPODIUM CAMBRICUM L. X P. INTERJECTUM Shivas *14, E. Sussex: Old walls
of ruined Greyfriars Chapel, Winchelsea, TQ/905.170. L. B. Burt & M. Thomas, 1991, BM, det.
R. H. Roberts.

¢27/1. AZOLLA FILICULOIDES Lam. *44, Carms.: Ornamental pond, Gellideg Farm, SN/
423.105. T. S. Crosby, 1991. Inlet to lake, Sandy Water Park, Llanelli, SN/493.005. G. Hutchinson,
1991, NMW. Ist and 2nd records. 49, Caerns.: Reservoir near Pistyll Church, SH/328.422. L. J.
Larson, 1991, NMW. Only extant locality.

28/1. BOTRYCHIUM LUNARIA (L.) Sw. 81, Berwicks.: Basaltic outcrop, Hareheugh Craigs, NT/
687.401. D. G. Long, 1991. 2nd extant locality.

142 PLANT RECORDS

46/7. RANUNCULUS SARDOUS Crantz 46, Cards.: Sandy field, Gwbert, SN/163.498. A. O.
Chater, 1991, NMW. Ist localised post-1930 record.

46/9. RANUNCULUS PARVIFLORUS L. 35, Mons.: Thin grass on bank, R. A. F. Caerwent, ST/
467.919. T. G. Evans, 1991. ist post-1930 record.

46/11. RANUNCULUS LINGUA L. *98, Main Argyll: Swamp, Lochan Iliter, NM/749.102. B. H.
Thompson, 1991, herb. B.H.T., conf. C. D. Preston.

46/19 X 22b. RANUNCULUS FLUITANS Lam. X R. PELTATUS Schrank *68, Cheviot: Whiteadder
Water at Whiteadder Bridge, Corporation Arms, NT/959.522. J. M. Croft & C. D. Preston, 1991,
CGE, conf. S. D. Webster.

46/22a X 21. RANUNCULUS AQUATILIS L. X R. TRICHOPHYLLUS Chaix *50, Denbs.: Farm pond,
Penley, SJ/408.412. B.S.B.I. meeting, 1991, CGE, det. S. D. Webster. Ist Welsh record.

46/22c. RANUNCULUS PENICILLATUS subsp. PSEUDOFLUITANS (Syme) S. Webster var. PSEUDO-
FLUITANS *77, Lanarks.: R. Clyde, Wolfclyde, NT/01.36. R. C. L. Howitt, 1979, herb. P.
Macpherson, conf. N. T. H. Holmes.

+50/aqu. THALICTRUM AQUILEGIIFOLIUM L. *79, Selkirks.: Railway embankment between
Bowland and Galashiels, NT/4.3. E. P. Beattie, 1969, E. *80, Roxburghs.: Riverbank, Ettrick
Water below Bridgeheugh, Selkirk, NT/476.307. E. Middleton, 1988, E, det. D. R. McKean.

57/1. CERATOPHYLLUM DEMERSUM L. *44, Carms.: Lake, Sandy Water Park, Llanelli, SN/
493.005. I. K. Morgan, 1991, NMW. 70, Cumberland: Recently dug pond, Bowscar, NY/
522.344. K. Raistrick, 1991, LANC. 2nd record.

58/3. PAPAVER DUBIUM subsp. LEcCoqi (Lamotte) Syme *44, Carms.: Burnt cliff, Dolwen
Point, SN/233.078. I. K. Morgan, 1991.

66/4. FUMARIA BASTARDII Boreau *67, S. Northumb.: Waste ground, Mason, NZ/211.735.
G. A. Swan, 1990, herb. G.A.S., det. M. G. Daker.

+68/1. ERUCASTRUM GALLICUM (Willd.) O. E. Schulz 70, Cumberland: Forestry track,
Spadeadam, NY/6.7. M. Gregory & J. Parker, 1991, LANC, det. T. C. G. Rich. 2nd record.

+71/1. HIRSCHFELDIA INCANA (L.) Lagr.-Fossat *50, Denbs.: Kinmel Bay, SH/990.808.
G. Battershall, 1991. *59,S. Lancs.: Railway siding, Fazakerley, SJ/388.983. V. Gordon, 1991.

79/3. LEPIDIUM HETEROPHYLLUM Benth. 26, W. Suffolk: Farm track, Eriswell, TL/754.764.
D. J. & Y. T. Leonard, 1991, herb. E. & M. Hyde, conf. T. C. G. Rich. Only extant locality.

+86/2. CAPSELLA RUBELLA Reuter *38, Warks.: Canal towpaths, Rowington, SP/187.699, and
Kingswood, SP/187.708. J. C. Bowra, 1991, WAR, det. P. J. Copson. Ist and 2nd records.

88/4. COCHLEARIA SCOTICA Druce *81, Berwicks.: St Abb’s Head, NT/9.6. A. Craig-Christie,

1884, E, det. P. S. Wyse Jackson.

+92/1. LOBULARIA MARITIMA (L.) Desv. 50, Denbs.: Colwyn Bay, SH/846.782. G. Battershall,
1991. 2nd post-1930 record.

+97/rap. CARDAMINE RAPHANIFOLIA Pourret *49, Caerns.: Bank of Afon Roe near Pont
Gorswen, SH/762.714. R. Lewis, 1990, NMW, det. E. J. Clement & T. C. G. Rich.

102/4. RorippA ISLANDICA (Oeder ex Murray) Borbas “46, Cards.: Flat rock by Afon Teifi
below Cenarth bridge, SN/269.416. A. O. Chater, 1991, NMW, det. B. Jonsell & T. C. G. Rich. Ist
Welsh record, and extension of range 250 km southwards.

102/5. RoRIPPA AMPHIBIA (L.) Besser *44, Carms.: Banks of Afon Llwchwr, Llanedi, SN/
59.06. I. K. Morgan, 1984, NMW.

102/6 X 3. RORIPPA AUSTRIACA (Crantz) Besser X R. SYLVESTRIS (L.) Besser *58, Cheshire:
Edge of lane, Partington, SJ/733.914. A. Franks & P. Newton, 1990, det..T. C. G. Rich.

PLANT RECORDS 143

4108/3. SISYMBRIUM LOESELII L. *25, E. Suffolk: Sandy bank behind Marina, Shotley Point,
TM/252.343. E. M. Hyde, 1991, herb. T. C. G. Rich, det. T.C.G.R.

111/1. DESCURAINIA SOPHIA (L.) Webb ex Prantl +42, Brecs.: Roadside verge, Llangattock,
SO/21.18. A. Newton & M. Porter, 1989. Ist post-1930 record. +48, Merioneth: Disturbed
ground by golf links, Morfa Harlech, SH/5.3. P. M. Benoit & D. C. Lang, 1990, NMW. Ist record
this century.

113/5 X 4. VIOLA REICHENBACHIANA Jordan ex Boreau X V. RIVINIANA Reichb. “44, Carms.:
Roadbank, Garn Fach, SN/571.156. R. D. Pryce, 1985, NMW, det. D. M. Moore.

113/6 X 4. VIOLA CANINA L. X V. RIVINIANA Reichb. *75, Ayrs.: Dune grassland, Lendalfoot,
NX/129.898. A. McG. Stirling, 1991.

113/9b. VIOLA PALUSTRIS subsp. JURESSII P. Fourn. *47, Monts.: Bare peat, Dovey Junction,
SN/703.982. P. M. Benoit, 1989.

¥113/10. VioLa coRNuUTA L. *77, Lanarks.: Woodland, Cleghorn Estate, NS/8.4. W. A. Scott,
1965.

123/la. SILENE VULGARIS Garcke 93, N. Aberdeen: Disturbed roadside bank, Auchterless,
NJ/706.411. D. Welch, 1991, ABD. Ist record since 1930.

123/14 Xx 13. SILENE LATIFOLIA Poiret X S. piorca (L.) Clairv. *75, Ayrs.: Shingle shore,
Pinbain, NX/136.913. A. McG. Stirling, 1991.
+124/cha. LYCHNIS CHALCEDONICA L. *35, Mons.: Forestry track, Bishop Barnet’s Wood,

Godefroy, ST/516.938. T. G. Evans, 1989, NMW.

131/10. CERASTIUM DIFFUSUM Pers. 712, N. Hants.: Abundant on roadside verges, Aldershot
military town, SU/870.517. C. R. Hall, 1991, herb. A. Brewis, det. F. Rose. 2nd record.

133/3. STELLARIA PALLIDA (Dumort.) Piré 81, Berwicks.: Basaltic outcrop, Hareheugh Craigs,
NT/688.400. R. W. M. Corner, 1991, herb. M. E. Braithwaite. 2nd record.

143/2. SPERGULARIA BOCCONI! (Scheele) Graebner +*15, E. Kent: Roadside verge on chalky
soil, Lower Bell near Maidstone, TQ/74.60. E. G. Philp, 1991, MNE, conf. J. Ratter.

144/1. POLYCARPON TETRAPHYLLUM (L.) L. 9, Dorset: Sandy shingle, Ferrybridge, S Y/667.756.
S. M. Eden, 1991. Only extant locality, in area where first recorded in 1774.

146/1. HERNIARIA GLABRA L. +*77, Lanarks.: Bare ground, site of 1988 Garden Festival,
Glasgow, NS/56.65 and 57.65. P. Macpherson, 1990, herb. P.M. Present in increased quantity in
£991:

149/1b. MONTIA FONTANA subsp. MINOR Hayw. 81, Berwicks.: Basaltic outcrop, Hareheugh
Craigs, NT/687.399. R. W. M. Corner, 1991, herb. M. E. Braithwaite. 2nd record.

*152/1. CARPOBROTUS EDULIS (L.) N. E. Br. *25, E. Suffolk: Shingle beach, Landguard
Common, TM/28.31. G. E. Steeds, 1990, det. C. D. Preston.

7153/bli. AMARANTHUS BLITOIDES S. Watson *26, W. Suffolk: Sandy arable field, Icklingham,
TL/768.735. E. Milne-Redhead, 1988, det. A. L. Grenfell.

154/14. CHENOPODIUM RUBRUM L. +*43, Rads.: Dung heap near New Radnor Station, SO/
219.605. D. C. Boyce, D. R. Drewett & R. G. Woods, 1991.

156/4 X pra. ATRIPLEX GLABRIUSCULA Edmondston xX A. PRAECOX Hulph. *68, Cheviot:
Saltmarsh, Goswick, NU/054.458. G. A. & M. Swan, 1991, herb. G.A.S., det. J. R. Akeroyd.

160/2. SALICORNIA DOLICHOSTACHYA Moss 44, Carms.: Channel to dock, Llanelli, SS/499.988.
_G. Hutchinson, 1987, NMW, det. D. H. Dalby. 1st post-1930 record.

~ 163/4. MALVA NEGLECTA. Wallr. *75, Ayrs.: Car park and shore, Girvan, NX/182.975.
A. McG. Stirling & A. Rutherford, 1991, E.

144 PLANT RECORDS

164/1. LAVATERA ARBOREA L. 50, Denbs.: Kinmel Bay, SH/994.807. G. Battershall, 1991. 2nd
record, Ist since 1859.

+165/2. ALTHAEA HIRSUTA L. 11, S. Hants.: Short coastal turf on reclaimed land, North
Harbour, Cosham, SU/643.050. R. A. Barrett, 1991. Ist record as established population since
1922.

168/12. GERANIUM ROTUNDIFOLIUM L. +*46, Cards.: Waste ground and scrub, Llandbadarn
Fawr, SN/598.810. S. P. Chambers, 1991, NMW.

169/1. ERODIUM MARITIMUM (L.) L’Heér. 14, E. Sussex: Bare chalk on cliff top, Flagstaff Point,
TV/538.967. R. M. Burton, 1991, herb. P. A. Harmes. Only extant locality. *35, Mons.: Grassy
bank near R. Severn, Sudbrook, ST/50.87. T. G. Evans, 1978. Rock garden and drive, La Cuesta,
Chepstow, ST/52.93. T. G. Evans, 1981, still present in 1991. Introduced with sand and now well
established. Ist and 2nd records.

169/2. ERODIUM MOSCHATUM (L.) L’Heér. 711, S. Hants.: Short turf by pavement, Portsdown
Hill Road (B2177), Widley, SU/668.064. D. P. J. Smith, 1991, herb. R. P. Bowman. ist record of
established population since 1924.

7170/3. OXALIS STRICTA L. *46, Cards.: Trackside in pasture, Glanrheidol, SN/663.792. A. O.
Chater, 1990, NMW.

+170/exi. OXALIS EXILIS Cunn. *46, Cards.: Abundant in lawns, Carrog, SN/562.724. A. O.
Chater, 1991, NMW.

+RHUS HIRTA (L.) Sudw. *70, Cumberland: Derelict industrial land, Mealsgate near Fletcher-
town, NY/201.423. M. Porter, 1991.

7183/2. LUPINUS ARBOREUS Sims *75, Ayrs.: Sandy ground near shore, Seafield, NS/328.205.
A. McG. Stirling & A. Rutherford, 1991.

+184/alp. LABURNUM ALPINUM (Miller) Bercht. & J. S. Presl 70, Cumberland: Damp bank E.
of Kershopefoot, NY/481.827. R. E. Groom, 1986, LANC, conf. A. O. Chater. 2nd record.

187/3. ULEX MINOR Roth +*59, S. Lancs.: Embankment of disused railway line, Culcheth, SJ/
646.950. V. Gordon, 1990.

+191/4. MELILoTUsS INDICcUS (L.) All. 12, N. Hants.: Disturbed sandy ground, Aldershot, SU/
858.508. T. Dove & A. R. G. Mundell, 1990, herb. A.R.G.M. 2nd record.

200/3. ASTRAGALUS GLYCYPHYLLOS L. 35, Mons.: Rough meadow, R.A.F. Caerwent, ST/
483.917. T. G. Evans and C. & G. Titcombe, 1991. 2nd extant locality.

205/1. ONOBRYCHIS VICHFOLIA Scop. +70, Cumberland: Bank near old railway, Maryport, N Y/
02.36. N. Botham, 1991, LANC. Ist record this century.

+206/6. VICIA VILLOSA Roth 12, N. Hants.: Abundant in disturbed sandy ground, Aldershot,
SU/858.508. T. Dove & A. R. G. Mundell, 1990, herb. A.R.G.M. 2nd record.

206/10. VictIa SYLVATICA L. 12, N. Hants.: Wood W. of Appleshaw, SU/296.485. C. Chatters
& I. Routh, 1991. Ist record since 1805.

+106/14 seg. VICIA SATIVA subsp. SEGETALIS (Thuill.) Gaudin *77, Lanarks.: Grassy verge,
Busby near Glasgow, NS/57.56. E. L. S. Lindsay & P. Macpherson, 1984, herb. P.M., conf.
A. McG. Stirling.

206/16. VICIA LATHYROIDES L. *1, W. Cornwall: Dry, stony soil on bank, Pendennis Castle,
SW/824.317. R. J. Murphy & P. E. Tompsett, 1991, conf. R. M. Walls.

207/2. LATHYRUS NISSOLIA L. +46, Cards.: Railway embankment, Eglwys-Fach, SN/67.96.
W. M. Condry, 1968, 1972. 2nd record. +*59, S. Lancs.: Weed amongst planted bushes, Risley
Moss, SJ/668.928. V. Gordon, 1991.

PLANT RECORDS 145

+ARUNCUS DIoICcUS (Walter) Fern. *70, Cumberland: Lakeshore, Ullswater E. of Lyulph’s
Tower, NY/410.203. F. J. Roberts, 1991, LANC.

211/11/32. RUBUS BRITANNICUS Rogers = *29, Cambs.: Woodland margin, Warren Hill, TL/
660.637. A. L. Bull, 1989, herb. A.L.B., conf. A. Newton.

211/11/51. RUBUS GLANDULIGER W. C. R. Watson *11, S. Hants.: Edge of clearing, Buckland
Wood, Lymington, SZ/31.97. D. E. Allen, 1990, BM, conf. A. Newton.

211/11/74. Rusus sitvaticus Weihe & Nees *12, N. Hants.: Wood margin, Sydmonton
Common, SU/49.62. D. E. Allen & F. H. Brightman, 1986, herb. D.E.A., conf. A. Newton.

211/11/81. RuBuUs ALBIONIS W. C. R. Watson *12, N. Hants.: Birchwood, Mount Pleasant,
Tadley Common, SU/60.62. D. E. Allen, 1990.

211/11/83. RUBUS CRUDELIS W. C. R. Watson “11, S. Hants.: Wood and heath margin, Chark
Common, SU/57.02. D. E. Allen, 1990, BM, conf. A. Newton.

211/11/107. RuBUs PAMPINOSUS Lees *12, N. Hants.: Wood margin, Hawley Common, SU/
83.57. D. E. Allen, 1984, herb. D.E.A., conf. A. Newton.

_ 211/11/121. RuBUs cISSBURIENSIS W. C. Barton & Riddelsd. *4, N. Devon: Plantation verge,
Challice’s Plantation, Eggesford, SS/68.09. L. J. Margetts & W. H. Tucker, 1991, det, A. Newton.

7211/11/139. RuBus prRoceRrus Muller *81, Berwicks.: Rough bank in village, Coldingham,
NT/904.662. D. P. Earl, 1991.

211/11/158. RUBUS CINEROSIFORMIS Rilstone *9, Dorset: Heathy common, Lambert’s Castle
Hill, SY/363.987. L. J. Margetts, 1991, RNG.

211/11/198. RUBUS AHENIFOLIUS W. C. R. Watson *12, N. Hants.: Roadside verges near
Pamber End, SU/60.58. D. E. Allen, 1990, BM, conf. A. Newton. *22, Berks.: Roadside,
Silchester Common, SU/62.62. D. E. Allen, 1990.

211/11/329. RuBus RILSTONE! W. C. Barton & Riddelsd. *9, Dorset: Forestry rides,
Champernhayes Plantation, Wootton Hill, SY/355.970. L. J. Margetts, 1991, RNG.

212/3. POTENTILLA STERILIS (L.) Garcke 93, N. Aberdeen: Rocky ground by waterfall, Turriff,
NJ/712.511. D. Welch, 1991, ABD. Ist post-1930 record.

7212/7. POTENTILLA RECTA L. *77, Lanarks.: Grassy waste ground, Glasgow, NS/56.64. P.
Macpherson, 1991, herb. P.M. :

+220/3/12. ALCHEMILLA MOLLIS (Buser) Rothm. *50, Denbs.: Waste ground by farm lane,
north-east of Llanrwst, SH/837.655. R. Lewis, 1990, NMW. This record was erroneously attributed
to v.c. 49 in Watsonia 18: 425 (1991).

+223/2. SANGUISORBA MINOR subsp. MURICATA (Gremli) Briq. *44, Carms.: Llandybie, SN/
610.145. R. D. Pryce, 1987. Carway, SN/455.067. F. H. Webb, 1991, det. R. D. Pryce. Both records
from hedgebanks on restored opencast sites. Ist and 2nd records. *50, Denbs.: Roadside,
Redbrook, SJ/508.412. K. Watson, 1991.

4224/inc. ACAENA INERMIS Hook.f. *77, Lanarks.: Stabilised shingle near Culter Water,
Coulter, NT/03.33. D. J. McCosh, 1981, herb. D.J.McC. Known here for at least 20 years.

_ 225/8 X 7. ROSA CANINA L. X R. sTYLOSA Desv. *46, Cards.: Hedge, Mwldan valley, Penparc,
SN/197.486. A. O. Chater, 1991, NMW, det. G. G. Graham.

225/cae. ROSA CAESIA subsp. GLAUCA (Nyman) G. G. Graham & Primavesi 50, Denbs.:
Hedge, Plas-yn-rhos, SJ/169.480. K. Watson, 1991. 2nd record.

225/cae. X 8. ROSA CAESIA subsp. GLAUCA (Nyman) G. G. Graham & Primavesi X R. CANINA
L. *29, Cambs.: Hedge, Fowlmere Watercress Beds R.S.P.B. reserve, TL/405.458. A. Asher,
P. H. Oswald & C. D. Preston, 1991, det. A. L. Primavesi. *46, Cards.: Hedge, Gwaryfelin,
SN/595.794. A. O. Chater, 1991, NMW, det. G. G. Graham.

146 PLANT RECORDS

225/8 X 14. ROSA CANINA L. X R. RUBIGINOSA L. *64, Mid-W. Yorks.: Hedge, Micklefield,
SE/445.336. P. P. Abbott, 1990, det. G. G. Graham.

225/8 X 15. ROSA CANINA L. X R. MICRANTHA Borrer ex Smith *48, Merioneth: Side of track, ©

Arthog, SH/6.1. P. M. Benoit, 1991, NMW, conf. A. L. Primavesi.

225/10. ROSA OBTUSIFOLIA Desv. *46, Cards.: Roadside verge S. of Glanrheidol, SN/663.789.
A. O. Chater, 1991, NMW, det. G. G. Graham.

225/13 X 12. RoSA MOLLIS Smith xX R. SHERARDII Davies *46, Cards.: Hedge, Coed
Cwmhwylog, Nanteos, SN/622.778. A. O. Chater, 1991, NMW, det. G. G. Graham.

225/15. ROSA MICRANTHA Borrer ex Smith *69, Westmorland: Limestone scrub, Heathwaite,
SD/451.768. M. Baecker, 1991, LANC, det. G. G. Graham.

225/17. ROSA AGRESTIS Savi 11, S. Hants.: Scrub woodland, South Charford Drove,
Breamore Down, SU/139.207. R. P. Bowman, 1991, herb. R.P.B., det. A. L. Primavesi. 1st record
since 1891 record at this site.

225/17 X 15. ROSA AGRESTIS Savi X R. MICRANTHA Borrer ex Smith *11, S. Hants.: Scrub
woodland, South Charford Drove, Breamore Down, SU/139.207. R. P. Bowman, 1991, herb.
R.P.B., det. A. L. Primavesi.

+226/7. PRUNUS LAUROCERASUS L. *12, N. Hants.: By cricket ground, Amport, SU/304.438.
M. F. Wildish, 1991.

+227/3. COTONEASTER HORIZONTALIS Decne. *12, N. Hants.: Water meadows, Winnall Moors
N.R. near Winchester, SU/48.30. M.S.C. team, 1986. Chalk cutting on old road, Weyhill, SU/
304.462. M. F. Wildish, 1991. 1st and 2nd records.

+227/atr. COTONEASTER ATROPURPUREUS Flinck & Hylmoe *77, Lanarks.: Scrubby waste
ground, Cambuslang, NS/649.602. P. Macpherson, 1987. Old industrial site, Cambuslang, NS/
646.605. P. Macpherson, 1988. Both herb. P.M., det. J. Fryer & J. R. Palmer. Ist and 2nd records.

+227/hje. COTONEASTER HJELMQVISTI Flinck & Hylmoe *77, Lanarks.: Bank of R. Clyde,

Glasgow, NS/58.64. P. Macpherson, 1989, herb. P.M., det. J. Fryer.

_ ~227/tra. COTONEASTER TRANSENS Klotz *11, S. Hants.: Edge of ride in conifer plantation,
Hurn Forest, SU/106.018. R. M. Walls & R. P. Bowman, 1985, herb. R.P.B., det. J. Fryer.
Published as C. affinis Lindley in Watsonia 17: 471 (1989).

229/1. CRATAEGUS LAEVIGATA (Poiret) DC. 50, Denbs.: Edge of lane, Isycoed Farm, SJ/
413.519. P. Day, 1990. 2nd record. *63, S.W. Yorks.: Hedge S.W. of Cawthorne, SE/273.062.
P. P. Abbott, 1991.

229/1 X 2. CRATAEGUS LAEVIGATA (Poiret) DC. X C. MoNOGYNA Jacq. *38, Warks.: Hedge,
Hampton in Arden, SP/199.829. S. M. Apted, 1991, WAR, det. P. J. Copson & J. C. Bowra.

232/5/1 X 1. SorBus ARIA (L.) Crantz X S. AUCUPARIA L. +*59,S. Lancs.: Field edge, Rimrose
valley, Waterloo, Crosby, SJ/327.985. V. Gordon, 1991.

232/5/7. SORBUS RUPICOLA (Syme) Hedlund *102, S. Ebudes: Bagh Uamh Mhor, Beinn na
Capull, Jura, NR/67.99. D. J. McCosh, 1991, E, det. A. McG. Stirling.

+232/cro. SORBUS CROCEOCARPA Sell 44, Carms.: Open oakwood, Llety-yr-ychen Fawr, Burry
Port, SN/455.015. I. K. Morgan, 1991, NMW, det. P. J. M. Nethercott. 2nd record. gr a):
Lanarks.: Scrubby wood, Shieldhall, Glasgow, NS/53.65. Laneside, Shieldhall, Glasgow, NS/53.66.
Both P. Macpherson, 1990, herb. P.M., det. P. J. M. Nethercott. Ist and 2nd records.

235/2a. SEDUM TELEPHIUM L. subsp. TELEPHIUM +*77, Lanarks.: Roadside west of East
Kilbride, NS/59.52. A. C. & P. Macpherson, 1984, herb. P.M., conf. D. R. McKean.

235/10. SEDUM FORSTERIANUM Smith +*39, Staffs.: With S. album on limestone rocks,
Milldale, S. of Alstonefield, SK/132.549. B. R. Fowler, 1991, herb. B.R.F.

PLANT RECORDS 147

+237/hel. CRASSULA HELMsII (Kirk) Cockayne 2, E. Cornwall: Shallow pool near Lamledra
| Farm, SX/011.410. R. E. Lees & R. J. Murphy, 1991. 2nd record. *64, Mid-W. Yorks.: Edge of
gravel pit, Ben Rhydding, SE/146.476. Yorkshire Naturalists’ Union meeting, 1991.

239/6 X 5. SAXIFRAGA HIRSUTA L. X S. SPATHULARIS Brot. +*77, Lanarks.: Woodland in old
- estate, Hartwood, NS/84.59. P. Macpherson, 1975, herb. P.M., det. C. A. Stace.

+240/1. TELLIMA GRANDIFLORA (Pursh) Douglas ex Lindley *79, Selkirks.: Riverside, island at
- confluence of Ettrick and Yarrow Waters, NT/447.273. D. Methven et al., 1988, herb. R. W. M.
- Corner.

¥241/1. TOLMIEA MENZIEsII (Pursh) Torrey & A. Gray *47, Monts.: Edge of R. Vyrnwy, Pont
Llogel, SJ/033.154. R. G. Woods, 1991.

+DARMERA PELTATA (Torrey ex Benth.) Voss ex Post & Kuntze *69, Westmorland: Laneside
_N. of Endmoor, SD/537.855. C. E. Wild, 1989, still present in 1991, G. Halliday, LANC.

7246/4. RIBES SANGUINEUM Pursh *49, Caerns.: Naturalized in woodland near Gwydyr
/ Castle, Llanrwst, SH/792.611. R. Lewis, 1991, NMW.

+246/odo. RIBES ODORATUM Wendl.f. “Va. Wwananks.: Bonk of R. Clyde, Hutchesontown, NS/
59.63. P. Macpherson, 1988, E, det. D. R. McKean.

253/1. LUDWIGIA PALUSTRIS (L.) Elliott +14, E. Sussex: Dew pond, South Hill Barn, Seaford
Head, TV/505.980. P. D. L. Maurice, 1991, det. J. Wheatley. lst record since c.1876.

254/46 X 3. EPILOBIUM CILIATUM Raf. X E. MONTANUM L. *42, Brecs.: Disused railway,
Llanhamlach, SO/08.27. M. Porter, 1987, det. T. D. Pennington. “46, Cards.: Waste ground,
Tal-y-bont village, SN/654.893. A. O. Chater, 1990.

254/+6 X 7. EPILOBIUM CILIATUM Raf. X E. TETRAGONUM L. “44, Carms.: Rank vegetation in
‘flood zone of stream, Pentre-cwrt, SN/390.386. G. Hutchinson, 1991, NMW, det. G. D. Kitchener
& B. Wurzell. 1st Welsh record.

256/42 X +1. OENOTHERA GLAZIOVIANA Micheli ex C. Martius < O. BIENNIS L. *38, Warks.:
| Waste ground, Emscote, SP/298.654. J. C. Bowra, 1988, WAR.
| 259/4. MYRIOPHYLLUM ALTERNIFLORUM DC. 61, S.E. Yorks.: Pond, Thornton Elers, Thorn-

‘ton, SE/73.45. D. R. Grant, 1990. Only extant locality.

+259/aqu. MyRIOPHYLLUM AQUATICUM (Vell. Conc.) Verdc. *28, W. Norfolk: Pond, Burnham
“Market, TF/826.418. G. Beckett, 1989, still present in 1991. *44, Carms.: Dafen Pond, Llanelli,
) SN/531.015. I. K. Morgan, 1991, NMW, conf. G. Hutchinson.

| 262/3. CALLITRICHE OBTUSANGULA Le Gall *67, S. Northumb.: Ditch near Healeywood, NZ/
| 232.849. G. A. Swan, 1991, herb. G.A.S., det. Q. O. N. Kay.

| }AucuBA JAPONICA Thunb. ~ *99, Dunbarton: Wooded streamside, Millig Burn, Helensburgh,
NS/29.82. A. Rutherford, 1985.

| 268/1 hib. HEDERA HELIX subsp. HIBERNICA (Kirchner) D. McClint. +*77, Lanarks.: Bank of
/ North Calder, Maryville, NS/68.62. A. McG. Stirling, 1986. 1st record of naturalized population.

285/4. APIUM INUNDATUM (L.) H. G. Reichb. 80, Roxburghs.: Side of Woo Burn, Ashkirk,
'NT/461.178. R. W. M. Corner, 1991, herb. R.W.M.C. 2nd extant locality.

287/1. SISON AMOMUM L. 44, Carms.: Edge of path, Penclacwydd Wildfowl Centre, SS/
1 530.986. B. Stewart, 1991, det. I. K. Morgan. Ist record since 1840s.

319/5. EUPHORBIA HYBERNA L. +*46, Cards.: Well established in scrub on slope, Llanbadarn
} Fawr, SN/598.810. S. P. Chambers, 1991, det. A. O. Chater.

| 320/1/3. POLYGONUM RURIVAGUM Jordan ex Boreau *25, E. Suffolk: Roadside verge, Tunstall,
| TM/35.54. A. L. Bull, 1991, herb. E. & M. Hyde, conf. B. T. Styles.

148 PLANT RECORDS

320/1/are. POLYGONUM ARENASTRUM Boreau “75, Ayrs.: Turf paths, Ardneil Bay, NS/18.48.
A. Somerville, 1903, CGE, det. B. T. Styles.

320/2. POLYGONUM OXYSPERMUM subsp. RAI (Bab.) D. Webb & Chater 28, W. Norfolk: |
Shingle beach, Snettisham, TF/646.329. M. Keene, 1991, conf. K. A. & G. Beckett. 2nd record. | |

+320/7. PERSICARIA AMPLEXICAULIS (D. Don) Ronse Decraene 64, Mid-W. Yorks.: Road |

verge, Wetherby, SE/406.475. P. P. Abbott, 1991. 2nd record.

320/14. PERSICARIA MINOR (Hudson) Opiz 44, Carms.: Mud in old oxbow of R. Tywi, Bishop’s | '
Pond, Abergwili, SN/443.209. I. K. Morgan, 1991, NMW, det. G. Hutchinson. 1st record this |
century. 58, Cheshire: Edge of Oakmere, SJ/575.678. A. Franks, 1991, herb. G. M. Kay, det. |
J. R. Akeroyd. 2nd extant locality. *75, Ayrs.: Stoney margin of Loch Maberry, NX/286.759. k i

A. McG. Stirling & A. Rutherford, 1991, E.

320/419 X +20. FALLOPIA JAPONICA (Houtt.) Ronse Decraene X F. SACHALINENSIS (F. Schmidt ex | I

Maxim.) Ronse Decraene *4, N. Devon: Hedge, Philham, SS/259.234. W. H. Tucker, 1991,

herb. W.H.T., conf. L. J. Margetts. *77, Lanarks.: Trackside in wood, Bothwell, NS/69.59. i

P. Macpherson & E. L. S. Lindsay, 1991, herb. P.M., conf. D. H. Kent.

+320/22. PERSICARIA CAMPANULATA (Hook. f.) Ronse Decraene *48, Merioneth: Edge of i

stream near Egryn Abbey, SH/5.1. P. M. Benoit, 1991.

325/2 bif. RUMEX ACETOSA subsp. BIFORMIS (Lange) Valdes-Berm. & Castroviejo. *46, Cards.:

With Crithmum on sea cliffs W. of New Quay, SN/373.595. A. O. Chater, 1988. Coastal grassland |

S.W. of Aberaeron, SN/426.607. A. O. Chater, 1991, det. P. D. Sell. 1st and 2nd records.

325/11 X 12. RUMEX crispuS L. X R. OBTUSIFOLIUS L. *50, Denbs.: Arable field, Penley, SJ/ |

408.407. D. Tinston, 1991, NMW.

325/15 X 13. RUMEX CONGLOMERATUS Murray X R. PULCHER L. *2, E. Cornwall: Footpath
S.W. of Pentire Farm, SW/934.800. G. D. Kitchener, 1989, det. J. R. Akeroyd.

343/2 X 1. SALIX ALBA L. X S. PENTANDRA L. *59, S. Lancs.: Disused railway line, Culcheth,
SJ/640.956. V. Gordon, 1991, herb. V.G., conf. R. D. Meikle.

343/2 X 4. SALIX ALBA L. X S. FRAGILIS L. *77, Lanarks.: Waste ground, Braehead, Glasgow,
NS/52.67. P. Macpherson, 1987, herb. P.M., det. R. D. Meikle.

343/6. SALIX PURPUREA L. 35, Mons.: Sandy bank of R. Usk, Llanllowell, ST/387.983. T. G.
Evans & D. Lewis, 1991, herb. T.G.E. Only extant locality.

*351/1. GAULTHERIA SHALLON Pursh *44, Carms.: Edge of conifer plantation, Gelli Aur, SN/
597.197. I. K. Morgan, 1991, NMW. |

+352/1. GAULTHERIA MUCRONATA (L.f.) Hook. & Arn. *46, Cards.: Dry roadside bank,
Eglwys Fach, SN/685.952. A. O. Chater & C. D. Preston, 1991, NMW.

1370/5. LYSIMACHIA PUNCTATA L. *79, Selkirks.: Steep bank, Clovenfords, NT/452.367.
R. W. M. Corner, 1991, herb. R.W.M.C.

372/1. ANAGALLIS TENELLA (L.) L. 81, Berwicks.: Flush in heather moor, Wheel Burn, NT/
565.515. M. E. Braithwaite, 1991, herb. M.E.B. 2nd record, Ist since 1853.

382/1. CENTAURIUM PULCHELLUM (Sw.) Druce *67, S. Northumb.: Dune slack, Hadston
Links, NZ/273.990. G. A. Swan, 1991, herb. G.A.S., det. F. Ubsdell.

387/1. NYMPHOIDES PELTATA Kuntze +*69, Westmorland: Old clay pit, Barrow-in-Furness, ©

SD/194.703. P. Burton, 1991, LANC. +*98, Main Argyll: Backwater of R. Awe at Fanans, NN/
033.293. D. Dugan, 1985, still present in 1991, B. H. Thompson, herb. B.H.T., conf. C. D. Preston.

}PHACELIA TANACETIFOLIA Benth. *41, Glam.: Railway cutting, Wingfield Road, Whitchurch,
ST/152.795. G. Hutchinson, 1991, NMW, det. R. G. Ellis. 2nd Welsh record.

PLANT RECORDS 149

392/6. SYMPHYTUM TUBEROSUM L. 711, S. Hants.: Edge of wood, Hedgemoor Copse, West
Tytherley, SU/265.313. R. P. Bowman, 1966, herb. R.P.B., still present in 1988. Ist record since
1929. +59, S. Lancs.: Hedge bank, Formby, SD/308.086. V. Gordon, 1990. 2nd record.

+392/7. SYMPHYTUM GRANDIFLORUM DC. *77, Lanarks.: Edge of wood, Carmunnock, NS/
60.57. P. Macpherson, 1991, herb. P.M.

400/3. MyosoTIS STOLONIFERA (DC.) Gay ex Leresche & Levier *63, S.W. Yorks.: Acidic
flush, Hordron Clough, Langsett, SK/175.994. P. P. Abbott, 1991. Southerly extension of
range. 80, Roxburghs.: Wet flush, The Schil, NT/866.215. R. W. M. Corner & B. H.
Thompson, 1991, herb. R.W.M.C. 2nd record.

406/1 ros. CALYSTEGIA SEPIUM subsp. ROSEATA Brummitt +*46, Cards.: Railway embankment,
Eglwys Fach, SN/673.962. A. O. Chater & W. M. Condry, 1991, NMW. Hedgebank, Tresaith, SN/
279.514. A. O. Chater, 1991, NMW. 1st and 2nd records.

+416/3. VERBASCUM PHLOMOIDES L. 46, Cards.: Disused railway, Felin-y-mor, Aberystwyth,
SN/581.804. A. O. Chater, 1977, NMW, det. I. K. Ferguson. Still present in 1990.

[416/4 x 1. VERBASCU:1 LYCHNITIS L. X V. THAPSUS L. 46, Cards.: Delete record published in
Watsonia 12: 173 (1978). specimen in NMW is V. phlomoides L., det. I. K. Ferguson, 1990].

416/7. VERBASCUM NIG&UM L. +*42, Brecs.: Garden weed, Gilwern, SO/24.14. S. G. & A.
Marshall, 1989, det. M. Perter. 1st localised record.

7420/5. LINARIA DALMATICA (L.) Miller *64, Mid-W. Yorks.: Verge of old A65 road south of
Settle, SD/813.620. E. Shorrock, 1991, det. N. Frankland.

— 424/2. SCROPHULARIA AURICULATA L. 71, Man: Pasture by R. Dhoo, Glenlough, SC/343.779.
J. Lamb & P. A. Sayle, 1991, conf. D. E. Allen. 2nd record, 1st as an undoubted native.

7425/cup. X 1 X 2. MIMULUS CUPREUS Dombrain X M. Luteus L. X M. GuTtatus DC. *41,
Glam.: R. Rhymney between Llechryd and Rhymney, SO/1.0. T. G. Evans, 1989, NMW, det. A. J.
Silverside.

426/1. LIMOSELLA AQUATICA L. 75, Ayrs.: Edge of North Craig Reservoir, NS/437.414. R. L.
Griffith, 1991, herb. R.L.G. 2nd record. *93, N. Aberdeen: Mud by dune slack, Sands of
Forvie, NK/01.26. J. J. Barkman & C. H. Gimingham, 1990, ABD. Northerly extension of range.

|
}

430/2 X 3. VERONICA ANAGALLIS-AQUATICA L. X V. CATENATA Pennell *36, Herefs.: Disused
gravel pit, Bodenham, SO/52.51. J. M. Croft & C. D. Preston, 1991, CGE.

7430/25. VERONICA CRISTA-GALLI Steven 46, Cards.: Waste ground by Church Hall, Llanba-
darn Fawr, SN/598.810. A. O. Chater, 1991, NMW. 2nd record.

i 7430/aus. VERONICA AUSTRIACA Subsp. TEUCRIUM (L.) D. Webb *39, Staffs.: Limestone spoil
i railway sidings, Waterhouses, SK/072.492. B. R. Fowler, 1991.

[434/3. MELAMPYRUM PRATENSE L. 29, Cambs.: Delete record published in Watsonia 18: 429
1991); specimen in CGE is Stellaria holostea L. ]

| 435/1/19 X 13. EUPHRASIA ROSTKOVIANA Hayne X E. NEMOROSA (Pers.) Wallir. *35, Mons.:
‘loor of disused limestone quarry, Blackcliff, ST/533.984. T. G. Evans, 1988, herb. T.G.E., det.
a. J. Silverside. 1st Welsh record.

i 440/2. OROBANCHE PURPUREA Jacq. 26, W. Suffolk: Lakenheath, TL/72.82. M. G. Rutterford,
990, conf. F. W. Simpson. Ist post-1930 record. 45, Pembs.: Dunes over limestone west of
{anorbier Church, SS/06.97. M. Higgins, 1991. Only extant locality.

440/4. OROBANCHE ALBA Stephan ex Willd. +*57, Surrey: On Thymus in garden, host planted
) years ago and not disturbed for at least 10 years, Abinger Hammer, TQ/098.460. P. Verrall,
791, conf. F. J. Rumsey.

/442/2. UTRICULARIA AUSTRALIS R. Br. *80, Roxburghs.: Branxholme Wester Loch, NT/
22.110. M. E. Braithwaite, 1991, herb. R. W. M. Corner.

150 PLANT RECORDS

+464/rus. PHLOMIS RUSSELIANA (Sims) Benth. *70, Cumberland: Railway embankment,
Langwathby, NY/575.324. F. Lawson, 1957, CLE, and R. W. M. Corner, 1990, LANC, both —
specimens det. G. Halliday.

472/1 int. PLANTAGO MAJOR subsp. INTERMEDIA (Gilib.) Lange *35, Mons.: Black Rock Point,
ST/51.88. A. O. Chater, 1987, NMW.

472/5. PLANTAGO CORONOPUS L. +*43, Rads.: Close-grazed turf on roadside bank, Llannerch

Cawr, SN/900.616. M. Porter, 1991, NMW.

+475/S5. CAMPANULA PERSICIFOLIA L. 80, Roxburghs.: Railway cutting, Long Newton, NT/—
587.277. R. W. M. Corner, 1991, herb. R.W.M.C. 2nd record.

485/3 X 4. GALIUM MOLLUGO L. X G. VERUM L. *93, N. Aberdeen: Roadside bank,
Auchterless, NJ/706.411. D. Welch, 1991, ABD. |

485/6. GALIUM PUMILUM Murray 12, N. Hants.: Isle of Wight Hill, SU/245.373. P. Wilson,
1991. 2nd record.

485/10. GALIUM ULIGINOSUM L. 46, Cards.: Fen, Banc-y-mwldan S.S.S.I., Penparc, SN/ |
197.483. W. Fojt, 1987, conf. A. O. Chater.

494/2. VALERIANELLA CARINATA LOIS. 50, Denbs.: Colwyn Bay, SH/845.782. G. Battershall,
1991. 2nd record. *81, Berwicks.: Rock outcrop, Muckle Thairn, Girrick, NT/665.374. M. E. |
Braithwaite, 1991, E, det. D. R. McKean. Northerly extension of range.

4502/3. BIDENS FRONDOSA L. 44, Carms.: Witchett Pool, Laugharne, SN/28.07. V. Gordon, |
1956, NMW, det. G. Hutchinson. 2nd record.

506/+4 X 7. SENECIO SQUALIDUS L. X S. viscosus L. *58, Cheshire: By gravel track, Alsager,
SJ/796.548. G. M. Kay, 1990, herb. G.M.K.
506/+4 < 8. SENECIO SQUALIDUS L. X S. VULGARIS L. *45, Pembs.: Waste ground, Goodwick |
Harbour, SM/95.38. J. W. Partridge, 1988, NMW, det. C. Jeffrey.

506/6. SENECIO SYLVATICUS L. 111, Orkney: Eroded bank at top of beach, Rackwick Bay, ND/
197.990. B. H. Thompson, 1991. Only extant locality.

+509/3. PETASITES JAPONICUS (Siebold & Zucc.) Maxim. 2, E. Cornwall: Roadside bank,
Lower Woon, SX/035.623. B. Molland, 1991, det. R. J. Murphy. 2nd record. 49, Caerns.: Tidal |
mudbank, Afon Ro near confluence with Afon Conwy, SH/77.69. R. Lewis, 1991. 2nd
record. *79, Selkirks.: Side of R. Tweed below Yair Bridge, NT/462.325. D: Methven, 1991. |

513/1. PULICARIA DYSENTERICA (L.) Bernh. *99, Dunbarton: Ditch bank, Bannachra Muir, |
Helensburgh, NS/33.83. K. Futter, 1991, E, conf. A. McG. Stirling.

|
+518/2. SOLIDAGO CANADENSIS L. 50, Denbs.: Kinmel Bay, SH/990.808. G. Battershall, 1991. |
2nd record. |

+518/3. SOLIDAGO GIGANTEA subsp. SEROTINA (O. Kuntze) McNeill *38, Warks.: Disused
quarry, Littlke Compton, SP/270.291. J. C. Bowra, 1980, WAR, det. C. Jeffrey & D. H. Kent. |

1519/8. ASTER LANCEOLATUS Willd. *46, Cards.: Tidal river bank, Afon Rheidol, Glanyrafon,
SH/612.804. Waste ground near railway, Aberystwyth, SN/589.811. Both A. O. Chater, 1990,
NMW, det. P. F. Yeo. Ist and 2nd records.

*+519/9. ASTER X SALIGNUS Willd. *46, Cards.: Tidal river bank, Afon Leri, Borth, SN/
616.898. Railway embankment, Glandyfi, SN/696.976. Both A. O. Chater, 1990, NMW, det. P. F. |
Yeo. Ist and 2nd records.

+522/1. CONYZA CANADENSIS (L.) Cronq. “77, Lanarks.: Waste ground near R. Clyde, )
Glasgow, NS/56.65. P. Macpherson, 1991, herb. P.M.

PLANT RECORDS 151

538/3. ARCTIUM MINUS subsp. PUBENS (Bab.) Arenes *46, Cards.: Disturbed ground by
caravan site, Ystrad Teilo, Llanrhystud, SN/546.695. A. O. Chater, 1991, NMW, conf. F. H.
Perring.

544/3. CENTAUREA cYANUus L. 61, S.E. Yorks.: Roadside verge, Bursea Lane, SE/803.342.
F. E. Crackles, 1991. Only extant locality. 70, Cumberland: Grass verge, lane off Linefoot-
Broughton Moor Road, NY/075.341. N. Botham, 1972. Only extant locality.

552/2 X 1b. TRAGOPOGON PORRIFOLIUS L. X T. PRATENSIS subsp. MINOR (Miller) Wahlenb. o25.
E. Suffolk: Roadside verge, Tunstall, TM/35.54. P. G. Lawson, 1991.

554/1. LACTUCA SERRIOLA L. *58, Cheshire: Embankment and ditchside, Helsby, SJ/489.774.
wie Kay & ©. C2G. Rich; 1991.

558/1/3. HIERACIUM SPELUNCARUM Arv.-Touv. *77, Lanarks.: Spoil heaps amongst scrub near
Bishopbriggs Golf Course, NS/59.71. D. J. McCosh & K. J. Watson, 1991, GLAM.

~ 558/1/45. HIERACIUM LASIOPHYLLUM Koch *44, Carms.: Dry rocks above Troed-rhiw-rudd-
wen, Rhandirmwyn, SN/7.4. I. M. Vaughan, 1972, NMW, det. J. Bevan.

558/1/99. HIERACIUM GRANDIDENS Dahlst. *93, N. Aberdeen: Dyke along shelter belt, Oyne,
NJ/671.262. D. Welch, 1989, ABD, det. D. J. McCosh & P. D. Sell.

558/1/136. HIERACIUM CAESIOMURORUM Lindeb. *93, N. Aberdeen: Rock ledge by waterfall,
Craig, NJ/472.247. D. Welch, 1990, herb. D.W., det. D. J. McCosh.

558/1/222. HIERACIUM SALTICOLA (Sudre) Sell & C. West *41, Glam.: Grassy waste ground,
Cardiff Docks, ST/205.743. G. Hutchinson, 1986, NMW, det. J. Bevan.

558/1/223. HIERACIUM VAGUM Jordan *41, Glam.: Disused railway line, St Fagans, ST/
117.768. G. Hutchinson, 1990, NMW, det. J. Bevan.

558/1/mem. HIERACIUM MEMORABILE Sell & C. West 88, Mid Perth: Rocky ledges, N. Coire,
Beinn Heasgarnich, NN/41.38. D. J. Tennant, 1991, herb. D.J.T. Ist record since 1891 specimen
from this site.

7571/1. LAGAROSIPHON MAJOR (Ridley) Moss *39, Staffs.: Canal N. of Brownhills, SK/
046.071. J. P. Martin, 1990, K, det. D. A. Simpson.

577/11. POTAMOGETON FRIESII Rupr. *80, Roxburghs.: Branxholme Easter Loch, NT/43.11.
P. Macpherson, 1991, herb. P.M., det. C. D. Preston.

577/16. POTAMOGETON TRICHOIDES Cham. & Schldl. *12, N. Hants.: Pond by R. Blackwater,
Hawley, SU/855.595. C. R. Hall, 1991, herb. A. Brewis, det. N. T. H. Holmes & C. D.
Preston. *36, Herefs.: Recently cleared section of Hereford & Gloucester Canal E. of Skew
Bridge, Monkhide, SO/61.43. J. M. Croft & C. D. Preston, 1991, CGE.

579/1. RUPPIA CIRRHOSA (Petagna) Grande *69, Westmorland: Old mine shaft frequently
inundated by sea, Blacks Pond, Askam in Furness, SD/207.763. P. Burton, 1991, LANC, conf.
C. D. Preston.

601/1. MUSCARI NEGLECTUM Guss. ex Ten. +*70, Cumberland: Quarry, Catlands, NY/2.4.
_ M. Porter, 1991.

602/1. COLCHICUM AUTUMNALE L. +*73, Kirkcudbrights.: Mull Island, R. Cree, NX/409.662.
J. McCleary, 1989.

7605/2. JUNCUS TENUIS Willd. *67, S. Northumb.: Grassy track, W. bank of R. Allen opposite
Plankey Mill, NY/795.622. G. A. Swan, 1972, herb. G.A.S. First recorded in NY/7.6 in 1955; still
present in 1991. Waste ground near Albert Edward Dock, NZ/348.671. D. N. Mitchell, 1988, herb.
G.A.S. 1st and 2nd records.

605/4. JUNCUS COMPRESSUS Jacq. 35, Mons.: Wet hollow, White House Farm, SO/42.14. P. C.
& J. Hall, 1991, NMW, det. P.C. H. & T. G. Evans. Only extant locality. *58, Cheshire: Boggy
area by pool, Middlewich, SJ/699.669. J. H. Clarke, 1991, herb. G. M. Kay, det. C. A. Stace.

152 PLANT RECORDS

605/12. JUNCUS FILIFORMIS L. *59, S. Lancs.: Exposed shore of Earnsdale Reservoir, TM/
669.221. P. Jepson, 1991.

607/6. ALLIUM OLERACEUM L. 26, W. Suffolk: Roadside verge, Shaker’s Lane, Bury St
Edmunds, TL/86.64. P. G. Lawson & E. Milne-Redhead, 1991. 2nd extant locality.

+616/2. IRIS VERSICOLOR L. 70, Cumberland: Amongst Juncus by lakeside between Fawe Park
and Lingholm, Derwentwater, NY/254.224. E. Sterne, 1985, LANC, det. B. Mathew. 2nd record.

628/2. LISTERA CORDATA (L.) R. Br. 46, Cards.: Moorland N. of Cwm Ystwyth, SN/812.778.
A. Jones, 1991. 2nd record.

640/3. OPHRYS SPHEGODES Miller 26, W. Suffolk: Open grassland, Lakenheath, TL/7.8.
L. Farreli, 1991, conf. J. J. Wood. 1st record since 1793.

643/1 x 3b. DACTYLORHIZA FUCHSII (Druce) Sod X D. INCARNATA subsp. PULCHELLA (Druce)
So6 *62, N. E. Yorks.: Calcareous flush, Dalby Forest near Thornton Dale, SE/8.8.
F. Horsman, 1991.

643/1 xX 5. DacTyLorHizA FucHsi (Druce) So6 xX D. PURPURELLA (Stephenson & T. A.
Stephenson) So6 *46, Cards.: Dune slack, Ynys-las Dunes N.N.R., SN/611.938. F. Horsman,
1991.

643/3b. DACTYLORHIZA INCARNATA subsp. PULCHELLA (Druce) S06 65, N.W. Yorks.: Marsh,
Combe Scar, SD/679.875. F. Horsman, 1991. 2nd record.

643/3a X 2b. DACTYLORHIZA INCARNATA (L.) So6 subsp. INCARNATA X D. MACULATA subsp.
ERICETORUM (Linton) P. Hunt & Summerh. *46, Cards.: Fen, Mwldan valley N.N.E. of
Penparc, SN/201.489. F. Horsman, 1991.

643/4. DACTYLORHIZA PRAETERMISSA (Druce) So6 62, N.E. Yorks.: Streamside, Keysbeck,
Stape, SE/798.953. P. Sykes, 1991, det. F. Horsman. Only extant locality. ;

643/6 cam. DACTYLORHIZA MAJALIS subsp. CAMBRENSIS (Roberts) Roberts 46, Cards.: Fen
near Cardigan, SN/1.4. F. Horsman, 1991. 2nd record.

643/6 cam. X 5. DACTYLORHIZA MAJALIS subsp. CAMBRENSIS (Roberts) Roberts X D. PURPURELLA
(Stephenson & T. A. Stephenson) So6 *46, Cards.: Dune slack, Ynys-las Dunes N.N.R., SN/
609.939. D. C. Lang & A. P. Fowles, 1990. .

643/7. DACTYLORHIZA TRAUNSTEINERI (Sauter ex Reichb.) So6 *11, S. Hants.: Flushed marshy
clearing, Exbury, SU/4.0. R.P. Bowman, 1984, det. F. Rose.

4646/1. ACORUS CALAMUS L. 50, Denbs.: Marchwiel, SJ/355.468. P. Goodhind, 1991. 2nd
record. 80, Roxburghs.: Side of R. Teviot below Roxburgh Castle, NT/713.337. J. M. Croft,
C. D. Preston & O. M. Stewart, 1991, herb. R. W. M. Corner. Oniy extant locality.

4647/1. CALLA PALUSTRIS L. *47, Monts.: Roadside marsh, Meifod, SJ/160.126. H. Webster,
1989. Known for over 20 years but now almost overwhelmed by Petasites japonicus. 1st Welsh
record.

+648/1. LYSICHITON AMERICANUS Hultén & H. St John 46, Cards.: Stream bank, Tyglyn, SN/
498.598. R. N. Stringer, 1991. 2nd record.

650/4. LEMNA GIBBA L. 44, Carms.: Ditch north of Glanrhyd Farm, Pembrey, SN/406.043.
I. K. Morgan, 1991, NMW, conf. G. Hutchinson. Ist record since 1840s. Garden pond, Towy View
near Ffairfach, SN/640.225. I. K. Morgan, 1991. 2nd extant locality.

650/min. LEMNA MINUTA Kunth *44, Carms.: Ditch between Bury Port and Pwll, SN/462.012.
Reed swamp E. of Bury Port, SN/458.010. Both I. K. Morgan, 1991, NMW, det. G. Hutchinson &
A. Orange. Ist and 2nd records. 3rd and 4th Welsh records.

653/2 < 1. TYPHA ANGUSTIFOLIA L. X T. LATIFOLIA L. *13, W. Sussex: Ponds, Chichester
Gravel Pits, SU/872.032. G. H. Forster, 1991. “14, E. Sussex: Ditch in pasture near R. Ouse,
Lewes, TQ/428.055. T. C. G. Rich, 1991, LANC.

PLANT RECORDS 153
654/3. ERIOPHORUM LATIFOLIUM Hoppe 81, Berwicks.: Calcareous flush, Lamberton Moor,
NT/955.582. M. E. & P. F. Braithwaite, 1991, herb. M.E.B. 2nd extant locality.

655/11. ISOLEPIS CERNUA (Vahl) Roemer & Schultes 47, Monts.: Saltmarsh, Dovey Junction
Station, SN/695.984. P. M. Benoit, 1989. Only extant locality; last seen here in 1939.

656/6. ELEOCHARIS UNIGLUMIS (Link) Schultes *17, Surrey: Old sand filter bed, Barn Elms
waterworks, TQ/228.774. M. Mullin, 1991.

658/1. CYPERUS LONGUS L. +59, S. Lancs.: By dried-up reservoir, Pilsworth, SD/755.086.
A. Franks. 1991. 2nd record.

659/1. SCHOENUS NIGRICANS L. 81, Berwicks.: Calcareous flush, Lamberton Moor, NT/
955.582. M. E. & P. F. Braithwaite, 1991, herb. M.E.B. 2nd extant locality.

663/23. CAREX STRIGOSA Hudson *63, S. W. Yorks.: Damp ditch in wood, Margery Wood,
Cawthorne, SE/278.095. P. P. Abbott, 1991, det. W. A. Sledge.

663/28. CAREX LIMOSA L. 50, Denbs.: Soligenous flush, Pentrevoelas, SH/890.545. J. A.
Green, 1991. 2nd record.

663/60. CAREX DISTICHA Hudson *48, Merioneth: Phragmites swamp near Llanaber, SH/5.1.
P. M. Benoit, 1991, NMW.

663/61. CAREX ARENARIA L. +77, Lanarks.: Waste ground behind High Street Railway
Station, Glasgow, NS/59.65. J. H. Dickson, 1989, GL. 2nd record.

663/81. CAREX DIOICA L. *43, Rads.: Molinia flush below Craig y Bwich, SN/900.619.
D. Reed, 1991, NMW, conf. R. G. Woods.

667/1 aru. MOLINIA CAERULEA subsp. ARUNDINACEA (Schrank) K. Richter *58, Cheshire:
Roadside ditch, Goostrey, SJ/797.729. G. M. Kay, 1991, herb. G.M.K.
+ERAGROSTIS CURVULA (Schrader) Nees “11, S. Hants.: Side of disused railway near

Mayflower Park, Southampton, SU/416.111. E. J. Clement & A. L. Grenfell, 1989, herb. R. P.
Bowman, det. T. A. Cope.

1670/6 meg. FESTUCA RUBRA subsp. MEGASTACHYS Gaudin *25, E. Suffolk: Earth bank,
Landguard Common, TM/28.31. A. Copping, 1987, det. C. A. Stace. *29, Cambs.: Roadside
verge, Little Shelford, TL/446.513. P. J. O. Trist, 1991, herb. P.J.O.T.

670/7. FESTUCA ARENARIA Osbeck *44, Carms.: Shingle at foot of calcareous cliff, Llansteffan,
SN/3.0. I. M. Vaughan, 1967, NMW, det. A. K. Al-Bermani.

670/9. FESTUCA FILIFORMIS Pourret *47, Monts.: On hummocks in boggy field near Llanfihan-
gel-yng-Ngwynfa, SJ/09.15. P. M. Benoit, 1989.

670/10. FESTUCA VIVIPARA (L.) Smith 50, Denbs.: Rocky outcrop at 450 m, Migneunt, SH/
778.425. G. Battershall, 1991. 2nd record.

671/42 X 1. LOLIUM MULTIFLORUM Lam. X L. PERENNE L. *38, Warks.: Farm track, Upper
Shuckburgh, SP/486.603. J. C. Bowra, 1991, WAR, det. P. J. Copson.

672/5. VULPIA CILIATA subsp. AMBIGUA (Le Gall) Stace & Aug. *48, Merioneth: Low
hummocks in dune grassland, Aberdyfi golf links, SN/5.9 and 6.9. P. M. Benoit, 1991, NMW. Ist
Welsh record.

673/2. PUCCINELLIA DISTANS (Jacq.) Parl. *48, Merioneth: Embankment, Mawddach estuary
at Garth Isaf, SH/6.1. P. M. Benoit, 1991, NMW.

673/5. PUCCINELLIA RUPESTRIS (With.) Fern. & Weath. 2, E. Cornwall: Abundant on mud on
sides of Millbrook Pond, SX/425.523. R. W. Gould, 1991, conf. L. J. Margetts. Ist record since
1917:

676/11. POA ANGUSTIFOLIA L. 2, E. Cornwall: Penlee Battery, Rame peninsula, SX/440.493.
R. W. Gould, 1991, herb. R. J. Murphy, conf. L. J. Margetts & R. J. M. 2nd record.

154 PLANT RECORDS

+680/2..BRIZA MINOR L. “13, W. Sussex: Halsey’s Farm, Chichester Harbour, $Z/871.975.
G. H. Forster, 1991.

683/1. BROMOPSIS ERECTA (Hudson) Fourr. “43, Rads.: Grassland in base of old quarry,
Stanner Rocks, SO/26.58. R. G. Woods, 1988, NMW, det. T. G. Evans. One plant.

+683/4. BROMOPSIS INERMIS (Leysser) Holub *14, E. Sussex: Edge of Sports Field, Beving-
dean Hospital, TQ/331.061. A. Spiers, 1990, herb. P. A. Harmes. Chalk downs, Castle Hill,
Brighton, TQ/367.070. G. Steven, 1991, det. P. J. O. Trist. Ist and 2nd records.

+683/13. Bromus LEPIDUS O. Holmb. *44, Carms.: Forestry rides, Pembrey Forest, SN/3.0.
I. M. Vaughan, 1965, NMW.

+683/19. CERATOCHLOA CARINATA (Hook & Arn.) Tutin *2, E. Cornwall: Roadside verge,
Saltermill, SX/430.637. T. Atkinson, 1991. “11, S. Hants.: Sandy roadside verge, Belvidere,
Northam, SU/430.129. P. D. Stanley, 1991, herb. R.P. Bowman, det. R. P. B. & P. J. O. Trist.

684/2. BRACHYPODIUM PINNATUM (L.) P. Beauv. *52, Anglesey: Pine plantation on sand dune,
Newborough Forest, SH/398.637. N. H. Brown & R. H. Roberts, 1990, NMW.

687/1. HORDEUM SECALINUM Schreber +73, Kirkcudbrights.: Field track E. of Castlecreavie,
NX/726.488. O. M. Stewart, 1991, E, det. P. J. O. Trist. Ist record since 1910.

687/jub. HORDEUM JUBATUM L. 49, Caerns.: Sandy beach, Pontlyfni, SH/433.532. L. J.
Larsen, 1989, NMW, det. R. G. Ellis.

689/1. KOELERIA MACRANTHA (Ledeb.) Schultes 44, Carms.: Limestone grassland near
Llandyfan, SN/647.176. D. Grey & S. Gouch, 1989. 2nd extant locality. *48, Merioneth:
Calcareous dune grassland, Aberdyfi golf links, SN/6.9. P. M. Benoit, 1991, NMW.

700/1. CALAMAGROSTIS EPIGEJOS (L.) Roth = 111, Orkney: Tall herb community on cliff top,
Hoxa, S. Ronaldsay, ND/40.93. E. R. Bullard, 1989, det. P. J. O. Trist.

700/3. CALAMAGROSTIS STRICTA (Timm) Koeler *59,S. Lancs.: Old pool, Charles St, Darwen,
SD/690.227. P. Jepson & N. P. Symonds, 1991, det. H. J. M. Bowen.

701/2b. AGROSTIS VINEALIS Schreber *44, Carms.: Cerrig Cyffion, SN/68.46. R. Walls, 1988,
NMW.

+701/8. POLYPOGON viRIDIS (Gouan) Breistr. *52, Anglesey: Disused limestone quarry,
Benllech, SH/51.81. J. E. Hawksford, 1991, det. R. H. Roberts.

+702/1. APERA INTERRUPTA (L.) P. Beauv. *77, Lanarks.: Canal towpath, Maryhill, NS/57.68..
K. Watson, 1988, GL. Princes Dock, Glasgow, NS/56.64. A. McG. Stirling, 1988, GL. Ist and 2nd
records.

707/4. PHLEUM PHLEOIDES (L.) Karsten *25, E. Suffolk: Chalky hollow on golf course, Stuston
Common, TM/136.786. S. Hooton, 1991, conf. A. Copping.

719/2. DIGITARIA SANGUINALIS (L.) Scop. *2, E. Cornwall: Pavement weed, Saltash, SN/
432.587. S. C. & P. S. Madge, 1990.

Watsonia, 19, 155-164 (1992) 5

Book Reviews

Wild plants of Glasgow. Conservation in the city and countryside. J. H. Dickson, with paintings by
Elspeth Harrigan and photographs by T. N. Tait. Pp. 208, including black and white illustrations,
maps and tables, and 30 pp. in colour. Aberdeen University Press, Aberdeen. 1991. Price £14.95
(ISBN 0—08-041200-9).

This book is the first fruits of the Flora of Glasgow survey which has been taking place over the last
decade. It is aimed at the general reader, and will be followed later by a more detailed and technical
publication. It is an extremely attractive book, which describes some of the extraordinary range of
plants, native and introduced, growing in the Glasgow area, including something of their history and
ecology, and the problems of conservation.

The book is divided into three sections. The first deals with the background and objectives of the
survey, and gives an insight into how the data are analysed. Dot maps are used to illustrate some of
the different distribution patterns which have been identified, and annotated site lists to indicate the
characteristics of different plants.

The second and largest section of the book takes a more detailed look at nine contrasting habitats,
and at some of the plants typical of each. These include some amazing finds, such as the wild Fig on
the Clyde, and the ‘extinct’ Mudwort in a dried-up reservoir, as well as more common plants and
some interesting hybrids. The final section consists of a chapter on conservation, and is followed by
an extensive bibliography, and notes on Field Guides and Societies to join. In short it is a popular
book which also succeeds in being scholarly.

Since it is intended for a general audience, popular English plant names are used throughout.
Latin names are included for all the plants which receive detailed attention, but not in the captions
to the illustrations, which I think is a pity. Personally I like to see both Latin and English used at first
appearance even in more technical literature, but this would have made the text too unwieldy in
places. Latin plant names are included in the Index (though one or two plants on the site lists have
been missed).

But these are quibbles. The whole volume is a work of art and beautifully produced on expensive
paper; the colour photographs are excellent, and some of the individual flower paintings are quite
stunning. In his Introduction Dr Dickson bemoans the unattractiveness of the books available to
him as a youngster, when he first took an interest in botany: he has made sure that no future
generation of Glaswegians will be able to make a similar complaint. Let us hope he can pull off a
double with the more technical publication still to come.

J. Muscotr

The wild flowers of Luton. J. G. & C. M. Dony. Edited by C. Boon. Pp. 64, with maps. Privately
published, Luton. 1991. Price £3.50 (inc. postage), available from P. Ellison, 90 Beverley Road,
Ruislip, Middx., HA4 9AS.

This, the last work by Dr J. Dony in a line of distinguished books on his local flora, is announced as
“an account of the wild flowers known to grow in the immediate past in Luton’’. Having already
accounted in detail for the flora of Bedfordshire in general in two books, we might be excused for
thinking that this could not be anything new. We would be wrong. In fact, it is two accounts in one,
and breaks new ground in several ways. About a third of the book consists of an annotated check-list
of the flora of Luton Borough and its immediate environs, covering some 4,803 ha, an area with a
remarkable range of habitat types considering its size. The main body of the book, however, consists
of a series of concise ‘site reports’ of places which “‘had a vegetation worthy of record”. Some 23
sites are dealt with in detail, each having a description, historical notes, grid reference, and a list of

156 BOOK REVIEWS

specially selected species recorded between 1987 and 1990. These selected species are those found in
128 or fewer tetrads in Bedfordshire as a whole (i.e. less than 33%). A special innovation is the use
of these species lists to construct a ‘plant rarity factor’ for each site, the higher the resulting score,
the greater overall botanical rarity represented by the site. These accounts are rounded off neatly
with a clear site map for each site, and there is a coloured general map of the area in the centre
spread which can be used both to identify specific sites and to see the extent and location of surviving
semi-natural vegetation.

Just as Dr Dony’s earlier Floras had pioneered the use of discrete ‘habitat studies’ as a way of
providing a factual base-line for describing the typical vegetation of specific habitats, so this takes
the process one step further, to provide an outline assessment, in botanical terms, of the
conservation value of sites. The result is a mini conservation review of Luton Borough, based on
sound facts. Many a consultancy would be incapable of producing such a document for a local
authority’s ‘green audit’, so much the current fashion. Luton has one virtually for nothing.

T. J. JAMES

Fern names and their meanings. A glossary for the fern grower and collector. J. W. Dyce. Pp. iv + 31.
British Pteridological Society, London. 1988. Price £3 (ISBN 0—9509806-1-7).

‘What do they mean’? ‘Why do botanists keep changing them?’ This small booklet is an attempt to
answer, at least for “‘the amateur grower of British ferns and their varieties and cultivars’’, the first
of those questions so frequently asked about Latin plant names. The second problem is also
addressed briefly but passionately in the introduction.

The booklet has four main parts: an introduction, sections on the etymology of the names of

ritish fern genera, and on the meaning of Greek and Latin word elements commonly found in fern
varietal names, and the heart of the book, the main glossary of varietal names. The latter runs to 18
sicGes and includes over 700 entries, all listed in the neuter form.

The author indicates in his introduction that his basic source of information is the glossary of fern
names prepared by Dr F. W. Stansfield and Rev. Canon Kingsmill Moore and published in the
British Fern Gazette between 1919 and 1921. What he does not say is that his glossary is almost
entirely a simple re-ordering of Stansfield & Moore’s work (where the varietal names were arranged
under each species) into a single alphabetical sequence, but in nearly all cases keeping their original
definitions verbatim. One or two of Stansfield & Moore’s own, rare, errors have however been
corrected: their picturesque definition of gemmatum (‘decked with gems’) has been changed to the
prosaic but more accurate ‘provided with buds’. Some epithets are misspelt or malformed; in some
instances this is obviously due to error on the part of the original fern grower or author
(gracilissimum should be gracillimum, as indicated by Stansfield & Moore) but others (e.g. minum
and majum which should be minus and majus) show faulty understanding of Latin.

In spite of its deficiencies, this is an admirably useful little book. It makes Stansfield & Moore’s
work available once again in a handy, accessible and inexpensive form to a new generation of fern
growers and collectors who may not have the early volumes of British Fern Gazette at their disposal.

R. R. MILL

Atlas Florae Europaeae: Distribution of vascular plants in Europe. Vol. 9, Paeoniaceae to
Capparaceae. Edited by J. Jalas & J. Suominen. Pp. 110, 155 maps. Committee for Mapping the
Flora of Europe and Societas Biologica Fennica Vanamo, Helsinki. 1991. Price FIM 350 (ISBN
951—9108-08-4).

Volume 9 of Atlas Florae Europaeae is of particular interest to British botanists because, in covering
mainly the Papaveraceae, it deals with genera such as Papaver, Fumaria and Corydalis (sensu lato)
which have species with rapidly changing distributions in this country. Fumaria parviflora, for

BOOK REVIEWS 157,

example, is in decline throughout Central Europe and is hardly known north of 50°N except in
Britain where it reaches 57°N, and the same seems to be true of Papaver hybridum, now apparently
extinct in Belgium and Holland.

Many of the 24 British species mapped in this part are introductions but their treatment is
somewhat arbitrary. Chelidonium majus is accorded native status throughout its range here whilst
all records of Berberis vulgaris are regarded as introductions, whereas both are probably best
regarded as ‘doubtfully native’. Although a map of Eschscholzia californica is included, no British
localities are shown, though it is said by Stace in his New Flora of the British Isles (1991) to be
naturalised and perennating on dunes, walls and cliff tops in Guernsey and in quarries and by
railways in Kent. There is no map of the ever-spreading Oregon-grape, Mahonia aquifolium. This
variable treatment is doubtless a reflection of our own uncertainty and it is an area to which the
B.S.B.1., with its wide network of recorders, could surely contribute in advance of future parts.

Members may also be able to contribute useful information from visits abroad. I was surprised to
find the native status of Laurus nobilis in Majorca questioned: I saw it in February 1991 on a cliff at
the mouth of the Torrent de Pareis where it was recorded in my 1880 Flora.

The maps in this Atlas will be even more fascinating when put alongside the results of the
Monitoring Scheme when they are published. Comparison with that scheme will not be as easy as
could be wished because some of the recent name changes are long and unattractive. Corydalis has
been split into Pseudofumaria and Ceratocapnos, whilst with Fumaria, F. martinii has been lumped
with the more widespread F. reuteri, and orthographic research has decided that F. muralis now has
a subsp. boroei rather than subsp. boraei. There are times when one would like to recommend that
taxonomists be paid a negative productivity bonus.

With this volume the number of maps published since the project began in 1966 has reached 2109
and perhaps 15% of the task is completed. Now is surely a time to salute the efforts of those two
botanists from Helsinki, Jaakko Jalas and Juha Suominen, who have masterminded the project as
organisers and editors over the 25 years since it began.

F. H. PERRING

Wild orchids of Dorset. M. N. Jenkinson. Pp. 120, with drawings, maps and 63 colour plates. Orchid
Sundries, Gillingham, Dorset. 1991. Price £17.95 hardback (ISBN 1-873035—01-2); £13.95
paperback (ISBN 1—873035—02-0). |

This book, by a police officer, is said to be “directed not so much at the committed orchid enthusiast,
but seeking to find new converts to the faith”. Indeed much enthusiasm for the field study of orchids
is engendered here, even though the perils of looking for Hammarbya paludosa are made
abundantly clear.

Short introductory sections refer to the objectives of the book, the structure of the orchid flower,
the geology of Dorset, the main habitats of orchids in the county (listed under five categories), and
the nature reserves. Much of the book is devoted to accounts of some 28 species of orchids which
occur regularly. For each species information is given on status, habitat, flowering period,
distribution and distinctive features, but there is no overall key. The distribution maps, based
mostly on records made during the last ten years, provide more information than previously
available, presence being shown within 1-km squares. Although readers are repeatedly reminded
not to pick flowers, the wisdom of the publication of such detailed distributions of uncommon
species is questionable.

Notable features are the full treatment of the helleborines (including the recent discoveries of
Epipactis purpurata) and of the dactylorchids, both subjects of special study by the author. In the
dactylorchids, nomenclature departs from Flora Europaea, and ‘‘a newly identified form, var.
bowmanii Jenkinson” (not validly published) is included under Dactylorhiza majalis subsp.
traunsteinerioides. This form, found also in Hampshire, is robust, has a rather dark flower colour
and a fairly narrow, very deeply three-lobed labellum. Hybrids are not prominently treated, but the
features of the labellum of many forms of the marsh orchids are illustrated.

There are a few unfortunate statements, e.g. ‘““physiological”’ features of the landscape, Epipactis

158 BOOK REVIEWS

palustris spreading by underground “runners, or elongated roots’, and one may doubt whether |
there would ever be ‘‘climax pine forest” in the New Forest. However, the valuable points for field
identification, and details of the phenology and occurrence, together with the original colour
photographs (for each species a general view and a close-up of flowers, mostly of good quality),
make this a useful and attractive book, particularly for those exploring the unspoilt countryside of
Dorset.

A. J. WILLIS

The Chelsea gardener: Philip Miller 1691-1771. H. Le Rougetel. Pp. 212, illustrated in black and
white and colour. Natural History Museum Publications, London. 1990. Price £14.95 (ISBN 0—565-
01101-4).

Philip Miller is, without doubt, the most celebrated English gardener of the eighteenth century. He
was a Sself-educated man, and what he could not learn from his father or other gardeners he learnt
from books. Having set up his own nursery in Southwark, he was already established in the small
circle of elite gardeners, when, in 1722, he was appointed Gardener (in today’s terminology,
Curator) of the Society of Apothecaries’ Physic Garden at Chelsea. He was soon to put the garden
on to a sound footing and to establish over the next 48 years an institution of international repute.

This is a book not just about Philip Miller and his achievements at Chelsea; it is a masterly analysis
of Miller’s influence on gardening and horticulture — and they were by no means synonymous in
those days. The author has arranged her account in 20 interlocking chapters, the first three giving
background, others discussing Miller’s links with botanists abroad, in Europe and North America,
with eminent gardens and gardeners, plant illustrators and the like. His links with Cambridge and
the effects his views were having on forestry and agricultural policy are reviewed; and the
importance of Philip Miller’s published works to botany at large is discussed in Chapter 20 by
William Stearn. His most important work, the Gardeners’ Dictionary, which went through eight
editions between 1731 and 1768, contained cultural information for the kitchen, flower and fruit
garden, and much descriptive botanical material beside. The last edition appeared after the
publication of Linnaeus’ Species Plantarum in which binomial nomenclature was introduced, and
Miller followed the convention and thereby validated many pre-Linnaean genera he had used in
earlier editions.

Miller was also a field botanist and a teacher, collecting in many parts of England from the
Cheviots to his home county of Kent. He was able to find the Deptford Pink (Dianthus armeria) in
Deptford, Aquilegia vulgaris in Bexley and Narthecium ossifragum on Putney Heath.

This book is a ‘must’ for anyone interested in the history of botany or gardening and the kind of
book that will be referred to time and time again. Hazel Le Rougetel is to be congratulated in giving
us such a delightful, concise account of what must have been many years of fascinating research.

A. C. JERMY

Shamrock, botany and history of an Irish myth. E. C. Nelson. Pp. xiv + 200 with 5 colour plates and
numerous black and white photographs and illustrations. Boethius Press, Aberystwyth & Kilkenny.
1991. Price IR£28 (ISBN 0-86314-200-1 hardback); IR£14 (ISBN 0-86314—199-4 paperback
without bibliography, pp. xiv + 158); now only available from the author, 14 Connaught Parade,
Dublin 7; p&p extra.

The curtain rises on a sentimental discourse of the importance of shamrock to the Irish, resolving to
strip away the facts from the fiction.

Act I opens on an Ancient Briton, St Patrick, himself of whom little is known, and who certainly
has nothing to do with shamrock! Indeed as we pass through the centuries nothing is heard of this
blessed plant in Latin, Irish or English, until we realise that shamrock is the anglicised version of
seamrog meaning a young clover. Variations on scothsheamrachor or clover-flowered thus abound

BOOK REVIEWS 159

in mediaeval manuscripts. The first time the word shamrock made its appearance was in Edward
Campion’s The first boke of the histories of Irelande (1577) where he stated that shamrocks were
eaten by the Irish!

By Act V we have caught up with the Herbalists where in 1571 Matthias de l’Obel wrote a strange
passage about the Irish passion for a meal of clover or meadow trefoil. This was followed by John
Gerard in 1597 who understood that Shamrock was clover and that seamr6ég covered both the red
and white clovers. By the early 1600s, it was the accepted view in England that shamrock was eaten
daily by the Irish or only in times of dire necessity (depending on the authors’ own prejudices).

Some decades later, seamrog was being defined as wood sorrel in an Irish dictionary. It was not
until 1726 that an Englishman, the Rev. Dr Caleb Threlkeld, set the record straight, recognising the
tradition around St Patrick and the shamrock as an explanation of the Holy Trinity, and also that
seamrog were clovers. He was also the first to record the now renowned ritual of drowning the
shamrock with alcohol and food.

By Act XI, the shamrock was beginning to be worn not just on 17 March but as a sacred political
badge throughout the year. The Volunteers were the first to use it in 1779, followed by the United
Irishmen in 1791. The shamrock motif was seen on flags, belts, cockades, glassware and seals, even
in a weekly journal The Shamroc and in verse “The Wearing of the Green’.

With the dawning of a new century, the romantic, mawkish image of the shamrock blossomed
forth. At the same time the ‘Age of Reason’ brought the actual botanical identity of the shamrock
into question. The first scientific survey of what plant was worn in Irish buttonholes was undertaken
by James Britten in the 1870s. He discovered that the plant mostly in use as the true shamrock was
Trifolium minus (now called T. dubium). Unfortunately, not many people took note of his
systematic findings. Indeed, ten years later, one Nathaniel Colgan added to the confusion by
establishing all the above plants as rivals, plus the red clover (T. pratense).

When the Currency Commission of 1926 decided to ban the shamrock motif from the new coinage
there was a mass uproar and by the time the Irish Post Office issued the first definitive stamps and
one commemorative one in 1933-4, each one had shamrocks in the design.

And so we finally come to the late 1980s, when Charles Nelson attempted to assess the present day
status of the shamrock in Ireland by means of modern communications. In fact Trifolium dubium
was revealed as the most commonly regarded true shamrock, while black medick (Medicago
lupulina), red clover, and wood sorrel (Oxalis acetosella) accounted for only a few gatherings. The
curtain closes on the fact that “‘shamrock is a young clover, nothing more, nothing less’’. That is
what it always meant ‘“‘and what it will mean until the end of time.”

Had this book been written by anyone else, it might have been both boring and pedantic, but Dr
Nelson’s sparkling dry wit and wicked sense of humour comes through again and again.
Accompanied by some delightful colour plates by Bridget Flinn and a hilarious and irreverent
forward by Bernard Loughlin, this book deserves to be read by a wide audience. Vivat trefolium!

S. ANDREWS

The Burren. A companion to the wildflowers of an Irish limestone wilderness. E. C. Nelson & W.
Walsh. Pp. 343, with 100 colour and several black and white illustrations. Boethius Press,
Aberystwyth, and The Conservancy of The Burren. 1991. Price IR£32 (ISBN—86314-213-3
hardback); IR£19 (0-86314-214-1 paperback); available from 14 Connaught Parade, Dublin 7;
pé&p extra. -

The Burren of north-western Co. Clare is Ireland’s premier botanical attraction. Remarkably, it
was first explored in detail only in 1851, the year that T. H. H. von Heldreich revealed Greece’s Mt
Olympus in remotest Thessaly to the botanical world. The two areas are actually very comparable,
for they are both karstic landscapes of bare limestone, subterranean water and few human
inhabitants, making travel difficult in the days before motorised transport. Each has a rich and
diverse flora and is amongst the most precious gems of Europe’s natural heritage. My own view
(coloured by my affection for Connemara) had long been that the Burren received too much
attention from botanists, especially those visiting from Britain, to the detriment of the study of other

160 BOOK REVIEWS

regions of Ireland. However, reading The Burren has successfully prodded me to take a positive
view of the region, the book stirring up as it does images of low, mysterious, grey hills, pastures
bright with Spring Gentians in May, and the massed flowering of Mountain Avens, Bloody
Cranesbill, Hoary Rockrose and other species in the native rock garden that the Burren presents to
the visitor in summer.

Charles Nelson sets out an eloquent, enthusiastic and erudite portrait of a landscape and its
plants. He provides background information to the flowers that make the Burren so special, gleaned
from a wide variety of sources, whilst maintaining an unobtrusive personal touch by allusions to his
childhood in Co. Fermanagh, his interest in tropical drift-seeds and the genus Fuchsia, and his
knowledge of Irish garden history. The style is discursive, the text packed with observational
sketches, anecdotes and quotations from the literature of botany, folklore and Irish history. This is
not a new Flora of the Burren: there are many references to David Webb and Maura Scannell’s
Flora of Connemara and the Burren (1983) — reviewed in Watsonia 15: 148-9 (1984). Nor is it a
guidebook; for topographical information the reader is encouraged to refer to Tim Robinson’s
excellent annotated map, The Burren, a map of the uplands of North-West Clare, Eire (1977). It is
indeed a companion, a book, in the author’s words, to be read by the fireside when “‘a westerly gale
hurls stair-rods of rain horizontally across Cappanawalla’’. It will certainly be a useful item in one’s
hotel or ‘bed-and-breakfast’ during a visit. I readily forgive the author for ignoring the mundane
plants that excite feeble minds like my own - there is, for example, no mention of any docks or
knotgrasses!

The author defines the region more or less on the basis of the (limestone) geology, although he is
sensibly a bit vague about the eastern limits around Kinvarra and Gort, where one can still see a
‘good’ Burren flora. The text is beautifully set out, the words interspersed with water-colour
illustrations by Wendy Walsh, deliberately conceived in a sketch format to give a sense of
immediacy. There are also black and white topographical scenes and some portraits. I should have
been happier with numbers as well as titles at the head of each chapter. Nor am I am at home with
the Irish word scailp to describe the deep crevices in the limestone; in Yorkshire, where my family
came from, they are grykes (as they are in Scandinavia).

Charles Nelson has done us all a considerable favour by his distillation of fact and experience, and
I urge the many lovers of the Burren to buy this fairly priced book, not least because all proceeds will
go to support the work of The Conservancy of The Burren. The future of this special place is
uncertain, the worst threat being increased pressure from tourism, notably the proposal to erect a
superfluous and environmentally insensitive ‘visitor centre’, and it is up to botanists above all to
promote its conservation. This book is a beginning.

J. R. AKEROYD

The northwest European pollen flora, Volume 6. Edited by W. Punt & S. Blackmore. Pp: v4 2755
with 103 black and white plates. Elsevier, Amsterdam. 1991. Price D.FI. 240 (ISBN 0-444-891641—
1).

This latest volume in the northwest European pollen flora project continues to document
systematically the pollen morphology of all families of vascular plants indigenous to, or regularly
naturalized in, northwest Europe. In this volume the families covered are Selaginellaceae,
Oleaceae, Geraniaceae, Juglandaceae, Cornaceae, Globulariaceae, Buxaceae and Ranunculaceae,
with the last family occupying more than half of the book. Each family account contains detailed
descriptions of pollen types (i.e. the morphologically discernible groups of pollen species) together
with keys to their identification, and is accompanied by excellent scanning electron and light
micrographs.

The arrangement of the survey is taxonomic and its clearest aim is to provide taxonomic
information on the range of form of pollen within families. But pollen identification has become an
important aspect of the work of many other plant scientists in disciplines as varied as allergy studies,
forensic science and the reconstruction of past vegetation and hence past environments. Palynolog-
ists with these interests will look to the Pollen Flora as a valuable source of data on pollen

BOOK REVIEWS 161

morphology, but the material contained in these volumes has to be used with caution in these areas.
The palaeopalynologist, for example, when faced with an unknown pollen grain, does not normally
have the assurance that it belongs to a particular family. A spiny trilete spore, such as that of
Selaginella kraussiana (G. Kunze) A. Braun as described in this volume, could also belong to a
bryophyte, such as Anthoceros or Riccia. Some pollen grains belonging to members of the Oleaceae
are very similar to those of certain Cruciferae; but such similarities cannot, of course, be dealt with
in a systematically arranged account.

The book will be of greatest value to those concerned with pollen identification when detailed
information and illustrations are needed to confirm identity or to provide greater taxonomic
resolution in important groups. The Ranunculaceae is of particular note here because most general
pollen keys provide limited taxonomic detail, yet distinction between ranunculaceous groups can be
of profound ecological value for the palaeoecologist. The ‘Ranunculus acris type’ pollen recognized
by most palynologists, for example, covers a multitude of ecologically varied species, and the
information provided here for more precise identification will be very welcome.

The keys are generally robust, though some expressions are loose and ambiguous, such as the
distinction between “reticulum relatively coarse” and “reticulum fine”’ in the distinction between
Olea and Phillyrea. This could have been improved by providing a count of lumina density across
the mesocolpium, which permits very adequate separation. I remain unconvinced whether some of
the fine distinctions, such as that between Erodium moschatum and other members of the genus, can
be effected by light microscopy alone. In the identification of fossil pollen using conventional phase
contrast microscopy, the work described in this valuable collection of papers will prove useful, but
must clearly be applied with great caution and always with the assistance of reference material.

P. D. Moore

New Flora of the British Isles. C. A. Stace. Pp. xxx + 1226, illustrated. Cambridge University Press,
Cambridge. 1991. Price £24.95 (ISBN 0—-521—42793-2).

This eagerly awaited and reasonably priced book fulfils almost all expectations and is essential for
anyone with a serious interest in the British and Irish floras. It is a manual for identification rather
than a descriptive Flora, and so needs to be judged for its usefulness rather than just the information
it contains. I make no apology for repeating the words ‘helpful’ and ‘useful’ rather often in this
review.

Everything in the Flora, from the taxa included (especially non-native taxa) to the nomenclature
and the construction of the keys has clearly been worked out afresh. Taking on trust its claim to
contain 2990 keyed and numbered species, it thus contains nearly half as many again as the 2030 that
are in the 3rd edition of the Flora of the British Isles by Clapham, Tutin & Moore (CTM). This gives
some idea of the amount of unfamiliar materials made available by Stace. Most of the extra species
are aliens, and in some genera the result is quite startling, Cotoneaster containing 45 keyed and
numbered taxa (five in CTM), Crocus containing nine (two in CIM) and even Trifolium 32 (22 in
CTM). There are 90 more grasses in Stace. His criteria for inclusion of non-native taxa are
eminently helpful, the aim being “‘to include all taxa that the plant-hunter might reasonably be able
to find ‘in the wild’ in any one year. Any such plant, whether native, accidentally introduced or
planted, affects wild habitats and is part of the ecosystem, and botanists and others might be
expected to need or want to identify it”. He thus includes a number of plants that persist rather than
regularly naturalise, as well as a wide range of non-persistent but recurrent casuals.

The dichotomous keys seem to work well, and many are quite original. The Carex key, for
example, broken up into sections like all long keys, differs in many ways from previously available
ones, often uses quite different characters, and works at least as well. I personally dislike the layout
of the keys, with alternately indented couplets, but one soon gets used to it. Multi-access keys are
often provided, but in only a few cases (notably Epilobium) are they the only sort available. The
ones in Sorbus, and in the novel account of the cultivars of Populus X canadensis, are especially
useful. The species descriptions are mostly less than 30 words, and have a limited value for
confirming identifications. As so many unfamiliar species are included for which fuller descriptions

162 BOOK REVIEWS

will be unavailable to most readers, this does create problems. The drawings, mentioned later, do
however somewhat compensate for this. With more familiar species, too, the brevity of the
descriptions can be worrying. Is there, for example, no mention of the hairs at the junction of leaf-
blade and sheath in Poa humilis (P. subcaerulea) because the author considers them not to be
diagnostic, or because there is just not the room to include this character?

Abbreviations are few and easily understood. The text in general contains an immense amount of
useful comment. Unnumbered and unkeyed “other genera” and “‘other species” are frequently
included, but I cannot help feeling that, as in most other Floras, they do more to salve the author’s
conscience than to help the reader. They stand out here because the rest of the Flora is so helpful.
Crop species are especially well treated and up-to-date. Red Data Book, scheduled, and Rare
species (in fewer than 100 10-km squares) are indicated. English names, many of them new, are
provided for all the numbered species. The bibliography is very brief, and it is perhaps a pity that no
direct references to more detailed accounts are given anywhere in the text.

The taxonomic stance of the book is a good balance between expert and consensus opinion. It is _
comforting to find one’s doubts about identifying segregates of such species as Galium mollugo,
Rhinanthus minor and Sedum telephium confirmed, but I am sorry to see the subspecies of Pilosella
officinarum so briskly dismissed (many of us record them, even though we may doubt whether they
are worth subspecific status). For once in a British Flora though, subspecies are generally very fully
treated. Varieties are rarely and rather unpredictably included. In the case of Fumaria the key
would probably have worked better if varieties had been included (as P. D. Sell, BSBI News 41: 16
(1985) recommends), and this is one of a number of cases where recourse to the Plant Crib is still
essential. All hybrids known to occur in Britain and Ireland are included. More are keyed and
described than in our other Floras, but inevitably, alas, the great majority are not. Here, as
elsewhere, the continuing need for a comprehensive critical and descriptive Flora of Britain and
Ireland is highlighted. Of the apomictic groups, Rubus, Hieracium and Taraxacum are treated only
in outline, but Sorbus and Euphrasia are treated in full.

Physically the book is about the same size as the 2nd edition of Clapham, Tutin & Warburg’s
Flora. The binding is sewn, and the cover is pliable plastic. My copy is still in perfect shape after two
months’ daily use. It is printed from camera-ready copy produced by a word-processor, and the type
is large and easy to read (though lacking such refinements as accents and italics). Traditional
typesetting would probably have reduced the number of pages by about a third and made the book
more suitable for the rucksack, but, as the author remarks, would even so have increased the price.
Errors of any sort are minimal. Main Argyll is given for Rorippa X hungarica instead of R. X
armoracioides. Veronica hederifolia should be cross-referenced to the illustrations instead of V.
serpyllifolia. The drawings of Ledum and Vaccinium uliginosum are labelled the wrong way round.
Salsola is missing from the index. The second half of dichotomy 8 on p. 564 should lead to 17 not 16.
Otherwise there seem few errors likely seriously to mislead the reader. Even the cm scale inside the
front cover is only 1% out. The brevity of the distributional information occasionally leads to slight
inaccuracy, but this is almost inevitable and detail of this sort is not what one will chiefly come to this —
Flora for.

There are some 150 pages of illustrations, and it is here that my only serious criticisms of the book
lie. The line drawings are mostly by Hilli Thompson and include excellent series of crucifer,
umbellifer and Rumex fruits. Her habit drawings of unfamiliar aliens are especially helpful and
compensate a good deal for the brevity of the description. Most give a good impression of the jizz of
the plants but some, for example those of Cotoneaster, are curiously diagrammatic, and several of
the series of drawings of leaves are unsatisfactory. In a few cases poor choice of specimens rather
than quality of drawing means that they fail to show, or even contradict, the diagnostic characters,
for example the number of intercalary leaves and angle of branching of Odontites vernus subsp.
serotinus, the leaf-lobes of Ranunculus omiophyllus, or the apical leaf-lobes of the Veronica
hederifolia subspecies. Other artists have also contributed vaiuable drawings, such as those of
Dactylorhiza \abella by R. H. Roberts and Oenothera flowers by J. Zygmunt. The many light —
photographs often seem to suffer from very poor reproduction, and it is difficult to see why the
blurred Potamogeton \eaf-apices were not drawn instead. The Euphrasia silhouettes are too small
and murky to be of much use, but those of Sorbus are excellent. There are also many scanning
electron micrographs (S.E.M.). Those of Epilobium seeds and Isoetes megaspores are good, and
the latter for once show that the sculpturing is not as easily diagnostic as most descriptions imply, but

BOOK REVIEWS 163

most of the rest are unsatisfactory. Even the very clear ones of Montia seeds, by the very nature of
the process, omit the diagnostic shininess or otherwise of the coats. Those of Tripleurospermum
achenes fail to show even the oil-glands, let alone their diagnostic shapes, and the complete series of
Carex utricles includes many unrecognisable and often shrivelled examples. S.E.M. pictures are
generally unhelpful as identification aids for botanists with a lens or light microscope, and with an
artist of Thompson’s ability and versatility to hand it is a mystery why they were used.

The New Flora was prepared in close consultation with D. H. Kent who was simultaneously
compiling a new nomenclatural check-list of British and Irish plants. Thus the nomenclature, like
every Other aspect of the Flora, is uncompromisingly up-to-date according to current knowledge,
resulting in a large number of changes to familiar names. These will best be reviewed when the
check-list itself is published. Meanwhile the reader’s dismay at them should be largely offset by the
fact that they are an integral part of an authoritative and user-friendly Flora that will be our standard
for taxonomy, nomenclature and identification for some time to come.

A. O. CHATER

Pleistocene palaeoecology of central Norfolk — a study of environments through time. R. G. West.
Pp. ix + 110, with 44 figures. Cambridge University Press, Cambridge. 1991. Price £40 (ISBN 0-
521—40368-5S).

This book continues the Pleistocene history of East Anglia from the author’s earlier book The pre-
glacial Pleistocene of the Norfolk and Suffolk coasts, concentrating on a part of the catchment of the
River Wensum near East Dereham. Within this area, sections exposed by commercial sand and
gravel workings over a period of c. 20 years have provided “‘an extraordinary wealth of evidence” on
environmental changes from the time of the first major glaciation of East Anglia. The presentation
of this evidence is organised chronologically. Five chapters deal with the local succession of deposits
identified as belonging to the Anglian, Wolstonian and Devensian cold stages and the Hoxnian,
Ipswichian and Flandrian temperate stages. The geological evidence and palaeobotanical analyses
produced by Professor West’s intensive studies are presented in meticulous detail in the form of
many clear sections (related to aerial photographs as well as maps), pollen diagrams, and tables of
fossil identifications. These numerous sections include glacial tills and gravels and overlying
fluviatile deposits with limnic sediments formed in depressions. His synthesis of this evidence
supports — through the relative positions of deposits identified by their palaeobotany — the
succession of Middle and Upper Pleistocene stages originally proposed for East Anglia by Professor
West and co-workers. At the same time he reiterates his caution that it remains possible that
additional stages may yet be identified in the terrestrial record.

The form of the book is that of an extended scientific paper. It furnishes an excellent example of
how primary data of the highest scientific value have been gathered, processed and synthesized to
provide the basis for reconstruction of past environmental — including climatic — changes. Not only
does this treatment fulfil the author’s claim that “‘the observations have led to a much better
understanding of the Pleistocene in Norfolk and are indeed relevant to the wider understanding of
the British Pleistocene” but it provides for the more general reader an insight into the methods by
which primary data are obtained and used to reconstruct climatic changes in the past. These
reconstructions can then be of use in comparison with models of possible climatic change in the
future. This book is indeed, as claimed on the fly-leaf, ‘‘a unique ‘case study’ of an investigation of
past climatic change’’.

Throughout the work there is a new emphasis on the importance to the palaeobotanist of
thorough acquaintance with the processes involved in accumulation of each sediment type, in order
to reach sound conclusions as to the taphonomy of fossil plant assemblages, whether pollen, spores
or larger fossils. The influence of this on the relationship between the fossil assemblage and the
vegetation from which it was derived is considered throughout. The presentation of the chronologi-
cally arranged botanical evidence is followed by two chapters on stratigraphic questions, related
especially to periglacial conditions during the cold stages and to inferred changes in water levels. A
further chapter discusses, in a review of the history of the Whitewater, Blackwater and Wensum

164 BOOK REVIEWS

valleys, the aggradation of the Beetley Terrace during a non-glacial post-Hoxnian cold stage. This
preceded the Ipswichian and therefore seems to coincide in time with the Wolstonian. The final very
concise chapter placing the work at Beetley “in the context of the East Anglian Pleistocene” should
be required reading for all students of the British Quaternary.

The reference list is of a realistic length to encourage further study. Hopefully the work as a whole ~
will stimulate others to undertake this kind of devoted fieldwork and patient study-in-depth in
promising areas.

W. TUTIN

Watsonia, 19, 165-168 (1992) 165

Obituaries

ADRIAN LEONARD GRENFELL
(1939—1991)

Adrian Grenfell was born on 6 April 1939 and died, aged only 52, on 17 November 1991. It was a
great shock to hear from Adrian’s wife, Diana, that my good friend had died of a heart attack whilst
jogging. We had all known that he was not in the best of health, but none of us realised just how ill
he must have been.

He and I have known each other for many years. We both attended the same Grammar School
together, although at that time Adrian was not interested in botany. He left school to start a career
with the Gas Board, later on changing to become an industrial chemist with a major Bristol
company. Redundancy in the early 1980s allowed him to become a self-employed printer and later
he moved into publishing. During the last seven years his expertise in printing and publishing was
put to good use on behalf of the Bristol Naturalists’ Society, the Wild Flower Society and the
Botanical Society of the British Isles.

He became interested in plants in about 1972 (reference to the B.S.B.I. list of members shows he
joined in 1976). At the same time he joined the Wild Flower Society and the local Bristol
Naturalists’ Society. He was keenly interested in mammals as well as plants and would often attend
meetings of the Mammal Section of the Bristol society. In the early days alien plants became his
main interest, brought on possibly by his friendship with Eric Clement, and Adrian became what is
known as a “Tip Man’. He and I, often in the company of other botanists including Clive Lovatt and
Trevor Evans, would frequently visit such sites as Brislington Tip, Bedminster Tip, Avonmouth,
Newport and Sharpness Docks, and other localities where aliens were to be found. Reference to the
‘Bristol Botany’ section in Proceedings of the Bristol Naturalists’ Society and B.S.B.I. News shows
the many interesting finds made during that period. He had to wait a long time to get into the now
Portbury Docks — so jealously was it watched. We were making comparisons all the time with N.Y.
Sandwith’s 1933 adventive plant list from the Port of Bristol — at one time Adrian intended to
publish an updated check-list.

On the local scene botanists eagerly looked forward to Adrian’s walks in the Bristol area. He
would lead us round such famous localities as the Avon Gorge, Leigh Woods, Brean Down and the
Mendip Hills. He was always pleased to help both the ‘starters’ and the more experienced botanists
with determinations. Adrian’s hybrids caused some amusement: so often, however, they were
confirmed by the correct authority. He truly had a remarkable eye for the unusual.

In the early 1980s Adrian and I were to enjoy several trips abroad to look at plants, sample local
wine, etc. I can always remember Adrian one evening dropping off to sleep in a chair after a hard

_day’s collecting and then determining in Portugal, with a volume of Flora Europaea in his lap and a
glass of wine in his hand. After that we decided that after one bottle in the evening all
_ determinations were suspect! We enjoyed trips to Crete, the Algarve, Mallorca and Scotland; later
on Adrian went to Zakinthos and Mt Olympus in Greece, the Alps (where he led B.S.B.I.
excursions), the Seychelles and Australia. All the time he was looking for interesting plants and
would always bring back something, usually grasses for his friends Ron Payne and Eric Clement.
_ From April 1983 Adrian took over the Alien News section in B.S.B./. News from Eric Clement,
_ editing it for eight years (see B.S.B.J. News 32-57). Adrian would look forward to his mail and
| would eagerly open packages of carefully pressed specimens and examine the soggy contents of
, polythene bags to see what was new. Many of these specimens have been kept and are preserved in
| his herbarium. Adrian was at the same time helping the Society in other ways. In 1983 he was on the
Meetings Committee and helped to organize the A.G.M. in Bristol, arranging accommodation and
the field excursions to the Mendips and Sand Point in Somerset, a very memorable day. In April
/ 1991, his successor as editor of Aliens and Adventives, Brian Wurzell, thanked Adrian for his ©
‘valuable contribution and wished him a speedy recovery from the illness which caused him to resign
' from the work. Sadly, Adrian only lived a further six months.

_ In 1979, one year after Early Star-of-Bethlehem (Gagea bohemica) was confirmed as occurring at

166 OBITUARIES

Stanner Rocks in Radnorshire, Adrian, Trevor Evans and myself went to the site, on a wintry
February day with snow in the air, and were rewarded by the sight of two flowers of this plant new to
the British flora. We knew that the next weekend Mary Briggs and a party of eminent botanists were
to visit the site — so new was it to the British flora. Adrian decided to leave a note addressed to Mary
under a stone to record that we had been there and to advise where the flowering plants were to be ©
seen. Mary found the note. Adrian later remembered this occasion when he found the same plant
high up above the Samaria Gorge in Crete.

In 1987, Adrian wrote a paper on the notable alien flora of the Avon Gorge, which was included in
The Avon Gorge, Special Issue No. 1 of Proceedings of the Bristol Naturalists’ Society. This was
elegantly produced by Grenfell Publications and offered for sale to B.S.B.I. members. Other
publishing ventures included several editions of the Proceedings, the Supplement of the Flora of
Gloucestershire by S. C. Holland, H. M. Caddick and D. S. Dudley-Smith (1986) and The difficult
and critical plants of the Lizard District of Cornwall by L. J. Margetts (1988). Every year Adrian
would send Professor A. J. Willis, the Bristol Botany recorder, many interesting records — all these ©
can be found in the Proceedings or in B.S.B.I. News. In the autumn of 1991, Adrian and I had just
finished an article on Bristol street trees, which will be published in the Proceedings in due course.
We had great fun driving up and down the streets of Bristol, identifying, collecting and tetera ie
material of a vast range of trees.

There is one plant that will always remind me of Adrian. It is not an alien but the British native,
Autumn Lady’s-tresses (Spiranthes spiralis). It was found by him growing on Eric Clement’s
Gosport lawn — and Eric had never seen it. Adrian always said with a grin that it would have to be
flat or mounted on a sheet for Eric to notice it!

On the basis of his work on alien plants, Adrian was elected a Fellow of the Linnean Society of
London in 1982. His private herbarium will be transferred to Bristol City Museum (BRISTM) and
kept there alongside those of other distinguished Bristol botanists such as I. W. Evans and J. W.
White.

Over the years Adrian built up a tremendous number of friends, many of whom stayed with him
when they visited the Bristol area. He was always willing to help in any way that he could. Adrian’s
warm friendship and dry humour will be missed by us all, particularly when we revisit a site where he
had shown us a speciality. We extend our sympathies to Diana and his son James.

T. TITCHEN

WILLIAM ARTHUR SLEDGE
(1904—1991)

Arthur Sledge was a Leodiensian. He was born in Leeds on 14 February 1904 and died there on 15
December 1991, having lived there all his life. He was educated at Leeds Grammar School and
graduated from the Botany Department of Leeds University in 1926. He gained his Ph.D. in 1928,
the year he joined the staff of the department as a Demonstrator. He was appointed as a Lecturer a
year later and eventually became a Senior Lecturer. After his official retirement in 1969 he retained
a room in the department and continued his work there as Honorary Research Fellow, usually
walking the 3 km from his home in Headingley. A severe heart attack two years ago restricted his
physical activities but he bore the resulting debilities uncomplainingly and with fortitude.

His interest in botany was triggered at his junior school by a pressed flower competition, which, of
course, he won. While he was out collecting specimens, his vasculum was spotted by Frank Paimer
who was taking part in a similar senior school competition. Frank and his father subsequently —
introduced the young Sledge to the joys of botanising on the Permian Magnesian Limestones a few
kms east of Leeds. From this was born an awareness of habitat and an interest in ecology and he was
later to become a founder member and council member of the Yorkshire Naturalists’ Trust (now the
Yorkshire Wildlife Trust).

Whilst still in his early teens, Dr Sledge had found F. A. Lees’ Flora of West Yorkshire (1888) in

OBITUARIES 167

the library and, learning that Dr Lees lived quite nearby, he became a frequent visitor to his home,
armed with eggs from his mother’s hens and a vasculum of plants for identification. In 1941, A
Supplement to the Yorkshire Floras by the Late F. Arnold Lees, edited by C. A. Cheetham and W.
A. Sledge, was published.

In 1920 Dr Sledge was introduced to Leeds Naturalists’ Club by the blind botanist, John
Wilkinson. Active members of the club at that time, who all helped to further Dr Sledge’s interest,
included such famous names as J. H. Priestley, Professor of Botany at Leeds, W. H. Pearsall,
Edward Percival, R. W. Butcher, and the bryologist, W. H. Burrell.

It was standard practice at Leeds Naturalists’ Club to report on Yorkshire Naturalists’ Union
meetings and Dr Sledge was inspired to join the Union. All the most knowledgeable naturalists in
the county belonged to the Union and from them much was gleaned. Later he more than repaid
what he had gained. In the field he was the ultimate authority who could be relied upon to give a
correct identification to all but the most esoteric of critical plants, along with interesting comments
on habitat and plant associations. His memory for sites of interest was unfailing, even those he had
not visited for 40 years and, when he felt there was good reason to give directions to them, these
directions were precise and accurate. All of us were in awe of him, but those who showed an
intelligent interest and desire for knowledge were subsequently greeted with a warm smile and
questions were answered fully, with quiet enthusiasm. In addition to sharing his experience in the
field, Dr Sledge edited the Union’s scientific publication, The Naturalist, from 1943 to 1975. His
dedication and skill resulted in The Naturalist being read and respected nationally and internation-
ally and, in spite of his modest, unassuming manner, his pride in it and his paternalism towards it
were evident.

He joined the B.S.B.I. in 1924 and was appointed Recorder for South-east, South-west and Mid-
west Yorkshire in 1949. He relinquished South-east Yorkshire to Eva Crackles in 1969 and
continued with South-west and Mid-west until 1987, when failing eyesight made him realise he
would not be able to cope with the Monitoring Scheme. In 1987 he was made an Honorary Member
of the Society for his “‘long and valuable service to Yorkshire botany and to the Society as Recorder
and Editor’’. He had edited the Distributor’s Report (Exchange Section), which was a supplement
to the Society’s Year Book, in the late 1940s. With our present emphasis on conservation this aspect
of the Society’s activities has developed into Plant Records.

Dr Sledge was very much a Yorkshireman but he was by no means parochial. During his
undergraduate days, much of his travel outside the ‘Broad Acres’ was by bicycle, accompanied by
R. W. Butcher. This included visits to East Anglia, the Gower, the Avon Gorge and Dorset in
search of their special plants. In those days specimens were picked and contributed to Dr Sledge’s
enormous and excellent herbarium. The British sheets were donated several years ago to Bradford
Museums Service and they now reside in Cartwright Hall Museum (CMM) along with those of Lees.
The foreign material is widely distributed internationally.

Dr Sledge was one of those lucky individuals able to combine his interest with his career. His first
publication was a contribution to the report of the British Association meeting which was held in
Leeds in 1927, for which he wrote sections on the flora of Fountains Abbey, Wharfedale,
Nidderdale and Malham. This was followed a year later by his Ph.D. thesis on the rooting of woody
cuttings. He travelled widely in Europe and, for research purposes, visited New Zealand, Madeira,
Sri Lanka and Samoa. His work was concerned mainly with systematics and, following his first visit
to Sri Lanka, he untangled the difficult taxonomy of the native ferns. His paper, written in
association with the late Professor Irene Manton, entitled Observations on the cytology and
taxonomy of the Pteridophyte Flora of Ceylon, published in 1954, remains a classic. Other papers on
the taxonomy of tropical ferns followed, as well as numerous articles on aspects of the Yorkshire
flora. All his work was carried out with meticulous care and his ciear systematic presentation made
him a first-class teacher. Even during his period of ill-health, Dr Sledge continued to work. He
recently completed an excellent historical review for the forthcoming West Yorkshire Plant Atlas to
be published by the West Yorkshire Ecological Advisory Service, and he was still collaborating with
Professor R. E. Schultes, formerly of Harvard University, on a Symposium to commemorate the
centenary of the death of Richard Spruce (1817-1893) pioneer botanist in the Amazon and Andean
regions. Professor Schultes and Dr Sledge shared a deep respect for this little-known Yorkshire
botanical explorer and they were instrumental in having a commemorative plaque put on the
cottage in the Castle Howard estate where Spruce had lived.

168 OBITUARIES

We in Yorkshire have lost our most outstanding contemporary botanist, and one of the best in
Britain, and we mourn him along with Marjorie, his wife, who has supported him wonderfully since
their marriage in 1939. We offer sincere sympathy to her and to their son, Christopher, and his

family.

P. P. ABBOTT

INSTRUCTIONS TO CONTRIBUTORS

Manuscripts must be submitted in duplicate, typewritten on one side of the paper, with wide margins
and double-spaced throughout.

Tables, figure legends & appendices should be typed on separate sheets and attached at the end of
the manuscript.

Submission of manuscripts

Papers and Short Notes: Dr B. S. Rushton, Department of Biological and Biomedical Sciences,
University of Ulster, Coleraine, Co. Londonderry, N. Ireland, BT52 1SA.

Books for Review: Dr J. R. Edmondson, Botany Department., Liverpool Museum, William
Brown St, Liverpool, L3 8EN.

Plant Records: the appropriate vice-county recorder, who should then send them to C. D. Preston,
Biological Records Centre, Monks Wood Experimental Station, Abbots Ripton, Huntingdon,
REI) 2LS.

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.I., 68 Outwoods Road, Loughborough, Leicestershire, LE11 3LY.

ee Recorps
Book REVIEWS
OBITUARIES -

UK ISSN 0043. 1582

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WILMASET LTD, BIRKENHEAD. MERSEYSIDE
Printed in: Great Britain by :
EATON PRESS: LIMITED, WES’ PFIELD ROAD. 9. WALLASEY, en

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 1992-93

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. R. Mill, 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). -

Watsonia, 19, 169-179 (1993) 169

BRITIS: @ RALIS

Presidential Address, 1992

P. MACPHERSON

Sieeapaihlls TE

COLONISATION OF THE GLASGOW GARDEN FESTIVAL SITE THREE YEARS ON:
IMPLICATIONS FOR RECORDIN®

I have been asked for how long I hesitated before accepting the honour of Presidency of this Society,
bearing in mind the custom of there being a Presidential Address. My :-nly was, ‘“‘not half a second —
at the time, I had completely forgotten this responsibility”. Once .::. :evelation dawned, search for
an appropriate subject became a recurring thought.

‘I felt that the occasion merited original work and, being an amateur field botanist, that had to be
amateur field botany. By profession I have been a neuroradiologist, specialising in depth in a small
branch of medicine. My inclination was therefore to do the same for this address and during last
summer it gradually became apparent to me that an intensive study of the flora then colonising the
site of the Glasgow Garden Festival was the most appropriate review that I could undertake. While
doing so, I realised that this raised general issues of plant recording.

THE FESTIVAL SITE

To set the scene, the Festival was held in 1988 in abandoned dockland and quayside on the south
side of the River Clyde (Fig. 1). The main area had been the very active Prince’s Dock with a
Canting Basin, the latter used by the ships to swing round to enable entry to an individual dock or
return to the river. The extension of the site to the east took over the Mavisbank Quay.

When the dock complex became redundant it was taken over and the docks filled in with a view to
housing, but temporarily let as the site of the Garden Festival. The area is just within v.c. 77
(Lanarkshire), for which I am Recorder.

The brochure stated that the Festival was part of a process of regeneration which was
transforming the South Side of Glasgow. One million trees and flowering shrubs formed the
backdrop to an ever-changing floral carpet of bedding displays and themed gardens. There were six
theme sectors in the main section, arranged like the petals on a flower, interlinked by winding
pathways and formal avenues. The effect was also likened to a Persian carpet.

There were gardens set up by commercial organisations, the National Trust for Scotland, botanic
gardens and wildlife groups, and a series of ten International Friendship Gardens from as far apart
as Finland, China, New Zealand and Mexico. The Canting Basin was used extensively for water-
related activities and referred to in one of the maps as Princes Harbour.

Above all however, the Festival was about family fun with concerts, street theatre, exhibits,
viewing tower, vintage trams and trains and play areas. The Director called it a festival in a garden
rather than a garden festival (Fig. 2).

_ Immediately the Festival closed, work was begun in preparation for the further development of
this 48 ha site. Some plants were sold, and others taken to form the basis of continuing feature
gardens elsewhere. The bulldozers moved in again and although there were a few residual, almost
intact borders and remnants, by 1990 practically all trace of the Festival had been removed and
looking at the site in 1991 when the main area had largely returned to being waste ground, one could
hardly imagine that it had ever been. In addition little hollows looked as though they had been like
that for centuries with Compact, Hard, Soft and Jointed Rushes. (I was once told by a professional
horticulturist that he wished I would stop pronouncing the scientific names of plants as though they
were diseases!)

I did not record plants which were at the site of original planting or had simply spread in the
neighbourhood. An example is that related to the waterfall which is still bordered by the planted
material and the little pond at its base containing such species as Fringed Water-lily (Nymphoides

GUTERAL HISTOR)
-4 MAR

BOTANY LIBRAF

170 P. MACPHERSON

Clydeside
Expressway

Cantin Basin —

Ficure 1. The site and environs of the Glasgow Garden Festival in its pre-festival state. V.c. 77, Lanarks.; v.c. 76,
Renfrews.

peltata), Water-plantain (Alisma plantago-aquatica) and Bulrush (Schoenoplectus lacustris). From
the waterfall a stream had been constructed which ran through a series of small ponds to the Canting
Basin; again with residual plants such as Wild Angelica (Angelica sylvestris), Yellow Loosestrife
(Lysimachia vulgaris) and Reedmace (Typha latifolia).

I was surprised to see the size of fish that local lads were catching in the ponds. They were very
pleased to be photographed and asked if their photos would be in the papers. I said, ““No, but I am
writing an article about the site and might show the pictures in London’’. One asked, ““Are you a
famous author we should have heard about?”’!!

There will be no point in any of you asking me afterwards what kind of fish these were. My family
will tell you that I have a two-track mind, neuroradiology and field botany, and I do appreciate the
great honour of being President of the two relevant national societies at the same time. The British
Society of Neuroradiology, although a smaller organisation, presents their president with a medal
and I take this opportunity of suggesting that you might so honour future Presidents with a badge of
office that can be worn at the Annual and Exhibition Meetings and other Conferences.

THE PLANT RECORDING

As a result of intensive recording the total number of species came to 325, which I have divided into
six groups (Fig. 3).

PLANTS ON THE SCOTTISH FIELD CARD
With regard to those plants which are on the Scottish Field Card (194 species), these in the main are
those that might have been expected to have arrived by natural dispersal. However, some could

171

PRESIDENTIAL ADDRESS, 1992

‘TEANSIJ Uspsiey MOBSETD dy} Jo INOAL] SNeWIAYIS *7 AUNDIY

ae 2.

| \ Ae N
(5

* Sa ee

172 P. MACPHERSON

1988 Garden Festival Site: plants recorded in 1991

Nursery Site Plants 3% _ Stowaways 2%

Doubtful Status 5%

Natural Spread
Piants not on
Field Card 11%

Field Card Plants 60%

Spread from
Planted Material 19%

FiGurE 3. The results of plant recording in 1991 on the Glasgow Garden Festival site. 325 taxa were recorded
in all.

perhaps more appropriately have been placed in one of the other categories. As examples of spread
from planted or presumed planted material, I cite Hemp-agrimony (Eupatorium cannabinum) and
Marjoram (Origanum vulgare), which, although common in other parts of the British Isles, are
relatively unusual in the area. Perring & Walters (1962) give only one definite v.c. 77 record for
each. When I tell you that on a bank were about ten plants of Greater Spearwort (Ranunculus
lingua), you may well raise your eyebrows. Greater Spearwort had been planted in and by the pond
at the base of the waterfall, i.e. diagonally about 100 m away, with a tree border between. How did it
get to the bank? Water Dock (Rumex hydrolapathum) growing happily on the pebbly shore of the
Canting Basin was almost certainly a feature of the planting along the stream, the seeds floating
down. There is only one other site for this plant in the Glasgow area.

Although Tormentil (Potentilla erecta) and Hare’s-tail Cottongrass (Eriophorum vaginatum) are
common in the neighbourhood, they obviously arrived at the site with peat.

PLANTS WHICH ARRIVED BY NATURAL SPREAD BUT ARE NOT ON THE FIELD CARD
By natural spread, at this stage, I simply mean plants not on the Scottish Field Card, which have
colonised from the vicinity of the Festival Site or whose introduction has nothing directly to do with
the Garden Festival. : |

Of the 36 plants in this category some have been wind-blown, like Butterfly-bush (Buddleja
davidii) and five species of Hawkweed (Hieracium). I was told by a referee not to bother coming
down to the riverside at Kew, but to look for Rumex obtusifolius var. transiens in Glasgow — and at
the site I made my first record.

Other plants have been bird-sown, such as five species of Cotoneaster.

Dense Silky-bent (Apera interrupta) was actually present at another part of the site while the area
was still dockland and may have persisted, while Fodder Vetch (Vicia villosa) has been present for

PRESIDENTIAL ADDRESS, 1992 173

six years across the river and one square down. Large Trefoil (Trifolium aureum) keeps appearing
and disappearing at various sites in Glasgow.

The rarest species were Canadian Fleabane (Conyza canadensis) and Least Pepperwort
(Lepidium virginicum) growing on otherwise bare, gravelly ground and both new v.c. records.

_ DEFINITE OR PRESUMED SPREAD FROM PLANTED MATERIAL

The complete list of 62 species is given in Table 1.

In the main the spread had been wind-blown on to bare ground produced by bulldozing or with
soil shifted by the bulldozers.

There was only a single plant of some taxa such as Achillea millefolium ‘Cerise Queen’, Anchusa
azurea and Artemisia stelleriana growing on bare ground and Saxifraga paniculata on a stony path.

Between two and five plants were noted of most species such as Artemisia ludoviciana and
Armeria maritima growing on bare ground. What was Thrift doing in the centre of Glasgow unless
from planted material? So although Armeria maritima is on the Field Card I have not crossed it off,
recording it only as a cultivar. Milk Thistles (Si/ybum marianum) were on a soil heap; Eryngium
planum was on a bank; one plant of Hemerocallis was on a soil heap and another was at the sheltered
edge of a bare open patch; Japanese Wineberry (Rubus phoenicolasius) was at the edge and

_ extending down on to the shore of the Canting Basin.

Sahara eR ee Se we
RES

There were over 50 plants of Pearly Everlasting (Anaphalis margaritacea) scattered widely and
about 50 also of Yellow Chamomile (Anthemis tinctoria) but with a more local distribution. By far

TABLE 1. DEFINITE OR PRESUMED SPREAD FROM PLANTED MATERIAL
ON THE GLASGOW GARDEN FESTIVAL SITE
Taxa (62 in all) not on the field card.

Achillea millefolium ‘Cerise Queen’
Achillea ptarmica

Hyssopus officinalis
Lamiastrum galeobdolon subsp.

Alchemilla mollis argentatum
Anaphalis margaritacea Lamuastrum galeobdolon subsp.
Anaphalis margaritacea var. revoluta galeobdolon

Anchusa azurea Leucanthemum X superbum

Anthemus tinctoria
Armeria maritima cv.
Artemisia absinthium
Artemisia ludoviciana
Artemisia stelleriana
Astilbe cf. japonica
Berberis sp.

Borago officinalis
Campanula medium
Campanula persicifolia
Carex pendula

Cornus sericea
Cotoneaster salicifolius
Cotoneaster X suecicus
Dianthus barbatus cv.
Elaeagnus angustifolia
Eryngium planum
Festuca brevipila

Lychnis coronaria

Lythrum salicaria cv.

Melissa officinalis

Mentha x villosa

Narcissus pseudonarcissus cv.
Narcissus tazetta

Nepeta X faassenii

Phalaris arundinacea var. picta
Polygonum bistorta ‘Superbum’
Potentilla fruticosa hybrid
Pulmonaria officinalis

Ribes sanguineum

Robinia pseudoacacia

Rubus phoenicolasius

Saxifraga paniculata

Senecio cineraria

Silybum marianum

Spiraea douglasii x S. salicifolia

Geranium endressii X G. versicolor
Geranium macrorrhizum
Geranium sanguineum

Geranium X magnificum

Spiraea sp.

Symphoricarpos cf. X chenaultii
Symphoricarpos albus

Tellima grandiflora

Geum cv. Tolmiea menziesii
Hemerocallis sp. Verbena rigida
Hippophae rhamnoides Vinca major

Hosta sp. Viola lutea cv.

174 P. MACPHERSON

TABLE 2. TAXA OF DOUBTFUL STATUS ON THE GLASGOW GARDEN FESTIVAL SITE

Angelica archangelica Oenothera biennis X O. glazioviana
Aster sp. Phalaris canariensis

Cotoneaster conspicuus Rosa rugosa

Ligustrum ovalifolium Rubus cockburnianus

Linum usitatissimum Salix alba x S. fragilis

Lysimachia punctata Salix discolor

Mimulus cupreus X M. luteus xX M. variegatus Solidago gigantea

Myosoton aquaticum

the most common plant in this category was Purple Loosestrife (Lythrum salicaria cv.) which had
been a feature of water-side planting but had spread to most parts of the site regardless of soil or
moisture.

DOUBTFUL STATUS
Some of those on this list (Table 2) might have been from planted material but their introduction
could equally well have been from other categories. 15 species were of doubtful status.

Japanese Rose (Rosa rugosa), White-stemmed Bramble (Rubus cockburnianus) and Early
Goldenrod (Solidago gigantea) have been growing for many years in the neighbourhood. With
regard to two plants of Garden Angelica (Angelica archangelica) growing well in from the river one
can speculate fancifully. The plants grow on top of all the wooden piles near the mouth of the nearby
River Kelvin (Macpherson 1984). The docks at the Festival Site were partly filled in with dredgings
from the river. However as the seeds are more likely to float than sink to the bottom, then perhaps
two were picked up as a bucket broke the surface!

Water Chickweed (Myosoton aquaticum) surprised me; one plant in a stony hollow at the edge
above the Canting Basin, but then it can be an import and flourish on dry ground, as in London
(Burton 1983). The Mimulus cupreus x M. luteus x M. variegatus was less than 2 cm high — the
name being much longer than the plant! Flax (Linum usitatissimum) and Canary-grass (Phalaris
canariensis) possibly came from bird seed.

‘STOWAWAYS.

Plants whose introduction I presume to have been in soil round plants brought for display. Ihave |

excluded plants which are on the Field Card, such as those brought in with peat as mentioned above,
or with woodland planting such as Wood Speedwell (Veronica montana) which leaves seven species
(Table 3).

Although there was much planting in and along the waterfall and stream I cannot imagine that

Orange Foxtail (Alopecurus aequalis) was brought intentionally, but there it was, one plant on the

muddy edge — from whence did it come?

Free access was available to only one tiny part of the site in 1990 and on part of a bare trodden
pathway there were six plants of Smooth Rupturewort (Herniaria glabra). 40 m farther on (in the
next 1 km square) there were a further nine. In 1991 there were more than 50 plants. This species is
even rarer than the Alopecurus in the British Isles.

I assume that Spiny Restharrow (Ononis spinosa) came with seed for sowing the grassy bank near
the waterfall. Like the two mentioned above and Sulphur Cinquefoil (Potentilla recta), also in this
group, this plant has not been recorded before in the vice-county.

TABLE 3. ‘SSTOWAWAYS’ ON THE GLASGOW GARDEN FESTIVAL SITE

Alopecurus aequalis

Geranium pyrenaicum

Herniaria glabra

Melilotus indicus

Ononis spinosa

Potentilla recta

Rumex acetosella var. tenuifolius

PRESIDENTIAL ADDRESS, 1992 175

NATURAL ENVIRONMENT RESEARCH COUNCIL
BOTANICAL SOCIETY OF THE BRITISH ISLES

Ka .

This is to certify that

cesé aot dy
| sta mas Dr Peter Macpherson

1s an

OFFICIAL PLANT RECORDER

Ficure 4. A personal ‘Access’ card designed by the author.

NURSERY SITE

In addition to the actual Garden Festival Site itself there is an adjacent area, connected by a short
tunnel under the intervening road, in which plants were stored, either in containers or shoved into
the ground while their sites were being prepared, or in some cases as substitutes to provide flowering
- colour throughout the season. It is an area of 3.5 ha.

It is still fenced off, but like most such pieces of ground, at least in Glasgow, if one searches one is
_ bound to find a gap where someone has cut a way in!

With regard to the main site, the Security Officer became interested in my survey and kept asking
if I had been to such and such a part where he had seen flowers — as opposed to weeds! At the
» Norfolk Recorders’ Conference I suggested that we be issued with a card and that seemed to meet
» with general approval, but as none was forthcoming I designed my own (Fig. 4). Of course it has no
official standing but never fails to impress.

I have included the Nursery Site (Table 4) because it was very much part of the festival complex as
far as plants were concerned and had eleven additions to what were seen elsewhere. Agrostis scabra
we see in hundreds, if not thousands each year in Glasgow. Crepis setosa reached Glasgow in 1984

but as a casual only (Macpherson 1990). In both 1990 and 1991 there were at least 40 plants on the
- bank of the track as it dipped down to go under the road, with two plants having crept into the
Garden Festival site. Genista hispanica had definitely spread from planted material as presumably
had the Hosta. :

| [ had nipped in during 1990 and the most interesting plant was a tufted, reddish sedge identified
for me as Carex buchananii, a native of New Zealand and possibly new to Britain as a wild plant. In
1990 one flowering plant was present. In 1991, the colony had increased to four (Macpherson &
Macpherson 1992) and a recent check showed that there had now been a further increase to six. This
‘plant plus one small Sensitive Fern (Onoclea sensibilis), more or less hidden away among a group of
_Cornus which had been left over near the boundary wall, would qualify for my ‘Stowaway’ category.

TABLE 4. TAXA IN THE NURSERY AREA ON THE GLASGOW
GARDEN FESTIVAL SITE

| Agrostis scabra Hosta cf. nakaiana
Carex buchananii Medicago sativa
Cichorium intybus Mentha X villosonervata
| Cotoneaster sternianus Onoclea sensibilis

Crepis setosa Solidago canadensis

Genista hispanica subsp. occididentalis

176 P. MACPHERSON
DISCUSSION

So now that I have all these records — what are the implications? What do I do with them?

With regard to those which have arrived by what could be natural dispersal I have no problem. I —

stroke off the Field Card and put.a dot on my index card. However I cannot simply accept all that are
printed on the Field Card without comment as some are very rare or absent from my vice-county,
and had probably spread from deliberate planting, e.g. Marjoram and Hempagrimony.

The next ‘acceptable group’ is comprised of those not on the Field Card, but which I assume have
not been deliberately brought to the site. Some are actually reasonably common in Glasgow and
could have spread on to any suitable ground. These I have no hesitation in recording without
comment e.g. Brassica rapa, Heracleum mantegazzianum, Senecio squalidus. With regard to rarities
such as Canadian Fleabane and Least Pepperwort, which I assume ‘fell off the back of a ship’, I have
noted the facts, done a drawing of their location and will monitor to see how long they persist as was

the case with Agrostis scabra which we first saw as a single plant in 1973 and for which I now have

eleven personal 1 km* records (Macpherson & Stirling 1988).

The other group not deliberately introduced are my ‘stowaways’. I already know that Herniaria
glabra and Carex buchananii are spreading and could easily become established if given a chance.

Of the vast majority of planted material, there is now no trace. With regard to the recording of
those which have spread or been spread from Garden Festival planting there is more room for
argument. I have filled out a card for those such as Anchusa azurea, and even added a dot. These
should at least therefore always be available for reference.

What of these so-called aliens? How are they to be regarded?

For reference I selected books from my bookshelves more or less at random. In no sense is this an
exhaustive review:

‘‘Human borne seedlings do not seem to me to deserve much consideration. I am not convinced
that we ought not to advocate definite action to prevent their settlement” (Raven 1953). A critical
comment taken a little out of context, but which one might argue could apply at least to the Giant
Hogweed (Heracleum mantegazzianum).

Or with disdain: ‘‘I almost immediately excluded aliens because they are coming and going in an
endless stream, occasionally leaving a mark on our landscape as in the case of Senecio squalidus, but
more often waifs and strays, left to perish on a dunghill” (Meikle 1953).

Or an intermediate view: “The alien flora is of little importance but must be accounted for as
many of the familiar plants of the countryside had their introduction either as aliens or as garden ~
escapes” (Dony 1976).

Or more tolerantly: ‘The British flora is changing whatever we do but all changes are not always
for the worst, some constituting interesting additions to the British flora and not necessarily
undesirable” (Brenan 1983).

A few years ago I met a fellow Scottish Recorder and reported that I had v.c. records of Chinese _

Ragwort (Sinacalia tangutica) and Peach-leaved Bellflower (Campanula persicifolia), both well
established, and which I would pass on. I was told not to bother as they would go straight into the
waste-paper basket. Campanula persicifolia is also present at the Garden Festival Site, so do I
consign the record, specimen and slide to the bin?

For some time I have had a certain reputation for recording ‘non-natives’ which can brighten up
dull routine recording (Dickson 1991), and so appreciate the contents of the New Flora of the British
Isles which states that ‘‘The aim is to include all taxa that the plant hunter might reasonably be able
to find in the wild in any one year”’ (Stace 1991).

I do appreciate that there is a pecking order:

Native — whatever that is! Swann (1977) stated that no two botanists will ever agree about what
constitutes a native species. It has been variously defined as: a species believed to have been in the
country before man (Lousley 1953; Ellis 1983); part of the natural vegetation for a long time (Briggs
1990); in a natural locality to which it has spread by natural means from a natural source (Dunn
1905); immigrated without the aid of man (Lousley 1953; Ellis 1983; Dickson 1991; Stace 1991):
arisen de novo in the country (Lousley 1953; Ellis 1983).

Non-native — unintentional introductions may be regarded as more imereeae than those brought

in intentionally. Some authors regard alien and introduction as synonymous while others limit

introduction to deliberate acts.

|
|
|

PRESIDENTIAL ADDRESS, 1992 177

Alien plants have been defined as species introduced by the intentional or unintentional agency of
man (Lousley 1953; Ellis 1983; Crackles 1990); species introduced by man and now more or less
established (Clapham et al. 1962).

Introduced alien — deliberately brought to the area for whatever reason (Dickson 1987).

Invading alien — unintentionally brought to the area or arriving in the area by natural means
(Dickson 1987).

There is then the question of Naturalised vis-a-vis Established. Some authors make a distinction
and regard Naturalised as a more important category: a species naturalised in natural habitats
(Hyde & Wade 1934); species naturalised in a natural or semi-natural habitat (Lousley 1953).

- Others regard them as synonymous: a species which has become self-perpetuating (Stace 1991); an

alien plant which has become completely established. Those who differentiate define Established

_ Alien as persisting only in a man-made habitat (Hyde & Wade 1934; Lousley 1953); or in absence of

competition from native species (Crackles 1990). The criteria have been widened to include an
annual which persists or is repeatedly introduced (Ellis 1983).
Both Naturalised and Established are regarded as more important than Casual (Temporary;

_ Ephemeral; Visitors). There is more unanimity regarding the definition of Casual, the general view

being that it refers to a species which does not persist (Lousley 1953; Clapham et al. 1962; Ingram &

_ Noltie 1981; Ellis 1983; Crackles 1990; Graham 1988). The terms Ephemeral Alien (Jermyn 1974)
and Adventive (Fitter & Fitter 1967) have been used as synonyms for casual.

How long does a plant have to persist before it can be regarded as Naturalised/Established? The
plants that I have recorded have persisted for two to three years. Lousley (1953) had suggested 25
years. I am unlikely to persist that long, so when may I put them into Plant Records?

What is an Escape and are all escapes equal? An escape has been defined as: of cultivated origin
but not naturalised (Clapham er al. 1962; Ingram & Noltie 1981); originally from seed or plants

| thrown out with garden rubbish (Jermyn 1974); spread vegetatively or by seed from a garden (Stace
21991).

I present a classification of introductions in what could be regarded as a pecking order (Fig. 5).
Other terms used in the literature include:—

Indigenous — native born; originating or produced naturally in a country; opposite of exotic
- (Kirkpatrick 1983); Exotic — introduced from a foreign country; alien; not native to a country as a

plant, etc; romantically — strange, rich or showy (Kirkpatrick 1983); Denizen — doubtful native in a

INTRODUCTIONS
NATURAL UNINTENTIONAL INTENTIONAL
+ Permanent Casual + Permanent Casual

Naturalised Established Naturalised Established

Escape from Dumped Sown;

Cultivation | planted

+ Permanent Casual + Permanent Casual + Permanent Casual

Naturalised Established Naturalised Established Naturalised Established

Ficure 5. Different modes of introduction and permanency in (arguably) decreasing importance from left to right.

178 P. MACPHERSON

natural habitat (Watson 1870; Lousley 1953); an alien species growing in a natural or semi-natural
community (Lousley 1953); species growing wild but originally introduced as cultivated crop or herb
(Graham 1988; Crackles 1990); native elsewhere in Britain but not in the area (Hall 1980);
Adventive — a plant growing unaided but not yet permanently established (Hyde & Wade 1934); not |
deliberately introduced (Crackles 1990); self-sown or bird-sown garden plant (Briggs 1990) or even
Colonist, a category implied in the title of this address — doubtful native in a man-made habitat
(Lousley 1953); native elsewhere in Britain but not in the area (Whitehead 1976); species which
grows only on man-maintained habitat (Lousley 1953; Graham 1988); alien which is established
(Jermyn 1974); synonym of Adventive which is a synonym of Casual (Fitter & Fitter 1967).

So is Pineappleweed (Matricaria discoidea) more acceptable, having been brought in unintentio-
nally, than Oxford Ragwort (Senecio squalidus) which was actively brought to this country; and that
in turn more acceptable because it escaped over the garden wall than Dotted Loosestrife
(Lysimachia punctata) which was thrown out and is now all over the West of Scotland? It has been
recorded in 70 of the 90 Flora of Glasgow tetrads. All three plants are on the Garden Festival Site. If —
I stop to think about it, yes, I agree with the pecking order, but while recording never give it a
thought.

I wish to make a comparison. After the Ice-age Britain was:—

a. wholly or mainly a bare area available for colonisation by plants (Heslop-Harrison 1953); and

b. a land available for colonisation by people.

As there were no people initially in the British Isles all here now are descendants of those who
were native elsewhere. The British Isles have been populated by waves of tribes: Iberians, Celts,
Picts, Angles, Saxons, Normans and so on. In the eyes of the Iberians the Celts were invasive
intruders, in the eyes of the Celts the Anglo-Saxons were incomers and so it went on. There is even
the question of being alien to a district. In the cities people are more readily absorbed into the
community but in the Highlands it 1s often said that families need to remain for a couple of
generations before they cease to be Incomers! There is also antipathy in parts of Wales to ‘white
settlers’. In contrast many people in England more or less regard Great Britain as synonymous with
England and therefore include the Scots and Welsh as English! In spite of the above no-one would
quibble if most if not all of us claim to be natives of the British Isles. I feel that this may have
something to tell us about our attitude to plant status.

With regard to plant distribution within the British Isles some Floras define Alien as not native to
the area under discussion, although it may be native in other parts of the British Isles. What are the
geographical limits to which this would apply? If a plant is gradually extending into Scotland from
England by natural dispersal, are the offspring which cross the border into Scotland to be regarded
as aliens? Or, to narrow it down, if a plant arises from seed blown on to suitable ground in Glasgow
from a neighbouring so-called native habitat are we allowed to call it Native in Glasgow, in the Flora
of Glasgow. Certainly ‘Alien’ would not seem to be appropriate. In other words are the boundaries
not rather arbitrary? Even if the term Alien is acceptable for the newly arrived, in general, afterone _
or two generations of people, or rather more in the case of plants, the term is not appropriate.

There is therefore an obvious diversity of opinion regarding the definition of terms or even of
their acceptability. ““The various terms are not easily, precisely defined as different botanists use
them in different ways” (Briggs 1990). “‘In assessing the status of many species, no two botanists
would ever agree, judgement is so much a matter of personal opinion and many of the terms used
are themselves arbitrary” (Petch & Swann 1968).

As a further example of differing terminology I present these definitions of Introduced Plant —
brought accidentally or intentionally (Clapham et al. 1962); deliberately sown or planted (Fitter &
Fitter 1967); deliberately brought for whatever reason (Dickson 1987); deliberately sown or planted
or garden escape (Graham 1988); brought by man but apparently naturalised (Ingram & Noltie
1981).

Some Presidential Addresses end with such statements as: Further work along the lines of my talk |
would be a suitable project for the Society (and that is usually the last one hears of it—like my access
card and possible Presidential Medals!).

I am going to be more positive and ask if I may set up a little sub-committee with a view to
producing agreed unambiguous British Terminology. It should be less controversial than the
English Names of Wild Flowers which, although many in Scotland object to, I have, with two
exceptions (did you notice one in my address?) been prepared to accept for the overall good.

PRESIDENTIAL ADDRESS, 1992 179

The requirement for clear knowledge of status without ambiguity is clearly indicated by the story

- of the young man from the West of Scotland who went into a London establishment that he would

- not have patronised had he known its reputation. He chatted to a young lady for some time and

eventually she leant forward, touched his arm and said ‘‘You do realise that I am a Call Girl?” He
leant back — *“‘Now is that not a co-incidence — I’m from Tiree”’.

ACKNOWLEDGMENTS

I am grateful to the following who identified plants in relation to this paper: J. C. Bowra, E. J.
Clement, M. G. Daker, J. Fryer, R. J. Gornall, R. M. Harley, I. C. Hedge, A. C. Jermy, D. H.
Kent, A. C. Leslie, D. J. McCosh, D. R. McKean, R. D. Meikle, D. M. Moore, A. Newton, H. J.
Noltie, J. R. Palmer, T. C. G. Rich, A. J. Silverside, D. A. Simpson, C. A. Stace, A. McG. Stirling,
P. J. O. Trist, B. S. Wurzell and to A. & R. Bowden, Hostas Nursery, Okehampton, Devon.
Illustrations were provided by N. MacPhee and E. L. S. Lindsay. Permission has been granted by
Scottish Enterprise to reproduce Fig. 2 of this article from the Glasgow Garden Festival souvenir
brochure.

REFERENCES

BRENAN, J. P. M. (1983). The British flora — a changing picture. Watsonia 14: 237-242.

Briccs, M. (1990). Sussex plant atlas selected supplement. Brighton.

Burton, R. M. (1983). Flora of the London area. London.

CLAPHAM, A. R., Tutin, T. G. & WARBURG, E. F. (1962). Flora of the British Isles. Cambridge.

~ Cracktes, F. E. (1990). Flora of the East Riding of Yorkshire. Hull.
_ Dickson, J. H. (1987). Native v. alien in the flora of Glasgow (or elsewhere). B.S.B.I. News 47: 17-18.

Dickson, J. H. (1991). Wild plants of Glasgow. Aberdeen.
Dony, J. G. (1976). Bedfordshire plant atlas. Luton.
Dunn, S. T. (1905). Alien flora-of Britain. London.

Exuis, R. G. (1983). Flowering plants of Wales. Cardiff.

_ Firrer, R. & Fitter, M. (1967). Penguin dictionary of British natural history. Harmondsworth.
_ GrauaM, G. G. (1988). The flora and vegetation of County Durham. Durham.

Ha t, P. C. (1980). Sussex plant atlas. Brighton.

_ Hesrop-Harrison, J. (1953). The North American and Lusitanian elements in the flora of the British Isles, in

LousLey, J. E., ed. The changing flora of Britain, pp. 105-123. London.

Hype, H. A. & Wane, A. E. (1934). Welsh flowering plants. Cardiff.

INGRAM, R. & Nortir, H. J. (1981). The flora of Angus. Dundee.

JERMYN, S. T. (1974). Flora of Essex. Dunmow.

Kirkpatrick, E. M. ed. (1983). Chambers 20th century dictionary. Edinburgh.

Lous_Ley, J. E. (1953). The recent influx of aliens into the British flora, in Lous.ey, J. E., ed. The changing flora
of Britain, pp. 140-159. London.

_ Macruerson, P. (1984). Garden Angelica by the River Kelvin. Glasgow Naturalist 20: 469-471.
_ Macpuerson, P. (1990). Bristly Hawk’s-beard eventually identified in the Glasgow area. Glasgow Naturalist 21:

612.
Macpuerson, P. (1992). Smooth Rupturewort in the Glasgow Garden Festival site. Glasgow Naturalist 22: 191-
192:
Macpuerson, P. & MacpHerson, A. C. (1992). Carex buchananii in Glasgow. Glasgow Naturalist 22: 194.
MacpH_erson, P. & StTirLinc, A. McG. (1988). Agrostis scabra in Glasgow. B.S.B.I. Scottish Newsletter 10: 8.

_ MEIKLE, R. D. (1953). Recent additions to the British flora, in LousLey, J. E., ed. The changing flora of Britain,

pp. 49-51. London.
PERRING, F. H. & WALTERS, S. M., eds. (1962). Adlas of the British flora. London.

_ Petcu, C. P. & Swann, E. L. (1968). Flora of Norfolk. Norwich.

Raven, C. E. (1953). The significance of a changing flora, in LousLey, J. E., ed. The changing flora of Britain,
pp. 14-18. London.

Stace, C. A. (1991). New flora of the British Isles. Cambridge.

SwANn, E. L. (1977). Norfolk and botany. Watsonia 11: 189-194.

Watson, H. C. (1870). A compendium of the Cybele Britannica, pp. 61-63, London.

WHITEHEAD, L. E. (1976). Plants of Herefordshire. Hereford Botanical Society.

Watsonia, 19, 181-184 (1993) 181

The correct Latin names for the Primrose and the Oxlip,
Primula vulgaris Hudson and P. elatior (L.) Hill

R. K. BRUMMITT
The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE
and
R. D. MEIKLE

Ranscombe Lodge, Wootton Courtenay, Minehead, Somerset, TA24 SRA

ABSTRACT

The correct Latin names for the Primrose and the Oxlip are maintained as Primula vulgaris Hudson and P.
elatior (L.) Hill (Primulaceae), despite recent assertions that the Primrose should be called P. acaulis (L.) L. and
that a question hangs over P. elatior. Additional evidence of Linnaeus’s intentions in his Flora Anglica is
provided in different printings of this work which have been previously overlooked.

INTRODUCTION

It may seem surprising and unfortunate that there should still be disputes over the correct Latin
names of the Primrose and the Oxlip after three centuries or more of the study of the European flora
and well over two centuries of the adoption of Linnaean binomial nomenclature. However, Greuter
(1989a) resurrected the name Primula acaulis (L.) L. for the Primrose. This has been adopted in the
Med-Checklist (Greuter, Burdet & Long 1989), and new subspecific combinations have been
published by Greuter & Burdet (Greuter 1989b). It has also been maintained that unless the
arguments for this are accepted it is impossible to maintain the name P. elatior for the Oxlip. The
decisions depend largely on interpretation of Linnaeus’s Flora Anglica (1754), a dissertation
defended by his student Grufberg. During preparation of the account of Primulaceae for the Flora
of Cyprus (Meikle 1985), we looked into this question and were satisfied that there was no threat to
the well-established P. vulgaris Hudson, and we maintain this position now. Re-examination of the
facts has brought to light an overlooked significant variation in the typography of the 1754 Flora
Anglica, which reinforces our opinions.

THE LATIN NAME OF THE PRIMROSE

In Species Plantarum (1753), Linnaeus did not provide a specific binomial for the Primrose because
he adopted a broad species concept, recognising the Cowslip, Oxlip and Primrose as one species, P.
veris, with three varieties, var. officinalis, var. elatior and var. acaulis respectively. (This taxonomic
concept persisted in British botany through all editions of Bentham’s Handbook of the British Flora,
to Rendle’s 7th edition in 1924.) In 1762 William Hudson published his Flora Anglica, including the
Cowslip and Oxslip in P. veris but raising the Primrose to specific rank with the new name P.
vulgaris. In 1765 John Hill also gave the Primrose specific rank, but took up Linnaeus’s varietal
epithet acaulis as P. acaulis, a name which has occasionally been adopted since. However, at specific
rank P. vulgaris Hudson has three years’ priority over P. acaulis (L.) Hill and so has been widely
adopted.

The current argument is over the rank given to the Primrose in Linnaeus’s Flora Anglica (1754).
Greuter (1989a) has concluded that here Linnaeus raised the Primrose to specific rank under the

182 R. K. BRUMMITT AND R. D. MEIKLE

Echium vulgare 227-1. Solanum nigrum 205 - 4.
Lycop/is 227-2. Duicamara 2665-1-2.

Primula verisofficinal.zgge3. Ramnus catharticus 466-1.
elarior 2. Frangula 465-1.

acaulis 1. Evonymus ewopzus 468-1.

farinofa 285-1. Ribes rubrum 456-1,
Menyanthes trifoliata 285-1. alpinum 456-2.
Nymphoitdces 368-2. nigrum ay6 -4.
Hoftonia paluftris 285-1. Hedera helix 459-1.
Lysmachia vulgaris 282-1, Illecebrumverssceslatum 160-1.
thyr(iflora 183-3. Glaux maritima 285-1.
Nummularia 283-1. Thefium Linophyllum 202-1.

nemorum 282-5. Vinca minor 268-1.
Anagallis arvenfis 232 = 4, major 268-2.
Di-

Figure 1. Linnaeus’s Flora Anglica (1754); lower part of p. 12, from Ray Society 1973 facsimile.

name P. acaulis, so that this name has priority over P. vulgaris Hudson (1762). Stearn (1973, p. 68),
however, in collaboration with the late J. E. Dandy, had concluded that Linnaeus merely repeated
his taxonomy of Species Plantarum in his Flora Anglica, and that apparent inconsistencies were due
to obvious slips of the pen or typographical errors. Greuter reproduced in facsimile the relevant
page of Linnaeus’s Flora Anglica (see Fig. 1), and argued that names given there which are neither
trinomial nor subordinated by indentation must be accepted as specific binomials. In the case of
Primula, ‘veris officinal[is]’ appears on one line, with ‘elatior’, ‘acaulis’ and ‘farinosa’ on successive
lines indented only very slightly under ‘veris’. The indentation, or lack of it, may here tend to
suggest that all four taxa were given specific rank. As Greuter has noted, the italicisation of acaulis
merely denotes that this taxon does not occur in Sweden.

However, when we looked again at the Kew copy of Flora Anglica (see Fig. 2) which we had
consulted ten years ago, we found that the typesetting is different from that of the Ray Society's
facsimile edition (1973), and that ‘elatior’ and ‘acaulis’ are indeed indented under ‘veris’ almost to
the position of the varietal epithet ‘officinallis]’, while ‘farinosa’ is indented almost as far. There are
similar inconsistencies between copies in indentation on the same page (see Echium, Ribes, Hedera,
R[h]amnus, Vinca) and on other pages, and it is clear that there were at least two different printings ~
of this dissertation and that the typesetting indentation was haphazard. We are grateful to Dr J. L.
Reveal for informing us that the Natural History Museum, London possesses both printings. It
appears that in the version represented at Kew the printer was instructed to remove excessive spaces
between some generic names and the following epithets without moving succeeding epithets, so that
indentations become quite different. Indentation thus appears meaningless as far as evidence of
rank is concerned. It would be false to argue that in the Kew copy the position of the epithets
alpinum and nigrum in relation to rubrum under Ribes (see Fig. 2) indicates that they represent
infraspecific taxa. But certainly, if the Kew copy alone were considered, the weight of evidence from
indentation alone would indicate that Linnaeus still regarded elatior and acaulis as varieties of P.
veris and not as separate species, contrary to Greuter’s conclusion.

But the clinching evidence of what Linnaeus did or did not do in Flora Anglica is provided by a
point already stressed by Stearn (1973, p. 68). For the Cowslip, Linnaeus employed the trinomial
Primula veris officinal/is], indicating that he still recognised varieties in the species. If he had raised
all three taxa to specific rank he would certainly have dropped the varietal epithet officinalis. To our
knowledge, in all his botanical works he never employed a varietal epithet in species in which he did
not recognise varieties. The aim of Flora Anglica was essentially to list the known flora of England
according to the binomial system, and although he did introduce a few novelties (Stearn 1973, pp.
63-68), in the case of Primula it is clear that he merely retained his taxonomy from Species

THE LATIN NAMES FOR THE PRIMROSE AND THE OXLIP 183

Echium vulgare - 227-1. Solanum nigrum 265-4.
Lycopfis 227-2. Dulcamara 265-1-2:.

Primula veris officinal. 284+ 3. Ramnuscatharticus 466~1..
elatior 2. Frangula 465 -14,.

acaulis I. Evonymus europeus 468-r..

farinofa 285-1. Ribes rubrum 456-T..
Menyanthes trifoliata 285-1. alpinum 456-2:
Nympboides 368-2. nigrum 456-4.

Hottonia paluftris 285-1. Hedera helix 459-1,
Lysmachia vulgaris 282-1. Illecebrum verticilatum 160-1.
thyrfiflora) 283-3. Glaux maritima. 285 °T.
Nummoularia 283-1. Thefium Linophyllum zo02-r.

nemorum 282-'s. Vinca minor 268-1.

_Anagallis arvenfis 282-1. major 268-2.
DI-

| Figure 2. Linnaeus’s Flora Anglica (1754); lower part of p. 12, from Kew copy.

Plantarum published the previous year. The erratic behaviour of a typesetter in 1754 does not alter
the facts and cannot be used as a reason to upset established nomenclature of a well-known plant.
Nor is the issue affected by the slightly more orderly, but still inconsistent, typesetting of the reprint
» of Flora Anglica in Linnaeus’s Amoenitates Academicae (1759, vol. 4), where all four epithets veris,
_ elatior, acaulis and farinosa were equally indented under Primula (p. 97). The inclusion of the
| varietal epithet officinalis still indicates that Linnaeus had not changed his taxonomy. Indeed,
I throughout all his works Linnaeus kept the Primrose and the Oxlip as varieties of Primula veris. The
_ correct name (and relevant synonyms) at specific rank for the Primrose is therefore as follows:

keveris:var-acaulis L:, Sp. Pl. 1: 143: (1753).

| :

| P. vulgaris Hudson, Fl. Angl., 70 (1762).

| P. acaulis (L.) Hill, Veg. Syst. 8: 25 (1765).

THE LATIN NAME OF THE OXLIP

|
|
|

|
1
}

Having concluded that Linnaeus raised the Primrose and Oxlip from varietal rank in Species
Plantarum (1753) to specific rank in Flora Anglica (1754), Greuter (1989a) has cited the correct
+ name for the Oxlip as Primula elatior (L.) L., Fl. Angl. 14 (1754) instead of the more usual P. elatior
| (L.) Hill (1765). He has warned that, if this is not accepted, the specific name for the Oxlip will have
_to change. The reason for this conclusion is that he considers that when Hill in 1765 published the
) name Primula elatior he did not make a new combination based on Linnaeus’s P. veris var. elatior
) but described a new species, which, according to Schinz & Thellung (1907, p. 333), is not the Oxlip
but the hybrid between the Cowslip and Primrose, the False Oxlip. This would then mean that the
i name P. X elatior Hill would have to be applied to the hybrid, that any later combination of
| Linnaeus’s elatior at specific rank would be an illegitimate later homonym, and that a new name
/} would have to be found for the Oxlip. Fortunately we cannot agree with this sequence of
| conclusions.

| ‘The fact that Hill may have described and illustrated the hybrid P. veris < vulgaris under the
j}name P. elatior is irrelevant if the latter is a combination based on Linnaeus’s var. elatior
| Unternational code. . . Berlin 1987, Art. 7.12). The question therefore is whether Hill made a new
_; combination or described a new species. Throughout the 26 volumes of the quarto edition of The
_ Vegetable System (1759-1775), in which the name appeared, Hill never cited any authors for the
_) binomials he used, nor did he cite any basionyms. He was, however, clearly using the Linnaean

“fl
#
5

a
~~

184 R. K. BRUMMITT AND R. D. MEIKLE

system, to which he referred directly in his introduction (Hill 1759, vol. 1, p. 24) with a comment
that “‘it will live . . . so long as there is science’’. It would be ludicrous to argue that the names he
used were independent of those published previously by Linnaeus simply because he used no author
citations. Such a conclusion would require that every name in all 26 volumes of the Vegetable System
should be listed in Index Kewensis as new species attributed to Hill.

Article 33.2 of International code. . . Berlin 1987 requires that a full and direct reference be given
in valid publication of a new combination after 1 January 1953, surely with the implication that
before 1953 such a reference is not necessary. All the circumstantial evidence, and in particular the
coincidence of all the epithets officinalis, acaulis, elatior and farinosa under Primula in the relevant
publications of both Linnaeus and Hill, points to the fact that Hill was merely taking up the earlier
Linnaean epithets and was not describing new taxa. The coincidence of the epithets themselves
refers us back to Linnaeus, which is confirmed (if it were necessary) by Hill’s direct reference to
Linnaeus in his introduction, as noted above. Publication of the combination by Hill, not Linnaeus,
was accepted in the original volume of Index Kewensis (1895) and by generations of botanists since.
It is comparable with the combination Helianthemum nummularium (L.) Miller, Gard. Dict. (ed.
8), [sub]. Helianthemum no. 12 (1768), also published with a passing reference to Linnaeus himself
in the introduction to the book, and with a reference under the generic riame, but without any
reference to the basionym Cistus nummularius L., Sp. Pl. 1: 527 (1753), and nonetheless universally
accepted as a new combination at the present time. To maintain today, when the need for
nomenclatural stability is being much discussed, that such names should be treated as newly
described species, with consequent new typifications, rather than new combinations, would not only
be highly undesirable, it would be contrary to the /nternational code. We are happy to conclude that
the correct name for the Oxlip should be maintained as follows:

P. elatior (L.) Hill, Veg. Syst. 8: 25 (1765).
P. veris var. elatior L., Sp. Pl. 1: 143 (1753).

ACKNOWLEDGMENTS

We are grateful to Werner Greuter for helpful discussion of the matter. We also thank Professor W.
T. Stearn for discussion, and are happy to have his full support for our conclusions above. We are
also grateful to Dr C. E. Jarvis and Dr J. L. Reveal for helpful comments and support. Thanks are
also due to the Library, Royal Botanic Gardens, Kew for permission to reproduce Fig. 2.

REFERENCES

GreEuTER, W. (1989a). Med-Checxlist Notulae Bibliographicae, 15. Primula acaulis, Primula elatior and the
“Flora Anglica”’ of Linnaeus. Candollea 44: 562-567. ;
GREUTER, W. (1989b). [Primula], in GREUTER, W. & Raus, T., eds. Med-Checklist Notulae, 15. Willdenowia 19:

GREUTER, W., BurpDET, H. M. & Lona, G. (1989). Med-Checklist 4: 378. Genéve.

HILi, J. (1765). The Vegetable System, 8: 25. London.

MEIKLE, R. D. (1985). Primula in Flora of Cyprus 2: 1075-1076. Kew.

SCHINZ, H. & THELLUNG, A. (1907). Begriindung vorzunehmender Namensanderungen an der zweiten Auflage
der ‘Flora von Schweiz” von Schinz und Keller. Bull. Herb. Boiss., ser. 2, 7: 331-346.

STEARN, W. T. (1973). Ray, Dillenius, Linnaeus and the Synopsis methodica Stirpium Britannicarum, in Ray, J.
Synopsis methodica Stirpium Britannicarum, editio tertia, 1724, & LINNAEUS, C. Flora Anglica, 1754 & 1759
(Ray Society facsimile edition) pp. 1-90. London.

(Accepted July 1992)

Watsonia, 19, 185-193 (1993) 185
Short Notes

DIANTHUS ARMERIA L. NEW TO IRELAND AND OTHER RARE PLANTS
IN WEST CORK

This note presents a preliminary report of extensive floristic research on Sherkin Island and adjacent
islands in Roaringwater Bay to the west of Baltimore, West Cork (vice-county H3). During field-
work from March to September 1992, based at Sherkin Marine Station, K.C. (joined by J.R.A., 25
August—-4 September) made so many significant records of rare native and ruderal species that it
seemed pertinent to make the most interesting results available. All data have been lodged at
Sherkin Marine Station. Some of our herbarium voucher specimens of the plants are fragmentary or
even lacking, as many of the plants are extremely rare and protected by Irish law. Nomenclature
follows Stace (1991).

Dianthus armeria L.
Horse Island, rocky pasture near the sea, 1 September 1992, J. R. Akeroyd & K. Clarke, DBN, herb.
Sherkin Marine Station.

Six plants in all were found, severely grazed by sheep and goats. The plants had anomalous few-
flowered inflorescences with secondary growth below, but could be distinguished by the annual/
biennial habit and flowers c. 1 cm in diameter, bright reddish-pink and scentless. The leaves, bracts
and stems were reddish.

Dianthus armeria, a plant of grasslands on sandy soils, has not previously been reported from
Ireland. It occurs over much of southern and central Britain, but is now rare and decreasing (Perring
& Walters 1976; Stace 1991). The Irish station is an area of some 12 X 3 mona south-facing slope of
submaritime grassland interspersed with a few outcrops of Old Red Sandstone. The species-rich
sward, dominated by Agrostis vinealis Schreber, Cynosurus cristatus L. and Koeleria macrantha
(Ledeb.) Schultes, is grazed to 5—8 cm, with grass-stems and chewed inflorescences of D. armeria up
to 18 cm. Further down the slope the grassland is coarser and enriched, with patches of nettles and
thistles. On adjacent rock outcrops a more heathy flora, dominated by Ulex gallii Planchon, includes
a large, unrecorded population of Ornithopus perpusillus L., a rare but often overlooked plant in
Ireland (fide J. R. Akeroyd & R. FitzGerald). No soil analyses have been carried out, but it may be
significant that the site is adjacent to old copper workings.

D. armeria has a wide distribution in Europe, where it extends northwards to southern Norway
and westwards to Galicia, Cornwall and mid-Wales (Jalas & Suominen 1986). Its occurrence in
southern Ireland, although a welcome surprise, is perhaps therefore not too unexpected. It should
certainly be sought elsewhere in W. Cork and on coasts from Co. Cork to Co. Dublin. It is the only
member of the genus Dianthus that is apparently native to Ireland.

Allium ampeloprasum L. var. babingtonii (Borrer) Syme
Sherkin Island, road from church towards Sherkin Point, garden of deserted cottage, July 1992, obs.
K. Clarke.

This distinctive leek has long been known from the Aran Islands and the coasts of Clare, W.
Galway (Webb & Scannell 1983) and from Donegal, and is also widespread in Scilly and the coast of
mainland Cornwall. It is considered to be a relic of ancient cultivation, probably introduced from

‘the Mediterranean region where the species has its centre of distribution, but perhaps a native
species (Webb & Scannell 1983). Var. babingtonii, endemic to western Ireland and south-western
England, is frequently associated with human habitation and old ruins. The Sherkin plants might
have been introduced, but they do provide a geographical link between the two main areas of
distribution of this variant.

Asplenium obovatum Viv. subsp. lanceolatum (Fiori) P. Silva
Cape Clear Island, South Harbour, damp stonework, 3 September 1992, obs. K. Clarke & J. R.
Akeroyd.

186 SHORT NOTES

We observed a single plant of this rare fern at South Harbour, where it had last been recorded, as
A. lanceolatum Huds., in 1896 (Colgan & Scully 1898). The warden of the Cape Clear Bird
Observatory later kindly directed us to a second locality to the west of South Harbour where the
plant had been rediscovered by another botanist on a damp stone wall just a few days previously,
allowing us to confirm the identity of our own plant.

Centaurium pulchellum (Sw.) Druce
Horse Island, damp grassland above patch of scrub on east coast, 10 m, 19 June 1982, L. C. Wright
LW0045, herb. Sherkin Marine Station.

We were unable to refind this species ourselves, but the herbarium material represents the only
recent record from Co. Cork. It is apparently the first record of this species from Roaringwater Bay
since 1818 when it was reported on Cape Clear Island (Colgan & Scully 1898). C. pulchellum has
recently been rediscovered at several old stations in Co. Wexford (fide J. R. Akeroyd & R.
FitzGerald), so may be overlooked.

Kickxia elatine (L.) Dumortt.
Sherkin Island, Foardree, open peaty ground on south-facing slope above sea, obs. J. R. Akeroyd &
K. Clarke, 29 August 1992.

A very rare, submaritime plant in Ireland, now restricted to Counties Cork and Wexford and |

usually found on cultivated land. It was found at Foardree whilst examining a population of Lotus
subbiflorus Lag. discovered there earlier (fide K. Clarke). However, we failed to find K. elatine,
described by Polunin (1949) as “frequent on cultivated ground’’, elsewhere on Sherkin or Cape

Clear Islands, although fragments of a formerly rich weed flora (cf. Polunin 1949, 1950) have |

survived in the islands.

Rumex pulcher L.
Sherkin Island, above Horseshoe Bay, obs. 28 June 1990, 25 August 1992, J. R. Akeroyd.

Ten plants of this rare Irish dock were located in two small colonies at the eastern end of Sherkin
Island. This confirms reports from the area by Polunin (1950) and earlier workers. As the species
has persisted for nearly 100 years on Sherkin, has been repeatedly recorded in Co. Wexford and is a
member of native plant communities (fide J. R. Akeroyd), it cannot be regarded as merely ‘“‘casual”’
(Perring & Walters 1976) and is probably native in Ireland.

Tuberaria guttata (L.) Fourr.

E. Calf Island, rock outcrops in heathland, 20 May 1992, K. Clarke C198, herb. Sherkin Marine |

Station.

This confirms an old record of the species from E. Calf, where it was not refound by Polunin |
(1950). The other Irish stations are all on islands or peninsulas in the extreme west of the country: on |
the coasts of W. Mayo and W. Galway and on Sheep’s Head and Three Castles Head, Co. Cork |
(recorded by J. R. Akeroyd & D. A. Webb in July 1987), respectively some 25 km to the north-west |

and west of E. Calf Island.

In addition to the above reports, we have made or confirmed records on Sherkin Island or
adjacent islands of many rare or local Irish plants, including Althaea officinalis L., Artemisia
absinthium L., Lotus subbiflorus, Trifolium striatum L. (cf. O’Mahony 1979), T. micranthum Viv.,
T. arvense L., Torilis nodosa (L.) Gaertner and Viola lactea Sm. The total number of flowering
plants and ferns on the islands of Roaringwater Bay, including Cape Clear and Sherkin Islands, now
stands at about 500, at least ten of them among the rarest of Irish plants — a flora of remarkable
richness. A full Flora of the area is currently being prepared.

ACKNOWLEDGMENTS

We are grateful to Matt Murphy, Director of the Sherkin Island Marine Station, for providing
accommodation and facilities for our research, and to Michael and Robbie Murphy for their skillful
and cheerful ferrying of us around the waters of Roaringwater Bay. Our work in 1992 on Sherkin

SHORT NOTES 187

Island would never have been possible without the solid groundwork provided by the studies of
Lucy Wright (1981-2) and Jennifer Shockley (1990).

REFERENCES

CoLGAN, N. & ScuLLy, R. W. (1898). Cybele Hibernica, 2nd ed. Dublin.

JALAS, J. & SUOMINEN, J. (1986). Atlas Florae Europaeae 7. Helsinki.

O’Mauony, T. (1979). Inula crithmoides L. and Trifolium striatum L. in the Cork flora. Bull. Ir. biogeog. Soc. 3:
7-10.

PERRING, F. H. & WALTERS, S. M., eds (1979). Atlas of the British flora, 2nd ed. Wakefield.

Po.unin, O. (1949). The Flora of Sherkin Island, S. W. Cork. Typed draft of Polunin’s paper (1950), with letter
from editor E. F. Warburg. Sherkin Island Marine Station.

PoLuNIN, O. (1950). Notes and additions to the flora of the islands of S.W. Cork. Watsonia 1: 359-363.

Stace, C. A. (1991). New Flora of the British Isles. Cambridge.

Wess, D. A. & SCANNELL, M. J. P. (1983). Flora of Connemara and the Burren. Cambridge.

J. R. AKERoYD & K. CLARKE
24 The Street, Hindolveston, Dereham, Norfolk NR20 5BU

PILOSELLA X FLORIBUNDA (WIMMER & GRAB.) ARVET-TOUVET (ASTERACEAE)
IN THE BRITISH ISLES

Stewart (1903) recorded Hieracium auricula L. from an old limestone quarry at Cave Hill, Belfast,
and remarked: “I have known this plant growing on the debris of the Whitewell limestone quarries,
Cave Hill, for at least six years. During that time it has neither increased nor decreased. It partially
occupies an area of about two square yards. The district around it is uncultivated, save for grass.
There are few gardens in the vicinity — none close at hand. I consider it an alien, but how it has
reached here is not easily understood”’. It was first recorded there in 1897 and last seen in 1910
(Wear 1923). Pugsley (1946, 1948) renamed it H. helveolum (Dahlst.) Pugsley. Sell and West (Sell
1967) made H. helveolum a subspecies of Pilosella lactucella (Wallr.) P. D. Sell & C. West. The
Same authors, when writing the account of Hieracium for Flora Europaea (Tutin et al. 1976),
decided it was better put under Hieracium xX floribundum Wimmer & Grab., i.e. Pilosella
floribunda (Wimmer & Grab.) Arvet-Touvet which is where H. helveolum was originally placed by
Dahlstedt as subsp. helveolum.

P. floribunda 1s almost certainly a hybrid between P. lactucella (Wallr.) P. D. Sell & C. West and
P. caespitosa (Dumortrt.) P. D. Sell & C. West, which can spread vegetatively as well as by seed. It
differs from P. caespitosa by its glaucous, less hairy leaves, and from P. lactucella by its much taller
habit.

On 21 June 1991, R. P. Bowman discovered 45 or more plants occupying an area of about 4 xX 2m,
in grass heath with Calluna vulgaris — Erica tetralix tussocks, on the wide verge of the B3056 between
Stephill Bottom and Pig Bush in the New Forest, S. Hants. v.c. 11, GR SU/35.05, herb. R.P.B. It
was the site of military emplacements during the 1939-1945 war, and P. x floribunda may have been
introduced during that period.

It is a good match with the eight plants from the Belfast locality in CGE, and with continental
material in that herbarium.

Pilosella < floribunda (Wimmer & Grab.) Arvet-Touvet in Bull. Soc. Dauph. 1880: 280 (1880).
Syn. Hieracium floribundum Wimmer & Grab., Fl. Siles. 2(2): 204 (1829); Hieracium floribundum
subsp. helveolum Dahlst., Hier. Exsicc. 4: nos. 14 & 15 (1891); Acta Horti Berg. 2(4): 13 (1894);
Hieracium helveolum (Dahlst.) Pugsley inJ. Ecol. 33: 347 (1946); Pilosella lactucella subsp. helveola
(Dahist.) P. D. Sell & C. West in Watsonia 6: 314 (1967).

Rootstock rather thick, sometimes producing leafy stolons. Stems 10—35(—45) cm, erect, rather
Slender, green below, darker above, with numerous long (to 7 mm), pale, dark-based simple
eglandular hairs throughout and with numerous to dense, small stellate hairs and numerous, short,
dark glandular hairs in the upper part. Leaves 50-180 x 6-20 mm, glaucous-green with a whitish

188 SHORT NOTES

midrib; the basal numerous, in a rosette, narrowly elliptical or narrowly oblanceolate, the
outermost rounded at apex, the inner obtuse or subacute and mucronulate, long attenuate at base to
a broadly winged petiole, with few to numerous, long (to 4mm), pale simple eglandular hairs on or
near the margin, and similar ones with red bases along the midrib (sometimes the hairs are found on
the surfaces and sometimes small stellate hairs are present on the lower surface); the cauline 0-3,
similar to basal but smaller; when stolons are present they bear numerous leaves which are similar to
basal but small. Inflorescence of 3-7 capitula, compact; peduncles 3-25 mm, with dense, white
stellate hairs, numerous, short, dark glandular hairs, and few to numerous, long simple eglandular
hairs. Capitula 15—20 mm in diameter. Involucral bracts 6-9 x 1-0-1-5 mm, dark green with whitish
margins, linear-lanceolate, rounded-obtuse at apex, with numerous, very small stellate hairs
particularly on the margins, numerous, short, dark glandular hairs down the centre, and few to
numerous, long, whitish, dark-based simple eglandular hairs. Ligules bright yellow, paler beneath,
glabrous. Styles yellow or slightly discoloured. Margins of receptacle pits shortly dentate. Achenes
2-3 mm, purplish-black.

In Stace (1991) it will key out to P. caespitosa, but differs from that species in its bluish-green
rather than yellowish-green leaves which are not as hairy on the surface. In Europe it occurs in the
north and centre southwards to north Switzerland and the east Carpathians. It is a triploid with 2n =
27. Little is known about its biology.

REFERENCES

Puas.Ley, H. W. (1946). List of British species of Hieracium. J. Ecol. 33: 345-347.

Pucs.ey, H. W. (1948). A prodromus of the British Hieracia. J. Linn. Soc. Lond. (Bot.) 54: 1-356.

SELL, P. D. (1967). Taxonomic and nomenclatural notes on the British flora. Watsonia 6: 292-318.

SELL, P. D. & West, C. (1976). Hieracium L., in Tutin, T. G. et al., eds. Flora Europaea 4: 358-410.
Cambridge.

Stace, C. (1991). New Flora of the British Isles. Cambridge.

STEWART, S. A. (1903). Hieracium auricula, Linn. Rep. botl Soc. Exch. Club Br. Isl. 2: Sie

WEAR, S. (1923). A second supplement to, and summary of, Stewart and Corry’s flora of the north-east of Ireland.
Belfast.

R. P. BOWMAN & P. D. SELL
22 Kennedy Road, Maybush, Southampton, SOI 6DQ

REGENERATING BALSAM POPLAR (POPULUS CANDICANS AIT.) X BLACK POPLAR
(P. NIGRA L.) (SALICACEAE) AT A SITE IN LEEDS

In 1983 poplar regeneration was observed on a patch of waste ground at a site in central Leeds,
Yorkshire. Vegetation surveys carried out in other towns over the next eight years failed to reveal
further examples and it gradually became clear that this was a most unusual occurrence. A literature
search revealed only one other incidence in the UK; this involved different poplar taxa at a sewage
works at Hackney, East London (Wurzell 1985). During 1991 the Leeds population, which is now
very well established, was investigated in more detail.

The parents, which are growing on a traffic roundabout at the junction of Kirkgate and Crown \
Point Road, are 20 well grown specimens of the Balsam Poplar (P. candicans Ait., all female) and
two similar sized Black Poplars (P. nigra L., both male). The origin of P. candicans is unknown; it |
may be a hybrid between P. balsamifera L. and P. deltoides Marshall and, if so, the regeneration |
could well be tri-clonal, a most unusual occurrence. The large quantities of seed which are shed in
late June collect as piles of white ‘fluff’ wherever there is a little shelter. Following rain, they |

germinate within 24 hours to produce extensive, dense swards of seedlings with dark blue-green ~

elliptic cotyledons. Most of these are subsequently killed by drought. In many years however a
number survive, particularly where the substrate contains fine material. This has led locally to very
dense stands of uneven aged young poplars up to several metres high; hundreds of individuals are
involved.

SHORT NOTES 189

The community they are invading is typical of dry, brick rubble demolition sites all over the
country (Gilbert 1989). Leading herbaceous species are Agrostis stolonifera L., Artemisia vulgaris
L., Chamerion angustifolium (L.) Holub, Poa annua L., Senecio squalidus L. and Taraxacum
officinale Wigg. which provide an open vegetation into which woody plants such as Buddleja davidii
Franchet, Fraxinus excelsior L., Malus domestica Borkh., Salix caprea L., Sambucus nigra L. and
the poplars readily self-seed. The only remarkable feature of the vegetation is the presence of the
poplars, now up to 3 m high and visually dominant. The climate of Leeds is not unusual in any way
and open, competition-free conditions are a universal feature of urban areas so the reason behind
this regeneration episode is enigmatic.

The hybrid plants are too young for features such as canopy shape, sucker development or trunk
morphology to be assessed, and to date none has flowered so their sex is unknown. However it is
already clear that they show a greater range of variability in leaf characters than either parent. A
hundred leaves were collected from strong shoots of each parent population and compared with a
similar number from the progeny (Table 1). The results suggest that by using combinations of leaf
characters it should be possible to identify populations that have arisen as hybrids between the black
and balsam poplar. The range of variation in leaf characters is wider than in either parent; the mean
expression of this variation is intermediate between that shown by the parents and new characters
are present such as the subpallid colour of the underside of the leaf (34%), rounded leaf base (20%)
and subacute leaf tip (17%). The variability of the parents needs to be fully understood before such
determinations are made (see Jobling 1990).

The opportunity was taken to investigate seed viability which is reported to drop to zero after only
a few days (Brendell 1990). A large sample of seed, collected from bursting capsules on 29 June
1991, was stored in daylight at room temperature. Initially, then at seven day intervals, subsamples
of c. 200 seeds were moistened and placed on damp filter paper in a petri dish. Germination
occurred within 24 hours but the dishes were left for seven days before being scored. The results

TABLE 1. A COMPARISON OF THE CHARACTERS OF POPULUS NIGRA, P. CANDICANS AND
THEIR HYBRIDS

Character P. nigra Hybrid P. candicans
Leaf shape Triangular 80% Triangular 28% Heart-shaped 80%
Diamond 20% Diamond 39% Ovate 20%
Ovate 26%
Heart-shaped 3%
Elliptic 4%
Leaf base Cuneate Cuneate 80% Cordate-
Rounded 20% subcordate
Leaf tip Acute through Acute through Acute through

Colour of underside
Serration

-Gland(s) at top of
petiole
Hairiness of petiole

Cross section of petiole*

Scent of unfolding
foliage

acuminate to
cuspidate

Green

Crenate

Absent

Hairy 80%
Subglabrous 20%

Flattened

Unscented

* This character was difficult to assess.

acuminate to

cuspidate 83%
Subacute 17%
Green 31%
Pallid 35%
Subpallid 34%
Crenate 52%
Serrate 47%
Entire 1%
Present 52%
Absent 48%
Hairy 49%
Subglabrous 4%
Glabrous 47%
Flattened 25%
Rounded 75%

Scented to some extent

acuminate to
cuspidate

Pallid (whitish)

Serrate

Present 68%
Absent 32%
Subglabrous 84%
Glabrous 16%

Rounded

Balsamic

190 SHORT NOTES

showed that a germination rate of 80% is maintained for the first week, after a fortnight it had
dropped to 25%, after three weeks to 1%, and after five weeks to zero. In this instance it would be
correct to record that germination dropped to zero after a few weeks.

At the London site, where hundreds of selfsown poplars have established at two adjacent disused
sewage works (the Middlesex and the Essex Filter Beds) ecological conditions are very different.
Here the substrate is mud in the bottom of seasonally waterlogged lagoons. The parents involved
are two varieties of black Italian Poplar which have crossed to produce abundant regeneration of the
hybrid P. X canadensis Moench ‘Serotina’ (male) X ‘Marilandica’ (female); the former has also
crossed with Balsam Poplar to produce the hybrid P. X canadensis ‘Serotina’ (male) x P. candicans
Aiton (female).

These examples from Leeds and London are the only recorded instances of alien hybrid poplars,
which are usually present as single sex clones, regenerating in Britain and are a further example of
how new taxa, with their dispersal centre in urban areas, are being added to our flora. In the Ruhr
district of Germany, I have observed that Black Poplars and Balsam Poplars hybridise freely, their
progeny are a conspicuous feature of dry urban wasteland sites in industrial areas. It would appear
that unlike Black Poplar regeneration, which is currently limited by a lack of suitably muddy
germination sites (Milne-Redhead 1990), regeneration involving alien poplars may be controlled by
the chance juxtaposition of compatible genotypes.

ACKNOWLEDGMENTS

I am grateful to E. Milne-Redhead for confirming the identity of Black Poplar and to B. Wurzell and |

A. Beaton for useful discussion.

REFERENCES

BRENDELL, T. (1990). Poplars of the British Isles. Princes Risborough, Buckinghamshire.

GILBERT, O. L. (1989). The ecology of urban habitats. London.

JoBLING, J. (1990). Poplars for wood production and amenity. Forestry Commission Bulletin 92. London.
MILNE-REDHEAD, E. (1990). The B.S.B.I. Black Poplar survey 1973-88. Watsonia 18: 1-5.

WuRZELL, B. (1985). The remarkable tree flora of the Middlesex Filter Beds. Manuscript. The London Wildlife

Trust.

O. L. GILBERT |
Department of Landscape Architecture, The University, Sheffield, S10 2TN

SEXUAL DIMORPHISM IN ERIOPHORUM VAGINATUM L. (CYPERACEAE)

In Britain, Eriophorum vaginatum L. is one of the major dominants in ombrotrophic pS
vegetation and comes into flower in early spring, before most other bog species. On 22 March 1991,
we had an opportunity to observe its floral characters in a central part of Borth Bog (Cors Fochno) in
Cardiganshire (v.c. 46), one of the largest intact raised mires in Britain. Tussocks of E. vaginatum |
are a prominent component of the vegetation at Borth Bog, so there was an abundance of material
on which to make observations.

The florets of E. vaginatum are wind-pollinated and strongly protogynous. Following pollen |
liberation, anthers are shed leaving the more persistent filaments protruding from the glumes.
Eventually, the filaments too are abscised, and the perianth bristles elongate rapidly and massively
to form the familiar cotton-like heads which aid seed dispersal.

What seemed curious about the floral biology of E. vaginatum at Borth Bog was that no stamens
were visible on the flowering spikes of some tussocks, even though they were at the appropriate
ontogenetic stage, with withered stigmas on the one hand and no sign of remnant filaments or
developing ‘cotton’ on the other. Suspicion that these plants were male-sterile was confirmed by
closer inspection; three tiny vestigial stamens, or staminodes, were clustered around the base of the

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Figure 1. Flowers of Eriophorum vaginatum collected from Borth Bog in March 1991: (a) hermaphrodite;
(b) male-sterile (glumes and bristles removed).

Ovary in every floret. These staminodes were clearly non-functional, with minute, empty, non-
dehiscent anthers, white or brown in colour, on filaments which had failed to elongate. The striking
difference between these structures and those of male-fertile plants is illustrated in Fig. 1.

Several male-sterile and male-fertile tussocks were examined in the field, and no variation in
stamen development was detected, either within or between inflorescences on the same tussock.
However, no thorough search for plants of intermediate phenotype was carried out. Similarly, no
attempt was made to assess the abundance of male-sterile tussocks on 22 March 1991, but they were
obviously frequent in the study area. No marked ecological separation of the two sex phenotypes
was apparent.

We re-visited the same section of Borth Bog on 21 June 1991 to see whether the two phenotypes
were setting seed. It was past the optimum time to make such an assessment as fruiting spikes were
disarticulating. Sexing had to be carried out with great care because staminodes were being shed
from male-sterile florets, as were remnant filaments from male-fertile spikes. Many of the remaining
intact fruiting heads of both phenotypes were barren. However, several florets in both male-sterile
and male-fertile spikes were found to contain ripening nuts. It appears therefore that the Borth Bog
population of E. vaginatum is gynodioecious (sensu Darwin 1877), with co-existing female and
hermaphrodite tussocks.

The only previous record of sexual dimorphism in E. vaginatum of which we are aware was
reported in a talk on pollination and seed dispersal in Danish Cyperaceae given by the eminent
botanist C. Raunkiaer to the Botanical Society of Copenhagen at a meeting on 14 May 1892

192 SHORT NOTES

(Anonymous 1893). Raunkiaer stated that he had found EF. vaginatum to be completely
gynodioecious on a small island in the Hvals@lille Sea; unfortunately, no further description of the
sexual characteristics was transcribed. Male-sterility has been reported for other species of
Eriophorum by several authors, including Dickie (1865) and Knuth (1906), but not for E. vaginatum
by Wein (1973).

Further investigations into the distribution of gender in FE. vaginatum and other British
cottongrasses are in progress.

ACKNOWLEDGMENT

We thank Anna Williams for translating the account of Raunkiaer’s talk.

REFERENCES

Anonymous (1893). Meddelelser fra den Botaniske Forening i Kj@benhavn. Bot. Tidsskr. 18: 18-29.
Darwin, C. (1877). The different forms of flowers on plants of the same species. London.

Dickie, M. D. (1865). Notice of two forms of Eriophorum angustifolium. J. Linn. Soc. Lond. (Bot.) 9: 161-162.
KnutTH, P. (1906). Handbook of flower pollination. Translated by J. R. Ainsworth Davies. Oxford.

WEIN, R. W. (1973). Eriophorum vaginatum L., in Biological Flora of the British Isles. J. Ecol. 61: 601-615.

D. P. STEVENS & T. H. BLACKSTOCK
Countryside Council for Wales, Ffordd Penrhos, Bangor, LL57 2LQ

CORYNEPHORUS CANESCENS (L.) BEAUV. (POACEAE) ON THE WEST COAST
OF SCOTLAND

There are three records between 1895 and 1961 of Corynephorus canescens (L.) Beauv. on the
Morar coast. In 1948 R. C. L. Burges collected ‘‘on sands one and a half miles south of Morar’’,
Westerness v.c. 97 in E. If this distance is stretched a little, is there similarity between this record
and that of S. J. P. Waters, 1961, “‘sand dunes by Camusdarach”’ which is slightly more than 3 km by
road from Morar? Or is it more likely that the Burges record refers to the earliest known site at
Toigal which by road is 2:5 km south of Morar? This site of C. canescens was found by F. Townsend,
as Weingaertneria canescens Bernh., on “Sand Dunes by the sea between Morar river and Arisaig,
Scotland, July 1895”’, in herb. Druce, OXF with duplicates in CGE, E, K and LIV. There has been
no further report of this site, except the reference to Burges above, until 24 July 1991 when Alfred
Slack, Elizabeth Norman and John Trist found the remains of the colony.

Beeby (1897) reported that one Eneas R. Macdonnel of Morar introduced Ca canescens to this
site; but this ‘introduction’ is Beeby’s interpretation of Macdonnel who in /itt., 1896 to Townsend
wrote “‘it is not indigenous but was introduced direct here and not by accidental admixture’. This
does not imply that Macdonnel introduced either seed or plant.

In 1991, after a thorough search of the dunes, we found a single plant of C. canescens within the
area described by Townsend. He reported to Beeby (1897) that ‘‘the grass occurs in plenty on the
sand-hills’. At Morar the dunes are highly mobile and bare of vegetation except for Ammophila
arenaria (L.) Link, which is largely confined to the crests, and a little Carex arenaria L. At the site of
the single C. canescens there was one plant each of Aira praecox L., Rumex acetosella L. and Carex
arenaria which were within 10 m of a small dune crested with Calluna vulgaris (L.) Hull.

Marshall (1967) reports that Corynephorus canescens is a plant of substrates which are extremely
low in mineral nutrients, and this is true of this site at Toigal, which showed available phosphorus
4 mg/l ', potassium 9 mg/l” ' and magnesium 7 mg/I_'. These values are very low and would offer
minimal plant nutrition.

The pH of the Toigal sand is high at 8-3. It was treated with dilute hydrochloric acid and shown to
have no calcium carbonate present, indicating that there must be another source of calcium. This
was confirmed by the detection of ammonium acetate extractable calcium in moderate quantity.

SHORT NOTES 193

This sand consists of multi-faced subangular particles of quartz, some of which are transparent,
opaque and ferrugineous; minute black particles of a mineral are also present and may represent 1—
2% of ferro-magnesian minerals.

The instability of these dunes has probably been a major influence in the past history of this
Corynephorus colony. Consideration may also be given to Marshall (1967) who has shown that this
taxon can thrive in a wide range of pH (3-7-8-5). Perhaps in this case we should only regard the high
pH value as one of several factors which have influenced the gradual decline of this grass colony.

C. canescens has a shallow rooting system of 7-5—10.cm in depth (Marshall 1967). In this site on the
coast it is subject to high winds which puts further stress on the sand to retain moisture. However
this area has an annual rainfall of c. 1500 mm (Met. Office 1991) which to some extent will offset the
disadvantages of a medium which lacks moisture storage. C. canescens grows best where up to 10cm
of sand accretion per year takes place though Marshall points out that where this exceeds 2 cm at
germination too much sand may be trapped and seedlings are smothered. Even in this event some
young plants may survive which have not germinated from seed and which owe their existence to
vegetative internodal growth.

Rabbits may have contributed to the decline. The single plant of C. canescens of 1991 had twelve
culms of which only three panicles survived the rabbits. Reference has been made to the sparse
vegetation of the dunes which would not attract rabbits. There are no rabbit-grazed swards about
the dunes but they are found around the settlement. As rabbits have probably been present here for
a long time, it would be difficult to assess their part in the decline of the C. canescens colony which
has survived for 97 years and now appears to be reduced to a single plant.

ACKNOWLEDGMENT

I thank Dr Bryan Davies for soil analysis and guidance in unfolding the calcium source.

REFERENCES

Beesy, W. H. (1897). Weingaertneria canescens Bernh. Rep. Botl Soc. Exch. Club Br. Isl. 1: 501-502.
MarsHaLL, J. K. (1967). Corynephorus canescens (L.) Beauv. J. Ecol. 55: 207-220.
METEOROLOGICAL OFFICE (1991). Rainfall statistics. Bracknell.

P. J. O. TRist
Glovers, 28 High St, Balsham, Cambridge, CBI 6DJ

Ler eioas

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Watsonia, 19, 195-200 (1993) 195

Book Reviews

The correspondence of Charles Darwin. Vol. 7: 1858-1559. Edited by F. Burkhardt & S. Smith.
Pp. xxxv + 671. Cambridge University Press, Cambridge. 1991. Price £35 (ISBN 0-521—38564—4).

463 pages of this volume are taken up with the correspondence during the dates given; a further 34
pages record letters located or re-dated since the publication of Correspondence vols |—6 (covering
the years 1821-1857). The eight appendices include a chronology, an abstract of Darwin’s theory of
natural selection and a copy of Wallace’s ‘On the Tendency of Varieties to depart indefinitely from
the Original Type’, which forced Darwin’s hand into publishing the Origin. Of the other
appendices, one concerns a memorandum to H. M. Government about the status of the nation’s
premier natural history collections vis-a-vis the British Museum — plus ca change . . . In addition,
there are comments on the manuscript, a bibliography, a biographical register and an index.

Of course, the nub of this volume is Darwin’s preoccupation with his book On the Origin of
Species by means of Natural Selection, the publication of which (1859) had to be brought forward as
a result of Wallace’s letter from the island of Gilolo in the Moluccas. As ever, there is much
correspondence with his friend Joseph Hooker and his mentor, Charles Lyell, which reveals
Darwin’s doubts about rushing into print over his theory — would that some modern workers might
so reveal their innermost feelings about their pet ideas. Nor does Darwin seek succour solely from
his great contemporaries; many of his letters seek information on points of detail from a host of less
noteworthy correspondents.

Apart from the scientific ‘dynamite’ enshrined in this correspondence, however, the reader
continues to be given access to a view of a stratum of Victorian life scarcely touched upon by
Dickens, for example. Nevertheless, the death, from scarlet fever, of his youngest child, Charles
Waring, at the age of two years is a reminder of the universal perils of the age. The agony of this
event, revealed in a letter to Hooker, is counterpointed by a letter to Darwin’s eldest child, William,
about a younger (George) son’s prowess at billiards.

With its revelations of the ways in which a great scientist arrived at one of the seminal biological
theories of this or any other age, together with a view of the society within which such ideas
developed, this volume is of widespread interest and value. It continues the feast of high scholarship
which is epitomised by this distinguished editorial and publishing treatment of Darwin’s Correspon-
dence.

D. M. Moore

Genetics and conservation of rare plants. Edited by D. A. Falk & K. E. Holsinger. Pp. xviii + 283.
Oxford University Press, Oxford. 1992. Price £35 (ISBN 0—19-506429-1).

Rare species are on the increase; the public expects botanists, who have highlighted the problem for
so long, to develop workable schemes that prevent the extinction of endangered species. This book,
resulting from a 1989 conference convened by the Centre for Plant Conservation in St Louis, helps
-to ensure that such schemes have a strong scientific basis. The complementary approaches of ‘off-
site’ and ‘on-site’ conservation are examined. Limited space is given however to the complexities of
re-introduction.
Building on what is known of rare plant population biology, the book then considers methods for
sampling, assessing and conserving their genetic variation. Finally the conclusions are drawn
together into a set of practical guidelines for conservation programmes. One of the strengths of the
book is that it calls on expertise in the fields of zoology, forestry and crop genetic resources. Here
the authors have successfully focussed their expertise on the specific problems faced in conserving
rare plants. It is a pity, however, that the chapter on germplasm management failed to emphasise
such practical problems as ‘empty’ seeds in seedlots of wild species.

196 BOOK REVIEWS

There are some very valuable contributions from leaders in the field of plant population genetics.
Hamrick ef al. point out the relative benefits of using morphology, isozymes and DNA studies in
assessing genetic diversity. In these ‘high tech’ days, it is comforting to know that measurements on
‘real’ plants still have a valuable part to play in such assessment.

Not surprisingly, the book draws mainly on examples from the Developed World. In the.

Developing World, the ratio of botanists to species means that the biology and even rarity of a
species is often not known. This makes development of conservation programmes difficult. Broad
generalities help. Biodiversity within wet tropical forests will be best conserved ‘on-site’ (see
contribution from Bawa & Ashton). In the dry tropics, however, where desertification threatens,
long-term conservation may often only be achieved through ‘off-site’ activities. The broad
statement on page 114 that tropical species’ seed cannot withstand drying and cold storage is
erroneous; many dry tropical species are quite amenable to seed storage. With the clock ticking,
there will be little time left for detailed studies in many cases to determine the best conservation
strategy. Indeed, many of these studies will only be possible once material has been taken into safe
keeping in seed banks and botanic gardens; so by using the sampling guidelines given in this book we
should conserve something now and worry about the precision of the approach later on.

In summary, this volume has been well edited, has a wealth of information bringing together some
850 references and has been produced to a high standard. A proportion of the book is accessible to
the general reader; the more genetical elements might have been more so had a glossary been
included. The book is an important step towards practical action. Botanists and conservationists will
find it a useful addition to the literature.

S. H. LININGTON

The Hamlyn photographic guide to the wild flowers of Britain and Northern Europe. R. Gibbons &
P. Brough. Pp. 336, including 163 pp. of colour plates. Hamlyn Octopus, London. 1992. Price £20
(ISBN 0-600-57452-0).

Here is yet another illustrated Wild Flower book, but this time it is hardly a field guide. Its size, 22 x
29 cm, is too large for most pockets, and its weight, 1-63 kg, is too heavy for most rucksacks after
packing waterproofs, food, camera, etc. It must therefore be considered as an indoor reference
book, as even a quick glance showed it to be more useful than the proverbial coffee table book.

After the general acknowledgments and bibliography are two pages of introduction and

explanations on using the book, and two pages of glossary terms with diagrams. Then follow 320
pages with descriptions of over 1,900 species and 1,500 colour photographs, 1,000 distribution maps
and more than 400 line drawings of important identification characters.

The text and illustrations are arranged according to the systematic order used in Flora Europaea,

and with a few minor exceptions the nomenclature also follows Flora Europaea. Grasses, sedges and. |

rushes are not included.

The distribution maps, text, line drawings and photo index on the left-hand page and 8-12 |
relevant photographs on the right-hand page do make reference easy. The photographs are lettered |

in a diagram of the plate layout, and the letter identifies the map, text and index of photographs.

Where possible the English name ts used in the marginal photographic index. Both the scientific and _ |

English names are given in the text.

On the whole the photographs are good. Some appear to have been selected to show important
characters separating ‘look-alikes’ such as Potentilla sterilis and P. micrantha. It is a pity that a few
photographs are enlarged more than other members of the same genus on the same plate. This could
be confusing to a beginner, even though size may be given in the text. Two examples are the smaller
species of Cerastium, and Sedum villosum is so much smaller than S. telephium.

The distribution maps, though of necessity tiny, are an added interest and the colour codes give an
idea of the status of the species where it does occur. The area covered is about the same as in
previous Floras of Britain and Northern Europe.

The line drawings are necessary for some species as even good photographs cannot show smaller
critical features, but there is space in the margins for a few more. A beginner with an unknown

flower and no knowledge of plant families will be faced with the daunting task of turning page after |

ee a a

BOOK REVIEWS OG,

page in search of a photographic match. There is no key, but descriptions of families appear in
systematic order in the text and species in the larger genera are grouped under headings of similar
characters. The last six pages contain a combined index of English and scientific names, which
makes for easy reference.

A few errors are inevitable in such a work, but the index has over 20 omissions.

While I find the book too large and heavy for use in the field, I certainly enjoy the photography
and find the distribution maps interesting, though a few do not agree with the distributions given in

the text.

V. GORDON

Bob Press's field guide to the trees of Britain and Europe. J. R. Press; photographic consultants E. &
D. Hosking, artwork by M. Tebbs. Pp. 247, with numerous colour plates and black and white
illustrations. New Holland, London. 1992. Price £17.95 hardback (ISBN 1—85368—103-—2); £9.99
paperback (ISBN 1—85368-104—0).

Field guides are a popular theme for publishers, and this is a fine example of the category. Richly
illustrated with colour photographs and line drawings, in a format which allows the inexpensive
production of versions in other languages, and with enough information to allow a reasonable
chance of successful tree identification, this book is likely to prove very popular. The simplicity of
style is similar to that of Oleg Polunin’s, but Bob Press has been able to use the pick of the
photographic agencies rather than relying mainly on the work of one person.

The coverage of tree species is wide, with over 450 species treated; native and naturalised species
are given equal emphasis, with all the main species illustrated by at least one colour photograph.
Line drawings in the left hand margin emphasise smaller features helpful for identification, and
there is a short glossary of terms. Keys are in two forms: a numbered dichotomous key relying on
both foliar and floral characters, and a synoptic key based entirely on leaf characters. The 52
families are described briefly in the introductory pages, facilitating comparisons and avoiding
interruptions to the text. Text descriptions include a rather generalised indication of the distribution
of the tree in Europe, followed (for naturalised species) by an indication of their country of origin.
There is an appendix with information on arboreta, and a short bibliography. Separate indexes to
common names and scientific names are provided.

The high quality of reproduction of colour photographs now achievable by modern publishers is
well displayed in this inexpensive book, which was printed in Singapore. Taxonomically up-to-date,
and concisely written, the book is unfortunate only in its title. Why, in 1992, does a publisher choose
to refer to “Britain and Europe’’?

J. R. EDMONDSON

Biology of plants. 5th edition. P. H. Raven, R. F. Evert & S. E. Eichhorn. Pp xvii + 791; lavishly
illustrated. Worth Publishers, New York. 1992. Price $59.95 (ISBN 0-87901—52-2).

This substantial book is the latest in a long tradition of all-embracing introductory Botany
textbooks. It is a marvellous introduction to botanical science, with a well organised and very
teadable text supported by excellent diagrams and photographs, almost all in colour, on nearly
every page.

Biology of plants has 31 chapters divided into six sections covering cell biology, genetics,
diversity, anatomy, physiology and ecology. Scattered throughout the book there are also essays,
separated from the main text, discussing topical themes such as ‘The Great Yellowstone fire’ or
“Jobs versus owls’. The level of treatment is generally aimed at the first year university student, and
is designed primarily for its North American home market. This is only really noticeable in the
chapter covering vegetation types, which perhaps leans too heavily on North American case
histories to the exclusion of more representative examples elsewhere in the world.

As would be expected, recent advances in plant physiology and molecular biology are well

198 BOOK REVIEWS

covered, but there are also some interesting reinterpretations of long-known phenomena; for
example, the ‘infection’ of grasses by ergot is now seen as a partnership in which the grass receives
protection from herbivores by the toxic fungus.

My main criticism of the book concerns the authors’ definition of what is a plant. They exclude
algae from their Kingdom Plantae, leaving only the bryophytes and vascular plants. The green algae.
are thus separated from their descendants, and giant kelps are left as odd bedfellows to the
unicellular Protista. Despite this divorce, the actual coverage of the algae, as well as the more
traditional ‘non plants’ — viruses, bacteria and fungi — is one of the most comprehensive of any
introductory textbook I have seen.

At around £40 for 800 pages Biology of plants is an excellent value textbook from A-level
onwards, but it would be a shame if its only audience were students. It is an excellent introduction to
the world of plants for anyone, and with superb plant paintings by Rhonda Nass and a beautiful
cover, courtesy of Van Gogh, it might even find its way on to one or two coffee tables.

A. S. GUNN

British plant communities. Vol. 2: Mires and heaths. Edited by J. S. Rodwell. Pp. x +. 628.
Cambridge University Press, Cambridge. 1991. Price £95 (ISBN 0—521-39165-2).

In 1989, I was, for all too brief a period, the last appointed Chief Scientist of the Nature
Conservancy Council (R.I.P.). While in the post, however, my most pleasant task was to bring the
National Vegetation Classification to its triumphal climax as a manuscript to be transmitted to the
publishers, Cambridge University Press. Last year I saw with pleasure the first volume (reviewed in
Watsonia 19: 49, 1992). Unlike volume 1, which dealt with woodlands and scrub, this second volume
is a truly ‘British’ volume, in that it deals with vegetation that is centred on Britain, and which makes
Britain special. Here is a tremendous compendium of information on mires and heaths — long
needed, but it will also be long used.

The format seems well suited to the task. The community descriptions ilo not lend themselves
readily to review, except for the passing comment that they are very thorough. The separate
introductions to mires and heaths are well crafted, and excellent synopses, although there are some
dense passages. For example, “The perspective looking towards the Continent from our own
generally oceanic standpoint is rather different from that hitherto proclaimed as normative from the
opposite direction” (p. 350).

At 38 plant communities, one might query if the mire classification were not too fragmented. The
difficulty of identifying homogeneity in mire vegetation, as well as the variation in substrate and
climate across Britain, all contribute to mires being such rich systems, and explain why so many
units are described, I found the ‘block’ diagrams gave an instant clarity of view for ecological
position. The discussion of the changing community context of Schoenus nigricans is also of interest
to illustrate that communities are but spatial and temporal kaleidoscopes of species. But here is also
a weak spot — for it would be marvellous to see that discussion set in a broader context of Ireland and
the western fringe of Europe north and south. Given the 20 year support from the (J.)N.C.C., and
the fact that they have an international branch, it would be good if future volumes could take a wider
view. Even if that is not possible, I hope someone will eventually fund and produce a synthesis of
British plant communities in their European context.

With regard to heaths, the point is well made that without appropriate management heaths
become very hard to distinguish, and tend to a uniform Callunetum; which has clear conservation
implications. It also underlines the very dynamic nature of these communities. Again, the
introductory description is sound, and the block/circle diagrams are helpful to explain lines of
variation. However, I believe it is simplistic to suggest that lowland dry heaths are a linked circle; for
my money there are two clear groups corresponding to the Ulicetalia minoris and Vaccinio-
Genistetalia, as discussed by Bridgewater (1980) in Phytocoenologia 8: 191-235. The Ulicetalia
minoris is a southern and western order and the Vaccinio- Genistetalia an eastern and northern
order, with some overlap between.

Boundaries between wet and dry heaths probably deserve more discussion throughout the

volume. So too does the position of Erica ciliaris heaths, which are given rather short shrift in this |

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BOOK REVIEWS 199

current treatment. The associations Ulici gallii — Ericetum ciliaris and Ulici minoris — Ericetum
ciliaris described by Bridgewater in 1980 I still regard as valid, but vegetation represented by these
associations is not even accorded variant status in this current work. Others will be able to test these
different views — provided it is done in a fully European context. As the author says (p. 13) “‘For the
classification is meant to be not a static edifice, but a working tool for the description, assessment
and study of vegetation’. Indeed, I sincerely hope that one by-product of the series will be an
increase in phytosociological papers in the British journals, as well as the expansion of British
papers in the continental ones.

Keys are provided for both mires and heaths. Do they work, and are they worth the effort? I used
them with sample data collected during the last three years. The results? Success every time, despite
some rather obscure wording of the couplets.

This is a book any botanist interested in phytogeography and vegetation will want to buy. And so
here is my biggest gripe, aimed not at the superb work done by the contributors, but at Cambridge
University Press. Having hired excellent and far-sighted editorial staff to secure works like this for
their list, why then price them out of most libraries’ reach, let alone interested botanists? If I want to
buy the book here in Australia, the price is $325!

But enough of gripes — this work is elegant testimony to those who developed the concept, and
participated through the years, but especially to John Rodwell who has the fortitude to have ridden
out the all-too-many rough patches, when it seemed the end would never come. And from my
personal knowledge, Lynne Farrell, once of the N.C.C. and now English Nature, did an excellent
back-room job, steering it through the shoals of Government bureaucracy. We should all look
forward to the third volume.

P. B. BRIDGEWATER

FLORA: The Computerised key to 786 species of British wild plants, version 1.10. Poly Enterprises
Plymouth Ltd, Seale Hayne Faculty, Polytechnic South West. Newton Abbot. 1992. Price £99.88,
with discount for multiple copies.

Recent years have seen a great increase in general use of computers and botanists have been quick
to explore the possibilities to which computer technology can be put to aid their studies. Several
programs have been developed with the naturalist in mind, building databases for use in plant
recording, mapping, etc., and some have delved into the realms of plant identification. Computer
aided keys have appeared for specialist groups (e.g. sedges and orchids), but FLORA 1s the first
published attempt at a general usage computerised key to British flowering plants.

The FLORA package contains a disk (with the databases and program), a User Instruction
Manual and a botanical reference manual. Once loaded on to the hard disk of your machine (taking
up a tiny 260K) the menu-driven program enables plants to be identified using a multi-access
approach, with a total of 48 possible characters to choose from. Characters are grouped into
categories, according to the approach of the user. Three general categories (primary, secondary and
tertiary) hold groups of characters in descending order of their usefulness in identification. The
characters are also grouped into categories relating to the flower, leaf, stem or environment. In
general, identification begins in the primary category, and character states are entered against those
characters evident from the specimen. For novices to the program there is a ‘beginners tour’ leading
you through all the characters in the primary category showing all their possible states. When a few
characters are entered the database can then be usefully searched for matches. The program sorts
through the database and the species are listed in descending order of likelihood. If need be you can
return to the category summary tables and enter more character states until the search gives a fewer
number of possibilities. At this stage you are advised in the instructions to turn to other botanical
reference books for descriptions and pictures.

Having used the program myself, and enlisted the help of others less experienced in plant
identification, I have found that the program is relatively easy to use, and generally successful. The
authors claim that as few as six characters need to be entered to effect an identification. My
experience is that unless you know which characters will narrow down the field quickly you will
probably have to key in considerably more. This leads me into my first criticism of the program, the

200 BOOK REVIEWS

lack of on-line help facilities. It would be very useful to be able to list the characters that will
differentiate effectively between your shortlist of species, and thereafter concentrate your efforts.

With the great diversity in plant form across the families it is very difficult to produce a general list
of characters and their states that will deal with all species in a satisfactory manner. The writers of
FLORA have had to simplify this variation, and on the whole they have produced a workable
system, and have tried to avoid using technical terminology. There are some rather odd character
states; for example Rumex flowers are described as ‘grass-like’, and the character ‘leaf venation’ not
only caters for parallel, pinnate and palmate venation but also for succulence and compound leaves.
In order to save storage space and to increase compatibility with non-graphics supporting
computers, the writers have decided not to include illustrated help screens, but rather to rely on the
botanical reference manual. Even allowing for the promised reprint (initial problems with printing
have left many of the drawings faint and some unusable) the booklet does not to my mind clearly
define the characters and this has led to mis-scoring of characters when testing the program (in the
botanical reference manual the same leaf shape, obovate, is included in both ‘paddle’ and ‘oval’
character states).

As with many partial Floras this program falls down in its depth of coverage. Grasses, sedges,
rushes and gymnosperms are excluded, and critical groups (e.g. Euphrasia and Salix) are treated as
aggregates. Looking at some of the datasets used there are an alarming number of missing data for
the species included. This leads to problems when separating some of the species. Although much of
the terminology is non-technical, a certain degree of botanical knowledge is required to operate the
program and sort out errors of scoring that frequently arise. I can see the potential for this program
in schools and field centres (particularly with the educational discount price) where it could be used
to encourage children to look at and identify plants. If FLORA was considerably cheaper then I
could see more botanists buying a copy, but at the quoted price most field botanists would be better
advised to invest in one (or two!) of the full Floras of the British Isles.

M. F. WATSON

Pe
a re ae ta

| Watsonia, 19, 201-203 (1993) 201

Obituary

JOCELYN MARY LEWIS RUSSELL
(1905-1992)

Jocelyn Russell, the able, lively and lovable personality, died on 14 May 1992, aged 86. She was
born at Ockham, Surrey, on 28 December 1905, the only child of Walter Lewis Castelden and
Charlotte Mary Katherine, née Thring, who died very shortly after. So, Jocelyn was brought up
until she was four by a sister of her father - who came from Canterbury and managed Henderson’s
Transvaal Estates. By his second marriage he had four children, to the youngest of whom, Mrs
Joanna Huntingford, I am greatly indebted for many details. He died in 1953.

From 1921 to 1923 Jocelyn was sent to the Godolphin School at Salisbury, where her
contemporary was another enthusiastic botanist, the late Mrs Barbara Garratt. This was followed
by an art school and work in her father’s office. On 1 January 1935, she married Basil Henry
Sackville Russell and they lived in Manchester, Portugal (1937) and the Belgian Congo (where he
was employed by Vacuum Oil), before returning to England at the outbreak of World War IT. Basil,
later a Squadron-Leader, joined the R.A.F.V.R. and Jocelyn the Mechanised Transport Corps.

After the collapse of France, her unit was drafted to the Middle East, joining in January 1941, at
Cairo, the Free French Forces with the British 8th Army. As those will know who saw the television
programme on 29 March 1989 (filmed 18 months before) ‘‘Tin Hats and Silk Stockings’’, she also
painted wild flowers when she was out there. Some 40 of these, dating from 1941 to 1944, with names
and localities, were on show at the Centenary Exhibition at Kew in 1953, who said they made a very
fine display and asked if they could eventually be returned there. Many were made at Bir
Hukayyim, where the battle raged for a fortnight in 1942. She made a notebook (which was shown
on the TV programme) grouped by families, of 115 of the plants she saw during the time before she
got to Tunis. For each she wrote out a full and detailed account with a clear, characteristic coloured
drawing. This material has also gone to the Royal Botanic Gardens, Kew.

Her arduous tour lasted four years and three months, from Syria to Tunis, then into Italy and
southern France; and her stalwart service was rewarded on 19 July 1944, with the French Colonial
Medal with clasp ‘Libye’ and on 20 June 1945 with the clasp ‘Tunisie’, and on 16 April 1947 with the
Croix de Guerre avec étoile en bronze. The citation included “‘A participé a toutes les campagnes,
faisant des liaisons aux postes avancés. A notamment assuré dans les combats de Tobrouck, le
ravitaillement en eau de la formation, malgré les mitraillages et les bombardments de !’aviation
ennemie, a continué ces missions de liaison avec la méme succes et la méme intrépidité”’; and on
27 July 1947 the Commemorative Medal of the Free French Forces.

While she was in Tobruk, she learnt that her husband, dropped by parachute from a flying boat in
Sumatra in 1942, shortly after meeting her in Cairo, was missing, presumed killed. Meanwhile, she
had met Constance Spry, the flower arranger, who appeared one day in her tent in Africa and
offered her a job after the war. This developed into a close friendship, Jocelyn travelling widely in
her Rednose cars, demonstrating the art. She even did Prince Philip’s private bouquet for Princess
Elizabeth on their wedding day, which she delivered personally in her Morris 8 to Buckingham
Palace.
_ Although she had been keen on flowers ever since she was a child, she learnt of the Wild Flower
Society only by chance when she was 43. She joined the B.S.B.I. the following year, 1950, and the
London Natural History Society in 1951. Thereafter these led her dominant interest in wild flowers,
she contributing every bit as much as she learnt, sending records to various Floras and the like. Not
surprisingly she rapidly took a leading part. For our Society she gave long service to the Meetings
Committee (1953-70), the Conservation Committee (1964-78) and the Council. She was a Vice-
President from 1976 to 1980. For the Wild Flower Society she held various posts, not the least being
her founding, in 1957 of the Branch ‘Parnassus’, the highest refuge for top members, running it until
1986.

Two of her discoveries were Dichondra micrantha in 1955, naturalised near Hayle in Cornwall

202 OBITUARY

FLOWER

SOCIETY

FicureE 1. The design for the front cover of the Wild Flower Magazine drawn by Jocelyn Russell in 1959.

and new to the British Isles; and while in Jersey she realised what none of us had, why Ranunculus
paludosus was so called, when it was always seen in flower in dry places. She showed that its habitat
had to be wet in winter.

In 1959 there was a competition for a design for the front cover of the Wild Flower Magazine, and
her garland with a flower for each month was the outstanding entry (Fig. 1). Since then it has also
been used on the front cover of the Wild Flower Diaries, the Compliments Slips, and even the Car
Sticker. In 1961 appeared R. W. Butcher’s New Illustrated British Flora, for which Jocelyn did many
of the plates, Dr Butcher describing them to her as “beautiful”, and to me as “‘very good’’. She left
numerous delicate and delightful coloured drawings of flowers (garden ones too) from the countries
she visited. Those with European flowers have been gifted to the Natural History Museum; those
from North and South Africa to Kew. She had coloured in exquisitely, and with date and place,
every possible species in the ///ustrations to Bentham & Hooker’s Flora and most of those in Dr R.
W. Butcher and F. Strudwick’s Further Illustrations of British Plants. These are kept, as the
treasures they are, by her family. They were kindly lent in November 1992, along with a sample of
her drawings, and shown at the A.G.M. and Tea Party of the Wild Flower Society; and also a
fortnight later at our own Exhibition meeting.

OBITUARY 203

Among the papers of this eager searcher are her notebooks from 1951 to 1968, detailing where to
find all manner of scarce, rare and very rare plants (and she had seen the rarest of plants, including
Epipogium aphyllum quite soon after it was rediscovered). There are others with copious notes on
what she found on her trips, often with itineraries which are now in the Natural History Museum.
These drawings and photographs show that she botanised at least in Scotland (1950-55, 1958, 1968,
1974-5); Wales (1952, 1954, 1956-7, 1968); Ireland (1952, 1954, 1958, 1967); Jersey (1954, 1956-7);
Guernsey (1956-7); Alderney (1956); Herm (1958); France (1946, 1950, 1958, 1963-5); Andorra
and Pyrenees (1963); Portugal (1968); Italy (1944-5, 1963-66, 1972); Austria (1964); Switzerland
(1964-8); Norway (1963); Greece; Turkey (1976); and South Africa (1969). She was in touch with,
and often close friends of, many of the leading botanists’ professional and amateur.

She was a skilful photographer, the evidence being in box after box, at least ten of them filled with

2 X 2slides, these going similarly to the Natural History Museum and Royal Botanic Gardens, Kew.

Bald facts such as all these give no impression of the warm, exuberant, cheerful companion she
was, with infectious enthusiasm. What a lot of good she did to, and for, many of us. There are
accounts of her in the Wild Flower Magazine Nos. 341, for September 1954, 407 for Autumn 1986
and 425 for Autumn 1992.

D. McCLINTOCK

Watsonia, 19, 205-207 (1993) 205

Report

ANNUAL GENERAL MEETING, 9 MAY 1992

The Annual General Meeting of the Society was held in the Jodrell Laboratory Lecture Theatre,
Royal Botanic Gardens, Kew, at 10.45 a.m. 74 members were present. Dr P. Macpherson,
President, taking the chair, welcomed members, particularly any who had joined in 1991 or who
were attending their first A.G.M.

Apologies for absence were read and Minutes of the 1991 Annual General Meeting, published in
Watsonia 19: 67-71 (1992), were approved and signed by the Chairman.

REPORT OF COUNCIL

The adoption of this report, which had been circulated to members, was proposed from the Chair.
Mr R. G. Ellis noted that the speciality of the late member Mr M. C. Clark was mycology, and with
this amendment the Report was accepted unanimously.

HON TREASURER’S REPORT AND ACCOUNTS

The Hon. Treasurer, proposing the adoption of his Report and Accounts, which had been circulated
to members, highlighted the B.S.B.I. Database (Leicester), for which the hardware, at an
expenditure of £6,000, was now in place. Mr Walpole acknowledged with thanks the hard work of
our members and staff at the University of Leicester, whose enthusiasm had ensured the successful
input of the initial project, the List of vascular plants of the British Isles by D. H. Kent which was
now ready for publication. The Treasurer also commented on the small number of resignations in
spite of a rise in the subscription rates this year, and he invited queries on the accounts. There being
none, the Report and Accounts were accepted unanimously.

RE-ELECTION OF HON. GENERAL SECRETARY AND HON. TREASURER

Proposing the re-election of these Officers the President stated that the Society was fortunate to
have two such hard-working Officers, thanking Mrs Briggs for tackling with good humour the
continual flow of mail, the administration, and the dispersal of information including regular notes
in B.S.B.I. News which informed members of the Society’s activities and projects. He also thanked
Mr Walpole for his hard work on the finances, including the V.A.T. and the Accounts, and for his
wise counsel on Committees and Council. Both were then re-elected with acclaim and applause.

ELECTION OF COUNCIL MEMBERS
In accordance with Rule 10, nominations had been received for Mr F. Horsman, Mrs A. Lee and Mr
P. Thomson. Profiles had been published oe the Annual Report and these members were elected
unanimously by the meeting.
ELECTION OF HONORARY MEMBERS

Council had nominated two members: proposing Mr J. F. M. Cannon, DrN. K. B. Robson outlined
Mr Cannon’s long years of service to the Society as a past-President, as a member of three of the

206 REPORT

Permanent Working Committees, and also as a general Umbelliferae referee. On the staff of the
Natural History Museum, Dept. of Botany, and later as Keeper of Botany there, Mr Cannon was
closely connected with B.S.B.I. activities centred at the Museum (the official address of the
Society). Proposing Mr E. S. Edees, Mr A. Newton had sent a note which was read to the meeting
reminding us that Mr Edees had been a member of the B.S.B.I. for 60 years, and v.c. Recorder for
Staffs. for 35 years. Author of Flora of Staffordshire, the first county Flora to be based on tetrad
mapping, and joint author with A. Newton of Brambles of the British Isles, Mr Edees had also been
a Rubus referee for more than 30 years, and his nomination was strongly supported by the local
botanists in Staffordshire. Their election as Honorary Members was unanimous and greeted with
applause.

RE-ELECTION OF HONORARY AUDITORS

The Hon. Treasurer, in proposing the re-election of Grant Thornton, West Walk, Leicester,
referred again to the honour for the Society to present their Accounts over the name of these
distinguished Auditors, and the re-election was passed unanimously with appreciation.

ANY OTHER BUSINESS

The Hon. Treasurer added thanks to the Secretaries and Chairmen of Committees, and all who had
assisted with the running of the Society’s affairs; and as Chairman of the Publications Committee he
warmly thanked the Editors of the Society’s Journals and publications, whose considerable work
was very much appreciated by the Society.

The President announced that Council had nominated Dr F. H. Perring as President-elect 1993-5
for election at the Annual General Meeting, 1993. Dr Perring replied that he was deeply honoured
and that he would take up this office with serious commitment.

On behalf of the Society the President recorded congratulations to Professor C. A. Stace on the
publication of his New Flora of the British Isles, an achievement in which we could share in that the
author’s published thanks listed many B.S.B.I. members, and he had also acknowledged the Society
as ‘‘a great source of inspiration to him’’. General appreciation was confirmed by the applause of the
meeting.

The Hon. General Secretary was seeking help with the heavy burden of the administration and |

correspondence and those present were requested to let her know of any volunteers.
There was no other business and the meeting closed at 11.15 a.m.

Mary BrIGGS

PAPERS READ AT THE ANNUAL GENERAL MEETING

Two papers were presented at the A.G.M. Dr P. Macpherson gave the Presidential Address on
Colonisation of the Glasgow Garden Festival site three years on: implications for recording (see
Watsonia 19: 169-179) and Professor G. Ll. Lucas spoke on Plant management for conservation
purposes.

Thanks were recorded by J. Ounsted and P. S. Green, Vice-Presidents, to Professor G. T. Prance,
Director, Royal Botanic Gardens, Kew for permission to use the Jodrell Laboratory, and to the
President and to Professor G. LI. Lucas for their Address and Paper.

The President then thanked all those involved in the arrangements for this successful meeting,
especially Dr D. A. Simpson, who with Mrs R. Simpson had organised the bookings and
refreshments, and who with Mr P. C. Boyce had guided members on tours round the Gardens in the
afternoon.

M.B.

|
| .
|
|

REPORT 207

FIELD EXCURSION HELD IN CONJUNCTION WITH THE A.G.M.
KEW GREEN AND RIVERSIDE, SURREY (V.C. 17). 10 MAY 1992

In all, 45 people attended this meeting associated with the A.G.M. at the Jodrell Laboratory, Royal
Botanic Gardens, Kew. The first part of the day was spent crawling around on Kew Green (some 40
botanists’ bottoms were seen) searching for the small plant species that grow on this sandy
grassland. It was too early for some of the more notable clovers but we did find Trifolium
subterraneum L., Ornithopus perpusillus L., Montia fontana L., Rumex acetosella L., Stellaria
pallida (Dumort.) Piré and Aphanes inexpectata Lippert. On our way to St Anne’s Churchyard we
saw Rumex pulcher L. The churchyard has long been known for interesting naturalized species and
we saw Phytolacca acinosa Roxb., Sisymbrium strictissimum L. and Sisyrinchium striatum Smith in
bud, young plants of Galactites tomentosa Moench becoming established and lots of Soleirolia
soleirolii (Req.) Dandy and Atropa belladonna L. in flower with both Allium paradoxum (M. Bieb.)
Don and Ranunculus ficaria L. subsp. bulbifer Lambinon demonstrating bulbils. During lunch
several people saw Saxifraga granulata L. also with bulbils, behind the cricket pavilion; it was 45 cm
tall. After lunch the march along the river bank was rather poorer in interest but still managed
Angelica archangelica L., Oenanthe crocata L., Ceratochloa carinata (Hook. & Arn.) Tutin,
Tragopogon pratensis L. subsp. pratensis, Diplotaxis muralis (L.) DC. and Bolboschoenus
maritimus (L.) Palla in its highest site on the tidal River Thames but we failed in our attempt to see
Barbarea stricta Andrz. After our long walk, some went on to Richmond Park and others returned
to Richmond or Kew having had a good day; on the way back I saw more than 15 herons fishing in
the river.

J. M. MULLIN

”n
o
fad

te)

INSTRUCTIONS TO CONTRIBUTORS

Manuscripts must be submitted in duplicate, typewritten on one side of the paper, with wide margins
and double-spaced throughout.

Tables, figure legends & appendices should be typed on separate sheets and attached at the end of
the manuscript.

Submission of manuscripts

Papers and Short Notes: Dr B. S. Rushton, Department of Biological and Biomedical Sciences,
University of Ulster, Coleraine, Co. Londonderry, N. Ireland, BTS2 1SA.

Books for Review: Dr J. R. Edmondson, Botany Department., Liverpool Museum, William
Brown St, Liverpool, L3 8EN.

Plant Records: the appropriate vice-county recorder, who should then send them to C. D. Preston,
Biological Records Centre, Monks Wood Experimental Station, Abbots Ripton, Huntingdon,
ei ASS

Back issues of Watsonia are handled by Dawson UK Limited,

Cannon House, Folkestone, Kent, CT19 5EE 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.I., 68 Outwoods Road, Loughborough, Leicestershire, LE11 3LY.

‘Hill.
one Nomis.

UK ISSIN 00143-1532

Filmset yee

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/e 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 1993-94

Elected at the Annual General Meeting, 15th May 1993
President, Dr F. H. Perring

Vice-Presidents, Mr A. O. Chater, 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. R. Mill, 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). |

Bev FS = thse 5 Fee Re HK Sortie eS

(HAYSBAL MISTGRY)

- 8SEP 1993

PRESENTED i
_BOTANY Lie BA ARY |

Watsonia, 19, 209-229 (1993)

The distribution of the Wild Service Tree, Sorbus torminalis
-(L.) Crantz, in the British Isles

P. ROPER

South View, Sedlescombe, Battle, East Sussex, TN33 OPE

ABSTRACT

The results of a survey of the Wild Service tree, Sorbus torminalis (L.) Crantz (Rosaceae), in the British Isles
initiated by the Botanical Society of the British Isles and the Biological Records Centre in 1974 are summarised
and analysed. It is confirmed that the tree is a useful indicator of ancient woodland and hedgerows and that it
shows a marked preference for two kinds of soil: those derived from clays and those derived from harder
limestones. The reasons for this are discussed as well as the occurrence of the species on other soils. It is
suggested that the very low rate of reproduction from seed is mainly the result of seed predation and that the
northerly limits of its range are influenced by the lower rate of seed production in places with cooler, less sunnier
summers.

The pattern of modern records in England and Wales reflects to some extent the part the tree has played in the
life of the countryside over many centuries. It has been conserved and planted in some areas for its fruit, its wood
and for ornament, but in other areas it is scarcely known by local people and held in little regard. Today it is
usually much commoner in the first of these areas.

Areas where the Wild Service tree grows have been divided into three types: those where it is relatively
abundant, those where it is scattered but widespread and those where it is rare. These often, but not always,
show a correlation with the solid or drift geology. Mapping in this way also shows that there are large areas of
England and Wales where conditions appear suitable but from which the tree has not been recorded.

INTRODUCTION

In 1974 the B.S.B.I. and the Biological Records Centre, Monks Wood (B.R.C.), organised a
national survey of the Wild Service tree, Sorbus torminalis (L.) Crantz (Rosaceae) and in the same
year I became coordinator of the survey.

The purpose of the survey was given by the Biological Records Centre as follows:—

“Wild Service is typical of ancient and undisturbed lowland primary woodland. Whilst the Adlas

of the British flora gives a good idea of its general distribution it may have been overlooked in

some areas. It will be of great use in recognising primary woodland for conservation to associate
its occurrence with individual woods, not just grid squares, and to trace its distribution in
hedgerows.”

The initial phase of the survey was conducted by the distribution of standard record cards (see
Fig. 1) from the B.R.C. to B.S.B.I. members, foresters, reserve wardens and others likely to have
an interest. On return to the B.R.C. these were forwarded to me. Appeals were also made on radio,
television and in the press to members of the general public which resulted in much further
information not only on the distribution of the tree, but on its economic and folkloric roles. Many of
these latter records came from people who were uncertain of the tree’s identity, but leaves or fruit
were sent for confirmation. Over 1000 records, both on cards and from other sources, were
received, many from areas where the tree had not hitherto been reported and even after an interval
of nearly 20 years records still occasionally arrive.

A comprehensive search of appropriate literature was also undertaken and this indicated areas
where the tree had once occurred, and might still exist, but the data have not found their way into
county Floras or other published material. In Pembrokeshire, for example, the tree was clearly well
known to local people long before any formal record appeared in the botanical literature.

P. ROPER

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SORBUS TORMINALIS IN THE BRITISH ISLES 211
GEOGRAPHICAL DISTRIBUTION

The Wild Service almost certainly entered Britain from the Continent after the last Ice Age,
although it was probably one of the later arrivals, and spread north and west along river valleys
(avoiding wetlands), or through the forests with which much of the country was covered. Its current
altitudinal limit in Britain is around 300 m and it has forked east and west at the southern end of the
Pennines and skirted the higher hills of the West Country and Wales. The northern advance, so far
as modern records show, continued to the southern Lake District in the west and, following the
Magnesian Limestone, to North Yorkshire in the east.

Deciding how best to present the geographical distribution of the Wild Service has not been easy.
Many prefer a national or Watsonian vice-county dot map method, but this is only useful for giving a
broad indication of the distribution across the country or in a vice-county. It also gives a potentially
misleading picture of a species that is occasionally found at quite high densities, but also occurs as
isolated plants in widely scattered locations. At first glance the species seems to be associated with
certain geological areas, but closer inspection reveals that this presents only a partial picture with
some baffling anomalies. The best explanation of its distribution in the wild is derived from an
analysis of its occurrence in relation to both solid and drift geology coupled with as much data as one
can obtain about socio-historical factors. The tree is widespread on the Weald Clay of Kent, Sussex
and Surrey for example, but almost absent from an area east of Tonbridge where the Weald Clay is
overlain by sand and gravel. Further east still it reaches one of its highest densities in an area where
its fruit were once in much demand as food (Hanbury 1770; Maynard 1925; Pratt 1854-57; Roper
1975) and where it is still widely known by a dialect name (Chequer Tree) and cultivated to a greater
extent than elsewhere. Its economic history, which has a considerable bearing on its present
distribution, is reflected in its vernacular name ‘Wild Service’. (Originally the species was known
simply as the Service — in many alternative spellings — and Wild Services were simply those growing
wild rather than in gardens or orchards. The usage is the same as in terms like ‘wild blackberries’ or
‘wild boar’. When Sorbus domestica L. was introduced in the 17th century it was called ‘sorbus
legitima’ and this was translated as ‘true service’ to indicate that it was the Sorbus to which classical
authors referred.)

I have based my observations largely on the ten mile (1:625000) solid and drift geology maps
published by the British Geological Survey. The maps give a broad indication of the type of
substrate and in some instances provide a very useful picture. However, drawing too many
inferences from cartography at this scale is unwise and as accurate a picture as possible of local
conditions should be obtained before reaching any firm conclusions on a particular site. The Wild
Service, although favouring particular habitats and areas, will survive almost anywhere in lowland
Britain, but this is quite a different matter from its being able to establish itself without any help
from man and to reproduce successfully.

Bearing all these provisos in mind I have illustrated the present distribution by considering the
more or less discrete areas in the British Isles where the species has been recently recorded (Figs 2
and 3) and its relative abundance within these areas. This distribution may reflect recording activity
to some extent, but generally there seem to be other reasonably satisfactory explanations for the
presence of the species and its density in a particular area. Equally interesting is its absence from
large areas that appear suitable and often lie adjacent to places where it is relatively abundant.

1. WEST CORNWALL

There are a few scattered records from the area west of Bodmin. All are confined to the Lower
Devonian and are often associated with the lower reaches of river valleys. Britain’s most westerly
record is from the Loe Pool south of Helston, although there is a planted tree at Castallack in
Penwith.

2. CAMEL VALLEY, CORNWALL

The area surrounding the Camel estuary and the Camel river and its tributaries in central northern
Cornwail had, in the recent past, a concentration of Wild Service trees of which a few remain
(Hamilton Davey 1909; Thurston & Vigurs 1922). The area is rather complex geologically, but most
locations are on the Upper and Middle Devonian formations.

212 P. ROPER

Ficure 2. Distribution by 10-km squares of the Wild Service tree in England and Wales. This map, though useful,
presents a potentially misleading picture due to the fact that some 10-km squares may contain only a single tree
whilst others may have hundreds often growing together in suitable areas (cf. Fig. 3). @ 1950 onwards, O before
1950, X introductions. Planted trees in Irish gardens have not been included as there are few precise locations.

3. LOWER TAMAR VALLEY

The tree is widespread along the river Tamar that divides Devon from Cornwall in the south. In
some places it has spread away from the river, in particular on to the soils derived from the Culm
Measures of the Upper Carboniferous in south east Cornwall. Old records from the south east of
Plymouth can be considered part of this grouping.

4. CENTRAL & NORTH DEVON AND NORTH EAST CORNWALL .
The Culm soils that stretch right across this area have a thin scatter of Wild Services, often in hedges
or on roadsides, although they are absent from a large tract of country west of Tiverton.

The association with the Culm Measures is marked and there are very few records from the

SORBUS TORMINALIS IN THE BRITISH ISLES 213

Ficure 3. Relative density of distribution of the Wild Service tree. The map shows each of the areas covered in
the text with isolated records as single dots. As well as these areas it is important to note that there are many
apparently suitable places where the Wild Service has not been recorded. Boundaries are those of Watsonian
vice-counties. The numbers refer to the sections in the text. Hl Areas were the species is generally not infrequent
and, in places, reaches its highest concentrations. Z Well-established but at lower concentrations. “| Vulnerable
or, in a few cases, apparently recently extinct (widely scattered, mainly as isolated trees with data often based on
a high proportion of old records).

extensive formations of granite, Middle Devonian or New Red Sandstone that lie adjacent. The
explanation is probably that the soils of the Culm are mainly clayey and water retentive, whereas the
others drain more freely.

5. SOUTH EAST DARTMOOR
This is perhaps an extension of 4 above. There is a concentration of locations to the east and south

214 P. ROPER

of Dartmoor on Culm Measures and other Carboniferous formations in this still well-wooded area.
There is also an old record from Chagford, the only one in Britain from granite, and a population on
Devonian limestone near Newton Abbot.

6. SOUTH EAST DEVON
I have only two old records from this area at Budleigh Salterton and Aylesbeare.

7. BRENDON HILLS
There is only one record from the southern section of the Upper Devonian formation that runs from
the Quantocks across Exmoor to Ilfracombe.

8. VALE OF TAUNTON

The species is scattered throughout the Vale and parts of the Quantocks on the Lower Lias, the

Devonian and the Keuper Marl, the latter a formation it favours, northwards through the Bristol

area, across Worcestershire and Warwickshire to the north Midlands as far as Nottinghamshire.
In Somerset there are some areas to the south and east that would appear suitable for the species,

but from which there are no records.

9. SOMERSET/DORSET JURASSIC
There are a few trees in woods on the Oolite east of Crewkerne. An old record from Puncknowle
near the coast to the south is from a geologically similar area.

On the whole, S. torminalis seems to avoid the Jurassic Cotswold limestone (Oolite), but it does
occur in small areas on this formation from Dorset to south Lincolnshire.

10. NORTH DORSET CLAYS
The tree occurs very sparingly in woods on the Oxford and Kimmeridge Clays in this part of the
county which is similar geologically to the Braydon Forest area west of Swindon in Wiltshire (see 35
below).

There is an old record (1799) from Broadley Wood west of Blandford Forum (Mansell-Pleydell
1895). This could either have been on chalk or, perhaps more probably, clay with flints.

11. LONDON CLAY IN DORSET
The Wild Service has a clear association with the London Clay almost everywhere this formation
occurs and Dorset is no exception. There is substantial documentary evidence indicating that S.
torminalis was widespread in the Lytchett Matravers/Wimbourne Minister area well into this
century (Mansell-Pleydell 1895; Marchant 1937). I have had no recent records, but a careful search
might well reveal that the tree still persists in this area.

Archaeologists have identified Wild Service charcoal from Maiden Castle, the Iron Age hill fort
on the chalk south west of Dorchester (Salisbury & Jane 1940), but this might not have been of local
provenance.

12. TEST VALLEY, HAMPSHIRE

There are scattered records from the valleys running from Redlynch and Alderbury south east of
Salisbury eastwards to the River Test and in the Test Valley itself south of Romsey. These are
largely associated with the London Clay and earlier Tertiary formations, although one site is on
chalk south west of Broughton and in the area south of Romsey the London Clay is much overlain
with alluvium, gravel and river terrace deposits.

13. NEW FOREST

The distribution of the Wild Service in the New Forest is one of the most difficult to interpret and it
perhaps reflects the way in which the forest has been managed as well as other factors. The trees are
concentrated in three areas:—

a. The south east of the Forest between the Beaulieu River and Lymington. This is partly on the
Tertiary period Hampstead Beds and Bembridge Marls and partly on the overlying glacial gravels
and sands from the more recent Quaternary period.

b. An area between Totton and Beaulieu Heath on the eastern fringes of the Forest. This is largely

SORBUS TORMINALIS IN THE BRITISH ISLES 215

on the complex series of Tertiary gravels and clays comprising the Barton, Bracklesham and
Bagshot Beds that overlies the London Clay.

c. The area around Cadnam, again on the eastern outskirts of the Forest. Geologically this is similar
to b above.

The tree occurs on the clay soils over the Hampstead Beds and Bembridge Marls in the northern
part of the Isle of Wight and here it clearly flourishes on this type of terrain. The Barton,
Bracklesham and Bagshot Beds, however, cover not only large areas of the New Forest, but extend
westwards nearly to Dorchester and over many square kilometres where Surrey, Hampshire and
Berkshire meet. The Wild Service is absent, or very rare, in all these places although there are
several records from the formation to the east of Southampton Water and in the Pamber Forest area
around Silchester on the Hants./Berks. border.

14. SOUTH EAST HAMPSHIRE

There is a well-defined grouping of Wild Service in south east Hampshire almost entirely on soils
associated with the London Clay, the Barton, Bracklesham and Bagshot Beds and overlying glacial
gravels. Some of these trees are probably relicts from the Forest of Bere.

15. ISLE OF WIGHT

The tree has been recorded, at one time or another, from many woods in the northern part of the
Isle of Wight and it still grows there, particularly beside estuaries and where there are low cliffs with
landslips. All these locations are on the clay soils derived from the Tertiary Oligocene as in the south
eastern New Forest.

There is one record from the Lower Greensand north west of Sandown in an area of complex
post-glacial geology. It was quite close to here at Nunwell that Sir John Oglander, in the early 17th
century, “planted above a hundred young elms and ashes, some chestnuts and service berries in the
grove of my house”’ (Bamford 1936).

The fruit also used to be sold, mainly to children in Ryde (Bromfield 1856), and it would therefore
seem that the distribution has been modified by human activity on the island, although it is
undoubtedly a native plant.

16. WEST ITCHENOR, WEST SUSSEX

The tree has long been known from a location by Chichester Harbour where glacial gravels overlie
London Clay. This is some 16 km from the nearest locations in Hampshire and 32 km from the
substantial populations in the Weald.

17. WESTERN WEALD
The Wild Service has one of its strongholds in the Weald of Kent, Sussex and Surrey and there are
concentrations on both the eastern and the western sides. In the west it is largely confined to the
Weald Clay but spills over on to the Hastings Beds between Haywards Heath and Horsham and
here and there on to the Greensand. There are two records from the chalk at Findon and near
Amberley, but the status of these needs further investigation.

In the west, records stop abruptly at Petworth, although the Weald Clay continues for some 24 km
along the Rother Valley.

The tree is also relatively scarce on the Weald Clay between Pevensey and Chailey in Sussex and
between Yalding and Hildenborough in Kent. In the latter instance it is missing from the extensive

. river terrace gravels associated with the Medway that overlie the Weald Clay.

18. EASTERN WEALD

There is a strong concentration of Wild Services in the area north west of the Romney Marsh
between Ashford in Kent and Robertsbridge in East Sussex. They grow on Weald Clay and the
Wadhurst and Guestling Clays of the Hastings Beds, but are almost entirely absent from the
sandstones and other light soils. I have searched many suitable looking woods on Gault Clay and on
the clayey soils overlying the Purbeck Beds in areas close to strong Wild Service populations without
locating a single tree, although the Midland Hawthorn, Crataegus laevigata (Poiret) DC., another
indicator of ancient woodland that often grows alongside Wild Service, is abundant in these places.

216 P. ROPER

This Wealden group of records spreads, at a thinner density, southwards and westwards almost to
the coast at Hastings and Fairlight and to the Ashdown Forest area.

It is in this part of the Weald that the species is still widely known as the Chequer Tree, as it is in
some other parts of south east England, and it has entered into the life of the rural community to a
greater extent than anywhere else in Britain. There are several farms, houses and one wood named
after the species and there is a clear association with the many Chequers Inns in the area (Maynard
1925; Roper 1975). The fruit, known as Chequers, were widely eaten within living memory (Pratt
1854-57 and D. Baird et al., pers. comm., 1975) and were probably used to make a cider-like drink,
hence their association with public houses. Because of this the tree is frequently planted in the area
and has clearly been given preferential treatment that it has seldom received elsewhere in woods,
hedges and on roadsides. The social history of the tree here and elsewhere in Britain and Europe has
been covered by Roper (1975, 1987).

19. BLEAN, KENT

The species is plentiful in woods on the London Clay in the Blean area of north Kent. There are also
a few records from the Sittingbourne and Rochester area of the London Clay and its underlying
Tertiary beds further to the west, and older references (Hanbury & Marshall 1899) indicate that the
trees were probably more widespread in the north Kentish clays.

20. SOUTHFLEET, NORTH WEST KENT

An area where the species occurs covering several square kilometres has been on record since the
16th century — “in Kent it groweth in great aboundance, especially about Southfleet and
Gravesend” (Gerarde 1597) — and it is still flourishing. Some trees grow on London Clay or
associated Tertiary gravels, but others appear to be on clay with flints or chalk. Careful investigation
of individual sites and the history of land management would be needed to establish why the trees
are found here but are absent from similar places nearby.

21. SURREY COMMONS
There are scattered records from the London Clay commons of central and west Surrey from
Wimbledon to just north of the Hog’s Back. The species closely follows the narrow belt of clay
between the sands and gravels to the west and north and the chalk to the south.

22. LONDON AND ESSEX :
This is a larger geographical area than many considered here, but records spread in a continuum,
with some local concentrations, from the Essex coast to the borders of Buckinghamshire and
Berkshire in the west and Kent and Surrey in the south. Virtually all are associated with London or
Boulder Clays and the gravelly soils that overlie them. Many represent survivors, or descendants, of
trees from the large forests of Essex and Middlesex which encompassed the smaller forests of
Epping, Hatfield and others. |

The earliest record I have from anywhere in Britain (other than the report of Iron Age Wild
Service charcoal from Maiden Castle, Dorset) is from Havering Park, Essex (enclosed as a royal
park in the Middle Ages). In 1260 two Wild Service trees (described in the text as “‘alyeras”’) were
brought from the park to the Tower of London for the manufacture and repair of crossbows, a
purpose for which the wood was renowned. Owing to a mistranslation of the Latin text these trees
have been considered by some authors to be hawthorns, but this is incorrect.

Today there are concentrations of records from the Basildon area and to the south east of
Chelmsford and these are associated with the Tertiary and Quaternary gravels overlying the
London Clay. It has also been noted that trees here tend to be associated with places where the
water supply is increased by the effect of differential percolation through the bedded strata.

In Epping Forest the trees grow mainly on London Clay, but north eastwards from Chipping
Ongar to the Ipswich area records are scattered across the Boulder Clay, or its associated gravels. It
is curious that, although the Boulder Clay sheet continues much further to the north, Wild Service
records become very scattered indeed beyond the Suffolk/Essex border. I am inclined to think this is
due to land and forest management over the last several thousand years rather than natural factors
and in earlier times the Wild Service could have been much more widespread in East Anglia.

To the west the tree is still remarkably well-distributed within the London Clay triangle of north

SORBUS TORMINALIS IN THE BRITISH ISLES 217

west London, often in urban situations such as on Hampstead Heath and in Ken Wood (Gilmour
1972). There are also scattered records from south east London and an old record from Wimbledon
Common, again on London Clay.

If these London and Essex records are plotted on a Quaternary geology map, their absence from
hundreds of square kilometres of glacial river terrace deposits along the valleys of the Thames, the
Lea and the Wey is very marked. It is remarkable that there are sufficient trees left to give such a
clear indication of the ecology of the primaeval wildwood that once covered what is now one of the
world’s largest cities. It also highlights the Wild Service’s ability to survive in unfavourable
circumstances for long periods.

23. SOUTH EAST BUCKINGHAMSHIRE

In the well-wooded area around Burnham Beeches that lies in the rectangle between the M40 and
the M4, and the A412 and the Thames at Cookham, there are a number of records from the river
terrace deposits that the species seems to avoid elsewhere. Careful investigation of individual sites
might provide the reason as might the history of woodland management in the area.

24. SOUTH EAST HERTFORDSHIRE

To the south west of Hoddesdon and the east of Welwyn, with an outlier west of Hatfield, there is a
concentration of records in a county where the species is otherwise rare. Most are on Boulder Clay
or associated gravels, with a few on London Clay.

In the 18th century the Wild Service tree was well-known in Hertfordshire: Miller (1768) said ‘in
many parts of Hertfordshire there are large trees now growing’’. He may have been referring to the
area north of Barnet where the tree still occurs in some quantity or to south east Herts. (Harper
1981). 92 years earlier M. Cook was planting them for the Earl of Essex on his estates at Hadham
and Cassiobury. Apparently they fruited well on the Boulder Clay at Hadham, but at Cassiobury
(Cook 1676) “‘we have them on a sharp gravel, the Fruit naught, and the trees bear very badly”’.

_ 25. WINDSOR FOREST, BERKSHIRE

| There is a strong concentration of the trees in the London Clay area of Windsor Forest, Berkshire.

| The trees are closely associated with the clay, avoiding the river terrace deposits to the north and the
gravelly or sandy Tertiary beds to the south.

26. SOUTHERN CHILTERNS

_ From the Reading area north across the Chilterns there are scattered records of the species over a 12
) X 18 km. well-wooded, geologically complex area typified by clay with flints, river terrace deposits
» and other gravels cut by rivers down to the underlying chalk or, on the southern fringe, the London
‘Clay. As with the trees in south east Buckinghamshire (see 23 above), a study of individual sites is
» needed to determine the conditions that favour the species in this area.

| 27. CENTRAL AND NORTHERN CHILTERNS

' There are very few records from the more elevated parts of the Chiltern range in Hertfordshire,
Buckinghamshire, Oxfordshire and Berkshire. As far as I can determine the trees are confined to
_ the clay with flints cappings on the hills.

28. PAMBER FOREST, HAMPSHIRE/BERKSHIRE BORDER

There is a group of records from the wooded area, formerly Pamber Forest, around Silchester on
the Hants./Berks. border. They are mainly on Quaternary sands and oravels that overlie the
London Clay.

__ The Tertiary formations running from the Kent, Essex and Suffolk coasts and with which the Wild
Service is so closely associated reach their westerly limit immediately beyond Newbury before
‘ See to the south of the chalk. There i is one record from the area west of Sine then a

218 P. ROPER

untypical and, in my view, do not constitute adequate evidence that the species once grew more
widely on chalk soils under British conditions.

30. SOUTH WILTSHIRE
The only modern record I know of from v.c. 8 is of a Wild Service in a hedge between Wilsford and
Beechingstoke. This is on the Gault Clay in the Avon Valley south east of Devizes. If the tree is
growing on Gault, it is the only current record of which I am aware from a formation which, in many
respects, seems well-suited to its requirements.

Aubrey (1685) said that “cervise-trees’’ grew at the foot of Heddington Hill and at Whitesheet (I
presume the one at Mere) and the trees here may have grown either on Gault Clay or on chalk.

31. WILTS./AVON BORDER
There is a concentration of records, mainly from the Greater Oolite and Cornbrash, but spilling
over on to the Oxford Clay along the Cotswold line from Castle Combe to Frome and roughly
associated with the old forest of Selwood. Most of the records are from the lower lying parts of the
River Avon catchment, but otherwise seem to have little in common. They lie in the counties of
Wiltshire, Avon and Somerset although they only extend over an area of some 32 X 12 km.
There are outliers to the east on an area of Kimmeridge Clay and the Lower Greensand near
Calne, and to the south on the Upper Lias near Evercreech.

32. WESTERN AVON
The Wild Service is widespread and sometimes fairly abundant from the Mendip Hills, through the
Bristol area and northwards along the Vale of Berkeley. It occurs on the Keuper Marl, the Upper
and Lower Lias and the Upper and Lower Carboniferous formations, in the latter instance
sometimes on the limestones that make up much of the Mendip Hills and the Avon Gorge. In this it
displays its two main habitat preferences: for open and often steep rocky areas and for nutrient rich,
heavy clays. :
Towards the north of the Vale of Berkeley the concentration of records increases in an area of |
complex Silurian and Cambrian geology. Similar areas occur across the River Severn to the north |
and these are more fully considered in 52 below.

33. FOREST OF DEAN
As Rackham (1986) has pointed out, the Forest of Dean was subjected to management regimes,
largely in the 19th century, that have altered the existing tree composition almost beyond

recognition. This is reflected in the virtual absence (very unusually for our ancient lowland forests)

of the Wild Service. There are only a few near Brean and more on the outskirts of the forest in the |
Cinderford area. These belong, however, to the group of records running northwards away from the
forest on the Silurian formations of the Malvern Fault.

Wild Services grow on the cliffs bordering the Severn Estuary at Chepstow and Lydney and in the |
Lower Wye Valley, but these again are not strictly part of the Forest of Dean.

36. LOWER WYE VALLEY
The Carboniferous Limestone of the Lower Wye Valley is renowned for botanical diversity and |

boasts a large number of Sorbus species. Wild Services are found in the open rocky woods and on |—
cliffs from the Goodrich area to Chepstow, though not on the stretch where the river flows through |

Devonian strata. This and Weston Big Wood near Weston-super-Mare are the only areas in Britain |
from which the hybrid (S. < vagensis Wilmott) between the Wild Service and the Whitebeam, |_
Sorbus aria (L.) Crantz, has been recorded. |

There are two Wild Service records close together from Llanfihangel Ystum Llywern in the valley |
of the River Trothy in Gwent. This site is only 10-12 km from the River Wye, but the tree is found |
almost nowhere else in the very extensive Lower Devonian countryside through which much of the | -

upper River Wye and its tributaries run, although it has its densest populations in some of the woods | |

immediately adjacent to the eastern boundary of this area and reappears beside the Wye on the -
Wenlock Series in Radnorshire (Powys) (Anon. 1976). i

SORBUS TORMINALIS IN THE BRITISH ISLES 219

35. FOREST OF BRAYDON, WILTSHIRE

Between Swindon and Malmesbury in north western Wiltshire there is a concentration of records
from the woodland remnants of the Forest of Braydon, which lie mainly on the Kimmeridge Clays of
the Upper Jurassic.

In general the species seems to be commoner on Oxford Clay and it is of note that the S50 km gap in
records between Braydon and Wychwood on the Oxon/Bucks. border is largely across an area of the
Thames Valley where the clay is overlain by river terrace deposits similar to those from which the
tree is absent in south west London and the Medway Valley, Kent.

36. SOUTH WALES

There are very few current records for South Wales and if the species has ever been commoner there
it could well have been largely restricted to the coast between Cardiff and the Gower and ‘the
immediate hinterland.

Many Sorbus species grow on cliffs close to the sea and their fruits fall on to the beach or into the
water, sometimes in considerable quantities. They can also be washed down rivers into estuaries and
must be able to travel for some distance by this means. From the beach to the cliffs is only a short
distance for a bird or fructivorous mammal and some coastal colonies may have originated through
seed being spread in this way rather than by overland routes.

Outside the Gower there are a few old records of probably planted trees in the Swansea area and
another at Wenvoe south west of Cardiff. If they were indeed planted they could, as in some other
places, have originated from wild trees in the area.

37. GOWER PENINSULA
There are several trees in the woods along the limestone cliffs on the southern side of the Gower
Peninsula growing in similar circumstances to those in the Wye Valley, the Avon Gorge and the
Arnside area of Westmorland.
Inland the Gower is almost covered in a sheet of Boulder Clay and there are no S. torminalis
records.

38. CARMARTHENSHIRE

V.c. 44, now part of Dyfed, has records from three scattered locations that imply that the tree was

once more widespread in lowland Wales. It is found at two sites in the heavily wooded Cwm Mawr

valley on cliffs of Pennant Sandstone capped by Boulder Clay into which suckers are spreading.
Further west there are five trees on a wooded estuarine Red Sandstone cliff of the Lower Devonian
(also capped with Boulder Clay) north of Laugharne.

Well inland at Poor Man’s Wood, near Llandovery, there are three trees in an oakwood that now
belongs to the Dyfed Wildlife Trust on clay derived from rocks of the Upper Ordovician.

39. RADNORSHIRE AND BRECKNOCKSHIRE, POWYS

The Wild Service grows on cliffs of the Wenlock Series of the Silurian bordering the Wye to the
south of Builth Wells and further upstream on similar cliffs by the River Ithon in v.c. 43, Rads. (now
part of Powys). It is also found on this formation in Brecs. (v.c. 42) and further to the north.
Further search will almost certainly reveal more locations in suitable lower lying parts of Wales,
both inland and on the coast. In many cases plants have been kept small due to grazing by sheep, but
several of these sites have now been fenced and the trees should now be able to grow to their full
size.
40. SOUTH PEMBROKESHIRE
The Wild Service was first recorded formally from Pembs. (v.c. 45) only in 1971. Since then it has
proved to be fairly widespread and even boasts a unique local name — maple cherry.
| The majority of trees grow in oakwoods on the low cliffs or banks bordering the extensive
estuarine system that runs out to sea at Milford Haven. The rock here is mostly Old Red Sandstone
/ of the Lower Devonian and these groups of trees are analogous to those associated with some of the
/south Cornish estuary systems.
Further coastal records occur on the Coal Measure sandstones on the western side of Carmarthen

eee ee

220 P. ROPER

Bay and there are a few inland records from this geologically very varied peninsula north to the
Newgale area (Anon. 1976).

Davis (1976) has observed that in this area the trees only fruit well after warm summers, |
confirming that it is a thermophilous species at the limit of its range in Britain. The same has been |
observed elsewhere in the west and the north in this country and elsewhere in Europe (Bisgen 1929;
Conwentz 1895; Termena 1972). In the south of England the tree usually fruits well, but will |
frequently miss one, and sometimes more seasons, another phenomenon well-known in trees that |
prefer a warmer and sunnier climate than our own.

41. TEIFI AND NEVERN, DYFED |
There are small Wild Service areas in woods adjacent to the lower reaches of the rivers Teifi and |
Nevern north of the Mynydd Preseli in northern Pembs. (v.c. 45) and southern Cards. (v.c. 46), now |
both in Dyfed. |

The soils here are derived from Upper Ordovician rocks of the same type as those at Poor Man’s /
Wood, Llandovery (see 38 above). .

42. ABERAERON, DYFED

There is a scatter of records from the low-lying area inland from the coast at Aberaeron in Cards.

(v.c. 46), mostly on steep, wooded river banks.
The soils here are derived from the Llandovery Series of the Silurian and cover much of western |

central Wales which is mainly free of boulder Clay and other drift. One record is from a hedge at |

Llangybi over 15 km from the coast and in a tributary valley of the River Teifi. The location is, |

however, still on the Silurian and falls within the grouping.

43. DOVEY VALLEY (CWM DYFI) |
There is one record of a plant at the top of a steep, rocky, wooded slope in a valley off the main |
Dovey Valley between Machynlleth and the estuary. The whole of this area, as with 41 above, ison |
the Llandovery Series of the Silurian. The site is just in v.c. 47, Monts. (now part of Powys), and |
quite different from those in the east of the vice-county (see 56 below) |

44. BARMOUTH
The most northerly of the west central Wales populations are the few trees that grow in steep, rocky
woods on Cambrian formations close to the Mawddach estuary. |

45. NORTH EAST SUFFOLK/SOUTH NORFOLK

There are a few records from the area to the south west of Southwold of trees growing on soils
derived from the underlying Boulder Clay. One correspondent sent me leaves from the Southwold
area and said he knew at least five sites for the species in the neighbourhood.

R. Mabey (pers. comm., 1976), has noted that members of the Yoxford Women’s Institute knew |
that the tree grew wild in their district. This indicates that there may be more sites for botanists to
discover in the area. |

There is one outlier of this group of records well inland near Mendlesham, another near |
Hempnall south of Norwich (Withering (1818) said the trees grew at Bath Hills near Bungay not far |
from here) and a garden relict north east of Bury St Edmunds. The Wild Service has been quite
widely planted in the past for ornamental and utilitarian purposes. The source of such plants is often
local and older garden trees may indicate a wild population nearby.

Most of the woodland cover in East Anglia was removed long ago but a map in Rackham (1986)
shows that this north east Suffolk/south Norfolk Wild Service area is broadly congruent with an area
where villages still had their own woods at the time of the Domesday Survey in the 11th century.
This area stretched north to the Foxley district in Norfolk (see 46 below).

a ee ne

|
H
;
&

46. CENTRAL NORTH NORFOLK

There are several Wild Services in Foxley Wood on the Boulder Clay sheet near Themeithorpe (E. |
Norfolk, v.c. 27) and a recent record from a wood at Mileham (W. Norfolk, v.c. 28), both to the |
north east of Dereham. Older records from the neighbourhood indicate that these are survivors | ~
from an earlier, more widespread woodland population. i

SORBUS TORMINALIS IN THE BRITISH ISLES 221

47. SOUTH MIDLAND LOWER GREENSAND

There are five Wild Service locations along the Lower Greensand ridge that runs from south western
Cambridgeshire to Leighton Buzzard in Bedfordshire. Although the correlation with the solid
geology seems too close to be accidental, most trees are in fact on Boulder Clay and elsewhere the
tree does not occur on soils derived from the Lower Greensand. Examination of the 1:50000
Ordnance Survey map shows that this ridge is more heavily wooded than the surrounding plain,
perhaps because the countryside was less easily cultivated due to its hilliness. The survival of the
Wild Service is almost certainly due to this rather than to the underlying solid geology.

48. BERNWOOD (BUCKINGHAM/OXFORDSHIRE BORDER)

There is a concentration of records of trees over the narrow, 16 km stretch of countryside from the

Quainton, Bucks. area to just south of Oxford and associated with the ancient Forest of Bernwood.
None of this area is covered in glacial drift and most of the sites are in woods on Oxford Clay, with

a few associated with the Greensand/Portland Beds complexes of the Kimmeridge Clay.

49. WYCHWOOD, OXFORDSHIRE
Before deafforestation, which followed enclosure in 1857, Wild Service berries from the Forest of
Wychwood used to be sold in Burford market and probably elsewhere locally.

The tree continues to be found at low concentrations in some of the few remaining woodlands,
nearly all of which are on soils deriving from the rather complex Oolite and Upper and Middle Lias,
that typify the Cotswold belt. The tree seems not to be found on the adjacent Lower Lias and is
scarce on this formation everywhere in the south and west and virtually absent from it in the north
and east. On a site near Kineton in Warwickshire on the Lower Lias some 32 km north of
Wychwood, the tree appears to be on the overlying Boulder Clay.

Scattered records occur westwards towards Cheltenham and the Vale of Evesham and there were
once, no doubt, trees in the woodlands throughout the whole of this area.

50. CIRENCESTER/STROUD

There is a somewhat isolated group of records of trees in Cirencester Park some of which are
planted, and records from the Tetbury and Painswick areas in the Cotswolds. These locations are
not dissimilar to those cited in area 48 above and again one can surmise that the tree was once more
widespread locally. The fact that there is a unique Cotswold name for the species — ‘lizzory’ or
‘lezzory’, clearly derived from ‘alizier’, one of the Old French words for the species (Boulger 1908) —
lends support to this hypothesis. |

51. VALE OF SEVERN

There are records of the tree, mainly from the Keuper Marl but also on the Lower Lias near
Pershore, from Gloucester to Stourport. Sometimes the tree grows on cliffs by the river, but more
often in woods and sometimes hedgerows nearby.

52. MALVERN

The Malvern and Suckley Hills and their southward extension to the outskirts of the Forest of Dean
are one of the strongholds of the Wild Service in Britain and are clearly associated with the complex
Silurian strata of the Ludlow, Wenlock and Llandovery Series that go to make up these hills. On the
spur of this formation that runs north west towards Hereford the Wild Service reaches one of its
highest densities in Britain with up to 125 trees per hectare in Haugh Wood near Mordiford. Stoke
Edith Wood, also in this area, is the type location and the source of the only British records of the
British Red Data Book moth, Stigmella torminalis (Wood), whose larvae, like those of several other
moths, mine the leaves of the tree (Emmet 1976; Shirt 1987).

53. WYRE FOREST

The Wild Service is not uncommon in the Wyre Forest and neighbouring parts of Worcestershire,
Shropshire and Staffordshire growing mainly on soils derived from the Westphalian Series of the
Upper Carboniferous. In the forest itself it is widespread, but occurs particularly along the central
Dowles Brook. While there are probably more soil nutrients here, it is possible that the tree was

222 P. ROPER

encouraged by the millers who had premises along the Dowles since its wood was among the best for
making cogs for mill machinery (Du Breuil 1854; Hanbury 1770; Hickin 1971).

54. UPPER SEVERN VALLEY :
The tree is found north of the Wyre Forest along the Severn Valley showing a marked preference for |
the Westphalian Series and largely avoiding the Lower Old Red Sandstone to the west as it does —
everywhere in western central England and eastern Wales. It is not found on the river terrace |
deposits, a distinctive feature of its distribution everywhere. Records become far scarcer north of |
the limit of glaciation, though there are a few from Boulder Clay-free areas east and north of
Bridgnorth.

55. WENLOCK EDGE AND THE WELSH MARCHES

As with the Malverns (52 above), the Wild Service has been recorded from much of the Wenlock
Edge and associated Silurian areas, with a concentration to the north around Ironbridge. There are
scattered records to the south west as far as the northern tip of Herefordshire and the Welsh border
at Knighton. There is one site near Pontesbury to the south west of Shrewsbury and two in the Welsh
hinterland, one on the Severn at Abermule and the other on the Afon Banwy (a tertiary tributary of
the Severn) south of Meifod. Both these are in Monts. (v.c. 47).

Unlike the more southerly areas of this type, the rocks of the Wenlock and other series of the
Silurian are overlain by Boulder Clay and glacial sand and gravel and all the S. torminalis records
from this area are on, or very close to, drift deposits. Sinker et al. (1985) say that in the Shropshire
region the tree is “‘a characteristic member of the ancient broad-leaved woodland community
together with holly, yew and small-leaved lime on sandstones and other freely drained acid rocks as
well as on limestone’.

56. WEST SHROPSHIRE LIMESTONE

There is one old and one more recent record from the Carboniferous Limestone west of Oswestry.
This area is well known for its apomictic Sorbus species that are so often confined to open limestone
areas and in this case the Wild Service is responding in the same way. The nature of the terrain
reduces competition for light from more vigorous tree species and allows the Sorbus spp. to reach
maturity whereas on richer soils in the same neighbourhood they would be shaded out.

57. UPPER DEE VALLEY | q
There is a small group of records from the Upper Dee Valley on the Shropshire/Clwyd border
between the point where the river debouches from the Vale of Llangollen downstream to Bangor-is-
y-Coed.

Unlike the lower part of the valley, this area is still heavily wooded. The sae is varied with
much overlying clay and gravel drift.

58. NORTH WEST MIDLAND PLAIN

There is a very thin scatter of records, many of them old, across the low-lying agricultural plain of
north-east Shropshire, west Staffordshire and Cheshire. Much of the soil is heavy and derived from
the thick sheet of Boulder Clay left by the last glaciation, while elsewhere there are glacial sands and
gravels. The underlying Permian and Keuper Marl appear here and there.

There seem to have been concentrations of the tree to the north and west of Shrewsbury, to the
south of the Potteries and in the Delamere Forest area, although these are based on very few old
records. Elsewhere there are, or were, isolated trees.

It is possible that the species was once widespread in this area, although perhaps always rare. If
one looks, for example, at the records from the sticky soils of the Culm Measures in central and
north Devon, one sees a similar, though slightly denser, pattern of scattered records.

59. VALE OF CLWYD

There is a surprising concentration of records in the Vale of Clwyd, mainly in the area around
Denbigh some way inland, but with an outlying station south of Prestatyn. Some of these records are
associated with Carboniferous Limestone, but the area is extensively covered with Boulder Clay
and the trees are growing in woods and hedges rather than on cliffs.

SORBUS TORMINALIS IN THE BRITISH ISLES 223

60. GWYNEDD COAST
One tree has long been known from the Craig y Gigfran on Garth Point, an area of Cambrian and
Ordovician rock on the coast of the Menai Strait in Bangor.

I also have an unconfirmed record of a tree alongside the Conwy Estuary on the Benarth Estate to
the south east of Conwy itself. My informant thought it might have been planted, but the geology
and the general situation are of the type favoured by the species and the area warrants further
investigation.

61. ANGLESEY
There is a long history of the species (possibly planted) at Trefarthen, near Brynsiencyn close to the
_ Menai Strait in the south-west of the island (Davies 1813) although it has not been seen recently as
far as I know. Non-native trees, including Sorbus spp., have been extensively planted in some parts
of Anglesey (Sell 1989), but there is no overriding reason why the Wild Service should not be native
here.

The underlying rocks in this part of Anglesey are metamorphic Horneblende Schists, but almost
the whole island is overlain by a mantle of Boulder Clay.

62. LIVERPOOL

There is a record of a Wild Service on the rocks at Knot’s Hole, Liverpool “‘in a situation quite
exposed to the salt water, and where it must occasionally be washed by the spray of the sea”
(Withering 1818). There have been no recent reports. In many ways this location is analogous to
those were the species grows on various parts of the coast of Wales, or further north around
Morecambe Bay. The rocks at Knot’s Hole are New Red Sandstone.

63. FOREST OF ARDEN

There is a strong concentration of records in the area south of Birmingham that is roughly congruent
with the ancient Forest of Arden. The trees are found in woods and hedges, almost entirely on
Keuper Marl.

64. NORTH EAST MIDLAND PLAIN AND FOREST OF CHARNWOOD
North of the Arden area the tree becomes very much scarcer. Its chances of survival Have been
reduced by urban development and agriculture and, as is the case in other areas, it does not grow as
successfully on the extensive Boulder Clay and drift as on other soils. As in the south Midlands and
places further north, it is absent from the very extensive Lower Lias.

There is a small concentration of trees on the Westphalian Series of the Carboniferous east of
Sutton Coldfield and another in the northern part of Charnwood Forest. Elsewhere the records,
mainly from Keuper Marl, are very thinly scattered and mostly date back some considerable time.

65. STAFFORDSHIRE AND SOUTH DERBYSHIRE

There are scattered records from the Needwood Forest and Bagot’s Park area of Staffordshire with
an isolated hedgerow tree from a neighbouring part of Derbyshire. Most are on Boulder Clay,
although the underlying stratum is Keuper Marl throughout.

66. NORTHAMPTONSHIRE AND ROCKINGHAM FOREST

The tree has long been known in northern Northamptonshire and north western Cambridgeshire in
the old Rockingham Forest area. The early 19th century poet John Clare, who came from this
neighbourhood, was familiar with the tree and called it by its local name of Surrey as well as Service
Tree in his writing (Clare 1832). J. R. Gilson (pers. comm., 1977) has reported that parish bounds in
the Rockingham Forest area used to be beaten with branches of the tree, and the branches also used
to be carried at the head of village processions (Grindon 1883). All this indicates that the tree has
long been familiar in the countryside here.

It now grows almost exclusively in woods, and occasionally hedges, on soils derived from the
Oolitic limestone formations that are so characteristic of this area and generally avoids places where
there is a Boulder Clay covering. This is in contrast to the examples immediately to the north where
the reverse is true and Boulder Clay seems to be preferred.

224 P. ROPER

67. RUTLAND AND KESTEVEN

There are still widespread records from this once heavily wooded area, although many trees were
destroyed during the replacement of deciduous woods by conifer plantations after the Second
World War.

The tree is particularly concentrated on the Boulder Clay spur followed by the Great North Road
from a few km north of Stamford to Grantham and records extend westwards on a similar substrate
to Dunsby in Lincolnshire. To the south east there are scattered records across the Vale of Catmose
area and from beyond Oakham.

Both in Rockingham Forest and this area, pheasant shooting has been popular, particularly on
the large estates, since the last century. Many Wild Service trees grow in the coverts where the birds
are raised or roost and the birds are known to be very fond of its fruit (Conwentz 1895). This may, to
some extent, have helped the tree survive as gamekeepers undoubtedly know what their birds like.
In some cases the species could have been deliberately introduced, particularly into the smaller
woods that are wholly artificial and that were established with game and foxes in mind.

68. HUNTINGDONSHIRE

There is a group of records from central Huntingdonshire and one to the south near St Neots. These
are on Boulder Clay and the underlying Oxford Clay. In the case of the latter, it is remarkable how
the cluster of records to the south-east of Sawtry (which includes Monks Wood) almost exactly
matches a small area where the Oxford Clay is not covered by glacial drift.

69. BARDNEY FOREST, LINCOLNSHIRE

Although much of the countryside to the east of Lincoln is now conifer plantation, there are still
some remnants of the ancient wildwood that persisted here until the middle of the 19th century and
the area is noted for its rich wildlife.

Records of the Wild Service are numerous and are concentrated in an area only some 14 X 20 km
in size. They grow almost entirely on the Boulder Clay, although a few records seem to be from the
alluvial soils in the Witham Valley.

The topography and geology of much of central England is not dissimilar to that in this area and
this strong population of the Wild Service may give some indication of the density it once reached in
many other places.

70. NORTH LINCOLNSHIRE
There is one record of a tree in a hedge on the chalk in north Lincolnshire far from any other Wild
Service trees. It is almost certainly introduced.

71. EAST NOTTINGHAMSHIRE

The Wild Service is well-distributed, mainly in woods, throughout the Keuper Marl of eastern |

Nottinghamshire with a couple of records, clearly belonging to this group, on the other side of the
River Trent at Gainsborough. This preference for the Keuper Marl in an area largely free of drift is
striking and, apart from one record on the New Red Sandstone near Thoresby, the species remains
on its preferred soil throughout the Sherwood Forest area.

The Keuper Mari continues north through Yorkshire to Teesmouth and the sudden cessation of
S. torminalis records is clearly related to the fact that from north Nottinghamshire the formation
vanishes under a great sheet of drift.

72. PEAK DISTRICT

There is one record from Wild Cat Tor on the Carboniferous Limestone at Matlock (Willmot 1975).
Other Sorbus spp. grow well on the limestone eyebrows and other habitats created by the geology of
this area and in this instance the Wild Service is responding to the environment like some of the
related Sorbus spp., especially the apomicts.

73. MAGNESIAN LIMESTONE .

The Wild Service appears in woods on Magnesian Limestone from its southern extremity and
follows it up the Nottinghamshire/Derbyshire border northwards to the Chadwick-le-Street area in
South Yorkshire. There are now some 20 known locations along this limestone belt and its

SORBUS TORMINALIS IN THE BRITISH ISLES 225

associated mudstones despite the fact that the formation is never more than about 9 km wide ©
(Willmot 1975). It is also largely free of drift and the species is clearly responding, as many other
plants do on this formation, to the soil conditions deriving from the solid geology.

Beyond Chadwick-le-Street it reappears, after a gap of over 60 km, on a Magnesian Limestone
cliff near Fountains Abbey in Mid-W Yorks. (v.c. 64). It seems perfectly natural on this site, which
is the most northerly so far discovered on this side of the country, but one should always be wary of
the status of plants with some economic value growing near ancient monasteries.

74. DERBYSHIRE COAL MEASURES

In 1974 four bushes of Wild Service were found in a hedge bordering a green lane at Staveley,
Derbyshire on Coal Measure shales only 8 or 9 km from plants on the Magnesian Limestone
(Willmot 1975). The species occurs on this formation in one place in Pembrokeshire, but otherwise
nowhere else, although it is very widespread.

75. MORECAMBE BAY AND THE SOUTHERN LAKES

The Wild Service occurs on Carboniferous Limestone rocks and in lowland woods with glacial drift
derived soils in a number of places around Morecambe Bay (Piggott 1973/74). In the Arnside area
many of these woods are known to be ancient and were part of the Chase of Harneshed and the deer
park created there in the 16th century. The wildlife and social history of the other localities where
the Wild Service is found also indicate that their vegetation is of natural origin.

Often in these steep rocky areas the Wild Service responds like other members of its genus,
particularly the apomictic species (on Jack Scout it grows with Sorbus aucuparia L., S. rupicola
(Syme) Hedlund and S. lancastriensis E. F. Warb.) but, in Grubbins Wood and elsewhere, it grows
in similar situations to those in the more heavily wooded areas of the south.

There are good reasons to believe that old records from Plumpton near Ulverston, Brigsteer and
Levens Park are authentic and new sites in this area may remain to be found. As recently as 1987 a
long-established Wild Service was found on a low cliff on the shore of Lake Windermere, a place
one would have thought had been quite well-worked botanically. This is the most northerly site in
Britain where the species can be accepted as native. There are a few old records from further north
on the Cumbrian limestones, but these trees are almost certainly planted.

From time to time one hears of ‘service trees’ in north-east England and southern Scotland.
Sometimes these are simply misidentifications, but the Swedish Whitebeam, Sorbus intermedia
(Ehrh.) Pers., is very widely known as the Service Tree in this part of the country and this is often
the species meant. The Wild Service has been planted at several places in Ireland (Forrest 1985).

CONCLUSION

The Wild Service tree is, as other authors have established (Peterken 1974), a useful species for
indicating primary woodland and ancient hedgerows, as well as some other types of relatively
undisturbed habitat, and it is more likely to be found in an area that is still, or was once, part of an
ancient forest, but its presence on any particular site does require interpretation. Apart from
biological and ecological factors and the history of land and forest management in the British Isles,
the tree has economic, aesthetic and other qualities that have favoured its survival in some districts
to a greater extent than would be the case with many other species.

While the present survey has been as comprehensive as possible, it is clear that the Wild Service
remains under-recorded. Although it grows to a large size — I know one tree over 18 m tall and witha
spread of 21 m, while trees of between 22 and 24 m have been reported from several parts of Britain
and of 25 m from the Crimea and the Caucasus (Elwes & Henry 1906; Howard 1947; Conwentz
1895) — it is easily overlooked, and there are many areas that have not been thoroughly searched.
Despite this, the limits of its range in the British Isles are now reasonably well defined: it is
essentially a lowland tree of England and Wales and has not been reported in the wild from
Scotland, Ireland, the Isle of Man or the Channel Islands.

Its range is unlikely ever to have extended to Scotland. In the Atlas Mountains, where the Wild
Service and the Whitebeam, Sorbus aria (L.) Crantz, grow (Jahandiez & Maire 1931-34; Quezel &
Santa 1962), the Whitebeam extends to higher altitudes and this is an analogue of the situation in
Britain where the Whitebeam is found further north in a cooler climate than the Wild Service. The

226 P. ROPER

Whitebeam and Sorbus devoniensis E. Warb. are both found in Ireland (Clapham ef al. 1962) and
the latter species has recently been reported from the Isle of Man (Proctor et al. 1989; Sell 1989). If
both these species could cross the Irish Sea, there would seem no reason why other Sorbus spp.
could not do the same, so it is worth continuing to look out for the Wild Service in these places.

HABITAT PREFERENCES

The Wild Service in Britain and elsewhere grows in several distinct and apparently dissimilar
habitats, although an understanding of the requirements of the species goes some way to providing
an explanation of its distribution. These habitats can be divided into four broad categories:—

1. Nutrient-rich soils, clays in particular (Brown 1894). The species is more abundant and grows
better on some clays than others. It does well on Weald and Wadhurst Clay in the south-east, and on
London Clay and Oxford Clay (in the latter instance especially where there are, or were, ancient
forests). It is widespread on the Culm Measures in the West Country and the Keuper Marl from
Somerset to Nottinghamshire. It is less common on Boulder Clay, but grows well where it occurs,
and is virtually absent from Gault. One possible explanation is that it grows better on clays with a
lower calcareous mineral content, although woodland management over the centuries is a more
likely factor.

Woodland soils overlying clays often do not share the characteristics of the underlying stratum:
they are usually lighter and more acid and may be modified by downwash or other local conditions.
While the Wild Service can send its roots below the surface soil layer, the natural distribution (as is
the case with any forest tree) is governed by the conditions in which it has to develop as a young
plant and the superficial soil qualities are therefore an important factor.

2. Areas of hilly or undulating gravel terrace and similar formations deposited in the glaciations of
the Tertiary or Quaternary periods. There is some evidence that its presence on these formations is
associated with bands where the water content is higher due to the slope and the layering of the
material (Kozlowski 1962). Such gravels are often accompanied by seams of clay which may
improve the nutrient content locally.

Those who cultivated the tree in the past noticed that they did not fruit as well on freely-draining
gravelly soils as on clay (Cook 1676). :

3. Coastal and inland cliffs, rocky hillsides, landslips and similar open or disturbed habitats.

4. Large river valleys, especially those containing cliffs and rocky slopes.

The Wild Service is capable of becoming a large forest tree reaching the canopy layer, and its ~ |

absence from lighter lowland soils may be due to the fact that it competes poorly with oak and other
trees (Longman & Coutts 1974). If it is overshaded by trees that grow more rapidly, the Wild Service
can survive, but will not flower and fruit. Away from nutrient-rich soils it depends on open habitats
where there is less competition and where it can flower and fruit more successfully. In these places
chances of seed survival are also higher due to their becoming lodged in crevices and similar
positions where predators cannot easily reach them. The young plants are also less likely to be
damaged by browsing or grazing due to their inaccessibility.

The present pattern of distribution indicates that the Wild Service was probably widespread, if
seldom abundant, on the stiffer soils in the forests that covered much of the lowlands of England and
Wales in the past. In many places it has been severely reduced, or eliminated, by human activity and
this has been compounded by its low rate of reproduction from seed compared with many other
native trees. There are two main reasons for the latter: climatic and ecological. Gabrielian (1961)
contends that the species evolved in dry, open woodlands and there is ample evidence to show that
climatic conditions affect the tree’s ability to flower and produce viable fruit. Warm summers
promote fruit formation (Davies 1976; Biisgen 1929) and increase the number of seeds per fruit. In
the warmer, drier parts of Britain many trees often fruit only every other year and in marginal
habitats, particularly towards the edge of the range, even less frequently. Termena (1972) has

SORBUS TORMINALIS IN THE BRITISH ISLES 227

shown that both temperature and humidity affect pollen viability and fruit production of the species
in the Bukovina area on the borders of the Ukraine and Romania. In northern Poland good crops of
fruit were only produced every 25 years (Conwentz 1895).

Although the fruit is avidly devoured by birds, Wild Service seems only rarely to be bird-sown
(the seed, with its thin testa, is probably digested in the bird’s gut) (Prime 1960). In large, lowland
forests wild boar and other animals, including domestic pigs and cattle, may well have been
important agents for the dispersal and burial of Wild Service seed: wild boar are known to like the
fruit and the effects of their trampling on woodland ecology have been well-documented (Conwentz
1895; Darling & Morton Boyd 1969; Goodwin 1975; Tansley 1968). Elsewhere seed was, and still is,
extensively predated by birds, small animals and invertebrates so that almost none remains (Corbet
1974; Janzen 1970; Tansley 1968; Termena 1972).

The wild boar as well as burying much seed by rooting and trampling, destroyed many small
rodents (Tansley 1968) as did the much higher numbers of predatory animals and birds that were
formerly widespread. Populations of voles and mice have increased substantially as predators have
declined and animals like rabbits, grey squirrels and pheasants (all of which eat seeds or seedlings of
Wild Service) have been introduced and have spread. :

Like many trees and shrubs within the family Rosaceae, Wild Service seeds need a period of some
three months of near freezing temperatures before germination will take place (Gordon 1982). In
places where winters are longer and colder than in much of Britain, germination will normally take
place in the first spring following seed formation whereas in Britain two or more years are often
needed and the seed is at risk for far longer. This is true of many tree seeds, but the first spring
germination that would have taken place more regularly during periods when the climate was colder
could have helped the Wild Service to reproduce from seed in slightly larger numbers in those days.

While a cooler, less continental, climate and increased seed and seedling predation coupled with
other factors may have reduced populations of the Wild Service and prevented recolonisation, its
survival in ancient hedges and woodlands has been helped by its ability to reproduce from suckers.
Wild Services live a long time: Mitchell (pers. comm., 1975) has estimated the age of large old trees
to be around 200 years and many of these may themselves have arisen from suckers produced from
an earlier generation of trees. Some mature populations — that in Epping Forest, for example — have
been shown to originate largely from suckers (Lloyd 1977) and O. Buckle (pers. comm., 1975) was
of the view that virtually all the Wild Services that he knew of in West Sussex (for which he was
B.S.B.I. vice-county recorder) had originated from suckers. No one knows how far back these
sequences may have extended since the original seeds germinated, but it is clear that the species can
survive for long periods before conditions recur in. which seeds germinate freely. Suckers are often
mistaken for seedlings and care must be taken in reaching a judgement on the origin of a given tree
(the root from which a sucker arose can often be discovered just beneath the soil surface).

As a result of the survey the Wild Service has, in the last 19 years, been shown to be more
abundant than was thought and it is clear that it had been overlooked in many places. Its range is
known to extend slightly further north than Morecambe Bay and further into Yorkshire than was
realised. It is also widespread, though rare, in the lower lying parts of Wales. There are, however,
many areas in the British Isles where it might be expected to occur from which it has not been
reported.

More evidence has come to light to show that the tree was formerly more abundant than today,

though probably never common in most areas. There is no doubt that the species is found almost
exclusively in ancient woodlands and hedges or on rocky outcrops, unless planted, and its value as
an indicator of primary woodland is confirmed.
_ The survey has also revealed that the tree has had a considerable social and economic role much
of which has not hitherto been recorded or gathered together and which has undoubtedly affected
the current pattern of distribution since the tree has been conserved and planted in some areas and
neglected or destroyed in others. This is the subject of a separate paper (Roper 1987).

Although the formal survey of the Wild Service is now complete, I am always interested to have
any new records or information about the natural or social history of the tree. There is much work
still to be done on its associations with insects, fungi, bryophytes and lichens and the uses to which
the fruit and the wood were put.

P. ROPER

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ACKNOWLEDGMENTS

I am grateful to all the B.S.B.I. members and the many others who have helped with the survey over
a long period of years and have patiently awaited the results. In particular I am indebted to Mary
Briggs and Franklyn Perring without whose initial support and continuing encouragement I would
not have become involved, and to Breda and Ernie Burt who showed me my first Wild Service.

REFERENCES

ANON. (1976). Sorbus torminalis in Powys, Cardigan and Carmarthen. B.S.B.I. Welsh Bulletin 25: 2-4.

AupryY, J. (1685). Natural history of Wiltshire, ed. Britton, J. (1847). London.

BaAmMrorD, F., ed. (1936). A Royalist’s notebook. The commonplace book of Sir John Oglander Kt. of Nunwell.
London.

Boutcer, G. S. (1908). Familiar trees. London.

BRoMFIELD, W. A. (1856). Flora Vectensis. London.

Brown, J. (1894), in NisBet, J., ed. The forester, 6th ed., enlarged. Edinburgh & London.

BuscEn, M. (1929). The structure and life of forest trees, 3rd ed. revised by MUncu, E. London.

CiLapHaM, A. R., Tutin, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed. Cambridge.

CiarE, J. (1832). The midsummer cushion, ed. TrBBLE, A. Edition published by the Mid-Northumberland Arts
Group, 1978. Manchester.

ConvENTZ, H. (1895). Beobachtungen tiber seltene Waldbaum im West Preussen. Abhandlung zur Landes-
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(Accepted January 1993)

Watsonia, 19, 231-245 (1993) Za

Cardamine bulbifera (L.) Crantz (Cruciferae) in the
British Isles

A. J. SHOWLER
12 Wedgwood Drive, Hughenden Valley, High Wycombe, Buckinghamshire, HP14 4PA
and
T. C. G. RICH

37 Hartfield Road, Forest Row, East Sussex, RHI8 5DY

ABSTRACT

The taxonomy, reproduction, dispersal, habitats and distribution of Cardamine bulbifera (L.) Crantz
(Cruciferae) in the British Isles are described, and all localities traced are listed. It is currently known from about
155 native and 27 introduced sites. It is a scarce plant in Britain; the main threats to its survival are woodland
replanting and clearance.

INTRODUCTION

Coralroot (Cardamine bulbifera (L.) Crantz)* was first recorded in Britain by J. Goodyer, ‘“‘at
Mayfield in Sussex in a wood called Highreede’’, in 1634 (Wolley-Dod 1937), and it is now known to
be a very local, native plant of ancient woodlands in South East and Central England. It is widely
introduced elsewhere in England, Scotland and Ireland. On continental Europe, the plant is also
local, occurring widely from central France eastwards (though rare near the Mediterranean coast)
to the Black Sea, the Caucasus and northern Iran, and northwards to 64° in southern Scandinavia.

At least 13 species of Cardamine have been recorded in the British Isles. In addition to C.
bulbifera, there are five other native species: C. amara L., C. flexuosa With., C. hirsuta L., C.
impatiens L. and C. pratensis L. (Rich 1991). Cardamine chelidonia Lam., non L., C. glanduligera
O. Schwarz, C. heptaphylla (Villars) O. E. Schulz, C. kitaibelii Becherer, C. pentaphyllos (L.)
Crantz, C. raphanifolia Pourret and C. trifolia L. have been reported as introduced and are
variously naturalized. C. bulbifera is easily distinguished by the large pinkish-purple flowers, scaly
rhizomes and axillary bulbils. Some authors (e.g. Rose 1981) transfer C. bulbifera to the genus
Dentaria L.

Cardamine bulbifera is considered to be a scarce plant in Britain (Stewart & Pearman 1991). The
purpose of this paper is to document its ecology, occurrence and current status.

TAXONOMY AND VARIATION

Cardamine bulbifera shows little morphological variation. Schulz (1903) and Hegi (1958) noted a
number of varieties and forms, of which only two have been noted in Britain, in addition to the
typical C. bulbifera forma bulbifera. Forma ptarmicifolia (DC.) O. E. Schulz is distinguished by the
broadly serrate teeth on the leaves; the leaflets also tend to be ovate and more asymmetrical (Fig. 1),
and the plant is generally bigger and with browner bulbils. It is probably native in scattered localities
through Europe to the Caucasus but appears commonest in the Alps, and has been introduced to

*Nomenclature follows Stace (1991), and Jones (1964) for Cardamine.

232 A. J. SHOWLER AND T. C. G. RICH

Figure 1. Basal leaf shapes of Cardamine bulbifera. a—c, forma bulbifera (native). a. Fennels Wood; b. Booker
Common; c. Hawkhurst. d-f, forma ptarmicifolia (introduced). d. Silverdale; e. Trudoxhill; f. Warley Place.
The scale bar is marked in cm intervals.

and naturalized in a number of localities in Britain. Forma Jactea (Wirtgen) O. E. Schulz has white
petals, and has only been cultivated in flower beds at Saville Gardens, Windsor Great Park (v.c. 17).
Rich (1991) provides a description of native British material.

Examination of herbarium material of typical C. bulbifera show clinal variations in leaf shape
across its natural distribution. The variation is most marked in basal leaves, but as these are rarely
collected the variation is illustrated from middle stem leaves which are generally present on
herbarium specimens (Fig. 2). Material similar in leaf shape to those of native British plants occurs
in most of western Europe, specimens from Belgium being identical. Plants with broad leaves tend
to occur mainly in the Alps. In Scandinavia, the leaflets tend to be longer and less toothed, and
eastwards towards the Commonwealth of Independent States, the leaf shape is often narrower. In
Bulgaria, Greece and Turkey, at the S.E. end of the range, the leaf margins often have pronounced
teeth (but not as pronounced as in forma ptarmicifolia).

ALLEL LL LL EL,

ee ime

CARDAMINE BULBIFERA IN THE BRITISH ISLES 233

Ficure 2. Clinal variation in representative middle stem leaf shapes of Cardamine bulbifera (distribution map
redrawn from Hegi 1958). Outlying localities are shown as dots (@). The leaves are not drawn to exact scale.

REPRODUCTION AND DISPERSAL

Amongst the British flora, C. bulbifera is unusual in that it mainly propagates vegetatively by the
axillary bulbils. The bulbils are probably reduced shoots with fleshy scale leaves (Hegi 1958), and
are typically about 1 cm long, and black to dark purple. They are readily dislodged from the plant
from June onwards. They fall to the ground and, after about four weeks, adventitious roots appear
from the axils of the scale leaves, and then a few first leaves. In the second year, the plant produces
the typical scaly rhizome with basal leaves, and in the third or fourth year, aerial shoots. Plants
grown from bulbils have flowered in their third years in Britain (Ferroussat 1982). Occasionally,
bulbils may start developing whilst on the plant; a plant brought to Maidstone Museum in late 1986
had some bulbils 10 cm long (E. G. Philp, pers. comm., 1989).

Many authors note that although flowering is common, C. bulbifera seldom produces ripe fruit or
viable seed. Schulz (1903) had only seen fruiting specimens five times, and noted that they were all
from near the coast, and that where it did fruit, bulbils were less numerous or not formed at all. This
latter observation does not appear to apply to British material, fruiting plants regularly having
bulbils. Hegi (1958) noted that fruit is only set under special conditions, usually in areas with high
humidity and fresh limestone soils. These observations appear to be based on only a limited
selection of material; examination of specimens at K and BM suggests that fruit set becomes more
frequent at the south east end of the range, and in Turkey, some populations reproduce solely from
fruit, lacking bulbils altogether. Reproduction by bulbils, does, however, seem to be the norm in
Western Europe, and there may be a trade-off between fruit set and bulbil production — if the latter
are stripped off the plant, fruit set is apparently more likely (Hegi 1958).

In the field in Britain, at least, it is generally not difficult to find a few fruits. Fruits are always
borne on the larger plants (typically 22-25 cm tall, compared to the normal average of 19-21 cm),

234 A. J. SHOWLER AND T. C. G. RICH

whose stems and foliage remain green and fresh until the seeds are shed. Thus if a patch of flowering
plants is revisited in June, there may be one or two fruits maturing on a small proportion of the
plants, whereas on the remainder, the small green ovaries which showed some initial development
will have dropped off and the plants will have started to die back. In July, when the fruits are ripe
(though green) the seeds may be scattered a considerable distance when the siliquae dehisce
explosively (as in some other species of Cardamine) (Showler 1988). These seeds can be germinated
successfully, and plants grown from seed may flower in their third year (Ferroussat 1982). Fruit set
appears to have been unaffected by the opening up of the tree canopy by the great storms of 1987 in
Britain (C. I. Pogson, pers. comm., 1988).

None-the-less, the sexual reproductive performance of the plant is well below its potential. The
plants flower for only a very short period of time in late spring, and Clapham (Clapham et al. 1987)
reports that they are rarely visited by insects. B. (1866) notes that the flowers have a faint, sweet
scent. Ferroussat (1982) noted that plants at Old Park Wood were pollinated by Orange-tip
(Anthocharis cardamines L.) and Green-veined White (Pieris napi L.) butterflies, together with the -
Raspberry Beetle (Byturus tomentosus Degeer (B. urbanus Lindemann)) and flies. Occasional visits
of beetles and flies have been seen during the current survey, though it is not known if these were
pollinators. Often only a small proportion of the plants flower, many reproducing solely by bulbils.
The flowers may require cross-pollination (many crucifers have well-developed self-incompatibility
systems) to produce seed. Some stamens and ovaries may have retarded development (Hegi 1958),
pollen grains may degenerate, and some ovules do not develop at all. There is no support for the
observation of Deakin (1871) that the flowers ‘‘are often imperfect”. The species is a high polyploid
with 2n=96 (duodecaploid) (Clapham et al. 1987), and the reduced fertility may also involve
abnormal chromosome behaviour. Hegi (1958) points out that such reductions in fertility are often
associated with taxa of hybrid origin, but there is no evidence to suggest this in C. bulbifera.

Cardamine bulbifera spreads vegetatively by creeping rhizomes and often forms patches.

Bulbils (and seed) may be dispersed naturally in a number of ways. Hegi (1958) reports that
bulbils are often carried by ants in continental Europe, but this has not been observed in Britain.
Many localities in Kent and Sussex are on, or close to, river banks and ditches, suggesting that
dispersal by water may occur; Rose (1966) suggested that this may govern its micro-distribution in
the Weald. Bulbils may also be transported by wheels, hooves or muddy boots; the majority of
localities in the Chilterns and close by (v.cc. 20, 21 and 24) are alongside footpaths, though this
could also reflect other ecological factors. It is also possible that small mammals play some part in
distribution, but this has not yet been noted.

Whatever the mechanism, dispersal is obviously a limiting factor of distribution. Plants are often |

locally abundant but absent from apparently suitable sites nearby. On a wider scale, the distribution
is often irregular and disjunct, and there are many gaps between populations (e.g. Hegi 1958). The
plant does not appear currently to be limited climatically in Britain; it survives well in many
introduced localities outside its native range, often persisting for long periods of time and spreading,

as at Glenbervie (v.c. 91), where, since 1934 or before, it has colonised many parts of the extensive |

grounds and now forms a very large population.

HABITATS

In South East England, C. bulbifera is associated with two distinct types of woodland; first, the wet,
generally acidic woodlands of the High Weald in Kent and Sussex, and second, the Chiltern Beech
woods which are drier with basic soils. These two types are referred to subsequently as the High
Weald and Chiltern.

The High Weald woodlands occur on both sides of the Kent-East Sussex border. They are for the
most part ancient woodlands, often now broken into quite small areas, called ‘shaws’ or ‘rews’ — thin
strips of woodland left between fields (Whitebread ef al. 1989a, b). They have survived clearance
primarily due to the presence of the many small, steep-sided streams or gills, which make the land
unsuitable for agriculture. The soil is generally acidic to neutral (the soil pH ranged from 5 to 7) and
clayey, and the tree canopy is predominantly Quercus robur L., Fraxinus excelsior L. and Carpinus

CARDAMINE BULBIFERA IN THE BRITISH ISLES 239.

betulus L. (Table 1). In the shrub layer, Corylus avellana L. is common, often with much Crataegus
laevigata (Poiret) DC. The herb layer is usually very dense with many spring flowers such as
Hyacinthoides non-scripta (L.) Chouard ex Rothm., Mercurialis perennis L., Ranunculus ficaria L.,
Lamiastrum galeobdolon (L.) Ehrend. & Polatschek and Allium ursinum L. Carex pendula
Hudson is common in the damper places, and it is here especially that C. bulbifera is to be found,
most frequently on the lower levels of the gills. It is often easiest to find the plant by walking up the
stream beds, but even then the plants may not be easy to spot as in dappled sunlight small patches
can be easily overlooked amongst H. non-scripta and other species of a similar height, and they are
often very localised. For instance, by the Kent Ditch (a small stream marking the Kent-Sussex
border for much of its length), C. bulbifera can occur on one bank but not the other 50 cm away.
Many populations only have 15-20 flowering plants, usually, though not always, with juveniles, all
occurring within a radius of 2-3 m. Plants are not always restricted to stream banks, and may be
found nearby in damp, shaded areas of woodland. At a number of stations colonies are found on
damp, sloping road verges, though these are often associated with ditches or water seepage.

Almost all the Kent and East Sussex sites are on Weald Clay or Wadhurst Clay and are of the
High Weald habitat, as are the native sites in Surrey (two now destroyed) and those in the eastern
part of West Sussex. However, at Harting (v.c. 13) (where the plants are probably introduced),
High Rocks (v.c. 16) and Hawkenbury (v.c. 16), the plants are found on sandy soils. The single plant
found near Brown’s Wood (v.c. 14) is from a sandy road verge.

In contrast to the High Weald habitat, the Chiltern woodlands where C. bulbifera is also found
are typically with a canopy of Fagus sylvatica L., often with some Prunus avium (L.) L. and Fraxinus
excelsior, an understorey of Corylus avellana and Ilex aquifolium L., and a sparse herb layer (Table
1). The woodlands are relatively dry and on slopes, though aspect appears unimportant. The soil is
generally a very thin layer of clay with flints over chalk; the pH of the soil around the roots of the
plant ranges from about pH 6-0 to 7-5, and often there is a considerable amount of leaf litter.
Hughes (1988) and Robinson (1988) acknowledge that C. bulbifera is a plant of ancient woodland
and almost without exception, these woodlands are classified as such (Table 2). The herb layer
confirms this, typically consisting of sparse Mercurialis perennis, Rubus spp., Hedera helix L.,
Galium odoratum (L.) Scop., Arum maculatum L. and Lamiastrum galeobdolon. Hyacinthoides
non-scripta is often absent reflecting its preference for the more acidic soils, and Carex pendula, a
common associate in the High Weald, is totally absent.

Cardamine bulbifera can be found in such Chiltern woodlands in Buckinghamshire, Hertfordshire
and Middlesex, generally in small patches or strips along the edges of footpaths or close to them.
The sparse ground layer and gently sloping terrain makes the plants more conspicuous than in the
High Weald woodlands. Again there are frequently only 20-30 flowering plants (though 100-150 is
not uncommon), but the number of juveniles ranges from very few to several thousand. The plants
will grow through a thin covering of Mercurialis perennis, Rubus spp., or Hedera helix, and
sometimes C. bulbifera grows in open areas where leaf litter has blown clear, usually as a tight patch.
There are only two Chiltern sites on road verges, but this may be because wooded verges are much
less common in the Chilterns than the Weald. There is no association with water, and only two
Chiltern sites are anywhere near water courses and one of these is man-made.

The plants in Staffordshire in the ancient Needwood Forest are found in a habitat which
approximates to the Chiltern type, but are on flat land and on Keuper marl, not chalk.

Plants can often be found in replanted ancient woodland, provided that there has not been too
much disturbance to the soil and ground layer. They are able to survive in deciduous woodlands, but
_ are lost under dense conifer shade. Trimen (1862) noted that in Sussex “‘it appears to like copses
recently cleared of underwood”’.

Elsewhere in Europe, C. bulbifera also occurs in a similar range of acidic and basic woodlands
(e.g. Keller 1988), though most authors only note that it is characteristic of base-rich Beech woods.
Hartmann (cited in Hegi 1958) suggests that in the more continental areas it prefers damper soils,
_and at lower altitudes, base-rich soils. In Southern Europe, it only occurs in the mountains, and
often on north-facing slopes.

Table 2 classifies the known stations by woodland type as identified in the N.C.C. Inventories of
Ancient Woodland (Hughes 1988; Robinson 1988; Whitebread et al. 1989a, b; Hutton 1990) for
each vice-county. It should be borne in mind that areas of less than two hectares are excluded from
these inventories so that some small sites are not classified.

236 A. J. SHOWLER AND T. C. G. RICH

TABLE 1. VASCULAR PLANT SPECIES RECORDED WITHIN 2 M OF CARDA-
MINE BULBIFERA IN THE HIGH WEALD AND CHILTERN HABITATS
Only species occurring in 15% or more of either site are included; a further 56 species
were also noted at lower frequencies.

Recorded in % of sites

HIGH WEALD* CHILTERNS

Species (n = 27) (n = 57)
Acer pseudoplatanus 15 12
Alnus glutinosa 19 0
Carpinus betulus 52 16
Corylus avellana 48 25
Crataegus laevigata 22 0
Fagus sylvatica 11 74
Fraxinus excelsior 26 333)
Ilex aquifolium i) 21
Prunus avium 0 16
Quercus spp. 26 9
Allium ursinum 19 2
Arum maculatum 4 30
Carex pendula 30 0
Circaea lutetiana 15 5
Galium aparine 4] 19
Galium odoratum 7 33
Geranium robertianum 15 ih
Hedera helix 70 37
Heracleum sphondylium 15 2
Hyacinthoides non-scripta 30 30
Lamiastrum galeobdolon 26 21
Melica uniflora i 24
Mercurialis perennis 45 58
Ranunculus ficaria 19 11
Rubus spp. 74 69
Urtica dioica 1) 16
* excluding High Rocks.
DISTRIBUTION

In Britain, Cardamine bulbifera is currently known as a native species in Sussex (v.cc. 13 and 14),
Kent (v.cc. 15 and 16), Surrey (v.c. 17), Hertfordshire (v.c. 20), Middlesex (v.c. 21), Buckingham-
shire (v.c. 24) and probably Staffordshire (v.c. 39) (Fig. 3). It has not been recorded in its sole
Berkshire (v.c. 22) locality since 1944.

In view of their isolation from the other sites in South East England, and the regularity with which
the plant is introduced, the Staffordshire sites must be viewed with caution. The first record from
Pendeford (Pitt 1796) was unusual in that it occurred in “hedge sides on this farm’’; however, Pitt’s
list also included other species of old hedgerows (e.g. Rhamnus cathartica L., Frangula alnus Miller)
and is packed with detailed, careful botanical observations and there is no obvious reason to reject
the record. There are at least three localities around Needwood Forest, which have sometimes been
suggested to be introduced (Edees 1972), and there is an undoubted introduction at Trentham.
Edees (1972), whilst quoting it as “‘rare and of doubtful status” accepts Pendeford and Blithfield as
native localities. D. P. Earl (pers. comm, 1987) notes that colonies he has seen recently “look native
on the Keuper Mar!l”’ but also points out that none of these are far from houses and two are close toa
road or track, suggesting that they may have originally been planted. The evidence from Chiltern
populations, also associated with tracks, suggests that this need not necessarily be so, but ‘looking
native’ is not always a good guide to status either (Webb 1985). As the early dates of recording lend

CARDAMINE BULBIFERA IN THE BRITISH ISLES 23

TABLE 2. ASSOCIATION OF NATIVE POPULATIONS OF CARDAMINE BULBIFERA WITH
ANCIENT WOODLANDS AS CLASSIFIED IN THE N.C.C. INVENTORIES OF ANCIENT

WOODLAND
Ancient woodland
No. TIP LRM i La ee Other

Vice-county sites Semi-natural . Replanted habitats*
13. W. Sussex 8 4 1 2
14. E. Sussex 33 19 4 10
15SSE~ Kent 3 5 = i

16. W. Kent 19 12 6 1
17. Surrey 1 = 1 —
20. Herts. 12 7 - 5)
21. Middx. 2 M2 - ~
24. Bucks. 56 38 13 5)
39. Staffs. 2 - 2 -
Total 136 85 (62%) 27 (20%) 24 (18%)

* includes sites too small (less than 2 ha) to classify.

support to its native status, and as the plant also occurs in disjunct localities elsewhere in Europe, we
currently accept the Needwood Forest plants as native.

Elsewhere in the British Isles, the plant has been introduced. It is often grown in gardens, perhaps
as an unusual plant — it is certainly a pretty, early-flowering plant which can tolerate dense shade,
though there are many other species with these attributes which flower for longer periods of time.
Clear evidence is hard to come by, but C. bulbifera appears to have been cultivated in large gardens
for many years (e.g. B. 1866). Once established, it can often increase rapidly and become almost a
weed, as at Uffculme (v.c. 3), and at Knightshayes Court (v.c. 4) and Cliveden (v.c. 24) where it
appears sporadically in flower beds. Escapes to adjacent verges or woodland, as at Bath (v.c. 6),
Wellingore Hall Park (v.c. 53) and Silverdale (v.c. 60), are not uncommon, and it may persist for
long periods of time. There are also many scattered records for introduced plants from the 1920s and
1930s, where it can no longer be found. Bulbils are easily collected, and the flourishing colony at
Trentham (v.c. 39) may well have originated from the plants at Yoxall, and the casuals in v.c. 95
could have come from Glenbervie, not many kilometres away. Interestingly, the majority of West
Country introductions are of forma ptarmicifolia suggesting a common origin. It would certainly be
instructive to learn more of the history of the known introductions.

The plant has also been recorded as introduced to some localities in Germany (Hegi 1958).

RECORDS

Records have been collated from national and county Floras, journals, correspondence with the
B.S.B.I. vice-county Recorders and numerous other botanists, and from the following herbaria:
BEL, BM, DBN, K, OXF, LANC, RNG, TCD and YRK. Full details, including population forms,
are lodged at the Biological Records Centre, Monks Wood.

In the following list of localities, vice-counties or sites where the plant is introduced are marked
with an asterisk (*). The records are given in order of vice-county, 10-km square, and tetrad (tetrad
nomenclature follows Ellis 1986). Exact grid references are given where known for extinct or
unvisited sites. The names for modern sites are largely those on the First or Second Series 1:25,000
or 1:50,000 Ordnance Survey maps. An estimate of the total number of plants is given for most sites
(sometimes only as the number of flowering plants counted). All undated sites have been visited
recently between 1987 and 1992 by A.J.S., often with help from co-workers, unless otherwise
stated. Sites where we have seen material of forma ptarmicifolia are noted.

*V.c. 3, S. Devon: Bere Ferrers (SX/4.6L), 1916, C. W. Bracken (Martin & Fraser 1939), presumed

bo
G2
(oe)

A. J. SHOWLER AND T. C. G. RICH

Ficure 3. Current 10-km square distribution of Cardamine bulbifera in Britain and Ireland. @ native 1987
onwards, © native pre-1987, < introduction (all dates), } record of uncertain status.

extinct. Plymstock (SX/5.5B), Plympton (SX/5.5N) and Harford (SX/6.5J), 1913, H. W. Smith
(Martin & Fraser 1939), presumed extinct. Dartington Hall (SX/7.6W), woodland area of garden,
C. Smith. Torbryan Plantation (SX/8.61), “introduced by Mr Ogilvie and now spreading” (Martin
& Fraser 1939), presumed extinct. Venn Ottery (SY/0.9R), over 800 plants, shaded lane verge,
escaped from adjoining garden, W. Tucker (forma pfarmicifolia). Uffculme (ST/0.1R), garden of
Yondercott House, thousands of plants over 0-25 hectare, in thin, damp woodland, Miss R. G.
Laidlaw.

*V.c. 4, N. Devon: Sherwill (SX/7.6), 1884, Wainwright (Martin & Fraser 1939). Knightshayes
Court, National Trust (SS/9.1S), gardens by house, Miss R. G. Laidlaw (forma ptarmicifolia).
*V.c. 6, N. Somerset: Millards Hill, Trudoxhill (ST/7.4L), in shrubbery and roadsides, escaped from

Millards Hill House, P. Green & R. G. B. Roe (forma ptarmicifolia). Smallcombe Wood, Bath —

(ST/7.6S), scattered throughout but more frequent near ruins of adjacent garden, R. Randall &

R. G. B. Roe (forma ptarmicifolia). Prior Park, Bath (ST/7.6), at one time abundant, destroyed

by building work (Murray 1896). Batheaston (ST/7.6Z), about 500 plants, shaded track behind

houses, probably dumped from garden, D. E. Green & R. G. B. Roe (forma ptarmicifolia).
*V.c. 11, S. Hants.: Millbrook (SU/3.1W), J. F. Rayner (1915); presumed extinct.

CARDAMINE BULBIFERA IN THE BRITISH ISLES 239

*V.c. 12, N. Hants.: Long since disappeared from East Ockley House (SU/5.5Q), D. H. Scott.
Small copse near Preston Candover (SU/6.4), 1879, H. R. P. Fitzgerald, and Basingstoke (SU/
6.5), 1916, G. W. Willis (Townsend 1904; Rayner 1929).

V.c. 13, W. Sussex: *Hotham Park, Bognor (SZ/9.9J), garden of former Bognor Museum, now
demolished (H. W. Matcham teste M. Briggs). *Hunstan Copse (SU/8.0, R/Q), H. L. F.
Guermonprez, ancient wood; searched for recently without success by N. J. H. Sturt, and
probably last seen in the 1940s by F. Rose (pers. comm. to Mrs M. Briggs, 1987). *Harting Combe
(SU/8.2C), one large clump, road side in Beech wood. A record for SU/8.2D (Hall 1980) is
probably an error for this site. Half-way between Midhurst and Petworth (SU/9.2); this record,
cited in Arnold (1907) and Wolley-Dod (1937), is based on a specimen said to be in herb. Borrer,
which cannot be traced at K. Although the area is suitable, there are no other records and this site
must be treated with caution. Warnham (TQ/1.3), “small copses in the parish’, 25 April 1862, H.
Trimen (Trimen 1862); exact site not traced, and presumed extinct. Langhurst Copse (TQ/1.3S),
wooded verge south of, 2000+ plants. Tickfold Gill (TQ/1.3T), by stream in wood, 29 plants
(“hundreds” were reported for this site in 1987, F. Rose etal. (pers. comm. to M. Briggs, 1987) so
possibly overlooked). Nuns Wood (TQ/1.3, X and Y), a few plants in the woodland at the
‘southern end, Mrs M. Briggs, 1961, and very sparse in western stream gulley in 1988. Great
Benhams (TQ/1.3Y), north of, about 200 plants, shaded bank of Boldings Brook by bridge; this
site is declining due to invasion by Heracleum mantegazzianum (A. Knapp, pers. comm., 1992).
Horsegills Wood, Rusper (TO/1.3Y), wet woodland, 600+ plants, Mrs M. Briggs, 10 May 1991.
Terry’s Cross (TQ/2.1H), 500 plants in hedge. (The record for TQ/2.2M (Briggs 1990) is an
error.) Faygate (TQ/2.3), near, C. E. Salmon (Wolley-Dod 1937), probably extinct. There is a 40-
column record card held at B.R.C. with details ““Rusper, 11 June 1957, Collyers School
Herbarium” which has the grid reference ““TQ/20—.32—”. As this grid reference refers to Roffey,
Horsham, about 5 km south of Rusper, and C. bulbifera is known near Rusper in TQ/1.3, this
record is dubious. Unfortunately, the Collyers School Herbarium cannot now be traced (Kent &
Allen 1984) and it is impossible to check if the grid reference was correctly transcribed, or was
added later to the B.R.C. card.

V.c. 14, E. Sussex: Withyham (TQ/4.3X), 140 plants in wood by Hewkins Bridge and also in
churchyard, Miss E. J. Rich, 1992. Mayfield, Lawyers Wood (TQ/5.2T), Miss E. J. Rich, 6 July
1987; fruiting plants seen in 1992. Highreede Wood, Mayfield (TQ/5.2), J. Goodyer 1634; exact
site not traced (Wolley-Dod 1937). Old Place Farm (TQ/5.2Y), one plant, E. side of road, Mrs P.
Donovan, 1992, and roadside copse, Miss E. J. Rich, fruiting in 1992. Coggins Mill, by bridge
(TOQ/S.2Y), Miss E. J. Rich, 1992. Heronry Wood (TQ/5.2Z), 15 plants, wet muddy area close to
entrance of wood near stream. P. C. Hall noted it as abundant in this wood when he was recording
for the Sussex plant atlas (pers. comm., 1991) so possibly some colonies were overlooked.
Furnace Wood (TQ/5.2Z), 6 m long patch on east verge of lane, Mrs P. Donovan, 1992. Banky
Wood (TQ/5.2Z), four scattered groups in wet woodland, some on stream bank. TQ/5.3N, Hall
(1980), further details not traced. Houndsell Place (TQ/5.3V), in ‘wild garden’ (wood), P. C.
Hall, 26 April 1970. Mark Cross (TQ/5.3), in a hedge by the road side about a mile (1.5 km) from
(Deakin 1871); presumed extinct. Great Wood (TO/5.3X), P. C. Hall, 31 May 1969. Coggins
Mill, (TQ/6.2D) stream bank 200 m east of, Miss E. J. Rich, 1992. Tidebrook (TQ/6.2E), large
numbers of plants along 50-100 m length of road verge in woodland, and 20 flowering plants in
damp, shady woodland to north. Mousehall (TQ/6.2E), 110 plants in lane east of Mill, and two in
garden of Mousehall Cottage on old dam embankment, Mrs E. Gibb, 1992. Sharnden (TQ/6.2E),
about 40 plants, wet woodland. Hawksden Park Wood, west of Hare Holt (TQ/6.21), about 15

- plants in hedgebank and ditch. Foxholes Wood (TQ/6.2G), J. Goodyer, 1634; not seen
subsequently (Wolley-Dod 1937). TQ/6.2K, Hall (1980), exact site not recalled, possibly Cox’s
Mill (pers. comm., P. C. Hall, 1991). Burwash Weald (TQ/6.2L), three sites: steep bank near
Mousehole Farm, 325 plants; Willingford bridge, 25 flowering plants on steep woodland stream
bank and then clumps along the north bank of the R. Dudwell for about 1-5 km S.W. of bridge (F.
R. Philps, pers. comm., 1987); Blackbrooks, about 100 scattered, flowering plants in wood.
Coalpit Wood and Wet Wood (TQ/6.2M), about 500 plants in damp woodland. Wood N.W. of
Stonegate Station (TQ/6.2, N & T), about 100 plants in damp Hornbeam wood. Rye Green Farm
(TQ/6.2R), two sites: woodland west of farm with about 100 scattered, flowering plants; Bog
Wood, 100 flowering plants on south bank, 60 on north bank. Bateman’s (TQ/6.2R), 140 plants

240 A. J. SHOWLER AND T. C. G. RICH

flowering along 25 m of dense hedge, and 500 plants in copse to north. Fonthill Farm (TOQ/6.2W),
farm track, 60 plants. Copse between Boarders Farm and Dudwell House (TQ/6.2X), P. C. & J.
F. Hall, 31 March 1968; site now destroyed. Shoyswell Wood (TQ/6.2, ?Y/Z), Mrs M. Warren,
1972 (there is some doubt about the exact location of this site as the original grid reference was
given as TQ/688.261). Burgham (TQ/699.284Z), un-named wood N.W. of, Mrs M. Warren,
1972. Wardsbrook Farm (TQ/6.2Z), about 500 plants in wood by stream. Singehurst Farm (TQ/ |
6.2Z), neglected, small, swampy wood S.E. of, probably only a few plants, Mrs L. B. Burt, 1984;
not seen in 1989 but probably overlooked. Rivenhall (TQ/6.3B), large patch (5 m long) on road
verge with a few plants in ditch behind, Mrs M. Vincent-Smith, 1992. Brown’s Wood, road verge
opposite Coker’s Down (TQ/6.3D), one flowering plant. (The record for TQ/6.3E in Briggs
(1990) refers to High Wood, v.c. 16.) Brown’s Wood, near Sunninglye Farm (TQ/6.3J), with 180
plants on verge in hedge, 30 flowering plants on verge in edge of wood, and 215 plants in wood.
Sluice Wood (TQ/6.3M), thinly scattered on stream bank in dense woodland, mostly on north
side of stream, one large patch on south side of stream. Win Bridge (TQ/6.3M), 30+ flowering |
plants in small copse by administrative county (but not v.c.) boundary (presumably same site as
Bayham Abbey (Arnold 1907, etc.)). Floshet Wood (TQ/6.3N), nine plants in wood by stream;
this site is in the administrative county of Kent. Park Wood, near Sidley Green (TQ/7.1, F/K), H.
J. Sargent (Wolley-Dod 1937); not seen recently. Goldspur Wood (TQ/7.1J), one very large
clump. Hollington Wood (TQ/7.1V), A. H. Simpson, 1918 (K); now apparently built on, though
recorded for this tetrad in Hall (1980). Beauport Park (TQ/7.1, ?W/X) (Wolley-Dod 1937); this
site is now a caravan park and the plant is presumed extinct. Bluemans (TQ/7.1X), wet, shady
woodland, with 50 flowering plants (many fruiting). Brookside Farm, wood (TQ/7.2C), 10+
plants on edge of recent woodland/verge. Fleet Wood (TQ/7.2D), about 40 flowering plants on —
edges of muddy bridleway in damp, shady conditions. Little Boarzell, Swiftsden House (TQ/7.2,
E & J), about 120 flowering plants (and more vegetative) mainly in the wild garden but also on
verge. Wood between Mountfield and Robertsbridge (TO/7.2, F/G), 1944, Miss Hanson
(Peatfield 1944). Etchingham, Gigmore Wood (TQ/7.2H), 20 plants in ancient woodland. Peagle
Wood (TQO/7.2J), 85 plants in wood on Sussex side of Kent Ditch, and damp woodland close to
road in adjacent tetrad (TQ/7.2P), about 3000 plants, many flowering. Copse in fork of A265 and
minor road to Merriments Shaw (TQ/753.284J), P. C. Hall, 8 May 1970; wood cleared and plant
extinct by 1988. TQ/7.2M, Hall (1980), further details not traced. Merriments Shaw (TQ/7.2P),
several patches with over 400 flowering and many vegetative plants in ancient woodland on Sussex |
side of Kent Ditch. Crossroads Farm (TQ/7.2P), about 200 plants in wood by A229 on Sussex side |
of Kent Ditch, and about 75 plants in wood 300 m south west of cross-roads. Terrace Wood (TQ/ |
7.2, S/T), about 200 plants on both sides of A229, plus one by stream. Bodiam Wood (TQ/7.2T),
one large patch near wet hollow. Records from ‘“‘about Hastings” presumably refer to sites at
Hollington, Beauport Park and Bluemans (Peatfieid 1943), and there are no localised sites in TQ/
oe

V.c. 15, E. Kent: Holman’s Wood (TQ/777.287U), woodland with wet gills, F. Rose; not seen 1991,
but probably still present. Kitchenham Farm (TQ/7.2Y), wood by Kent Ditch, E. G. Philp,
sometime between 1971 and 1980. Detling, Maidstone (TQ/7.5), A. D. Melvin, 21 June 1867
(BEL); this is a surprising record. N.E. of Hawkhurst (TQ/763.313Q), last seen 1950s (E. G.
Philp, pers. comm., 1991). Rolvenden, wood east of Halden Lane Farm (TQ/8.3L), 50+
flowering and many vegetative plants. Little Halden (TQ/8.3L), 17 flowering plants on steep
north facing bank by stream near ancient woodland, plus one downstream; this is probably the
same general locality as Littlke Oven Wood (Hanbury & Marshall 1899). Little Halden (TQ/
861.326R), F. Rose, 1949 (MNE), site now destroyed .

V.c. 16, W. Kent: TQ/S.3P, Philp (1982), likely to be near High Rocks, but further details not
known. High Rocks, Tunbridge Wells (TQ/5.3U), one large dense patch by railway with about
2400 plants, and two smaller patches to west with 250 plants; about 100 plants in scattered
colonies on dry sandstone ridges and rocks at eastern end of rocks; more plants north of rocks by -
stream. Hungershall Park, Tunbridge Wells (TQ/5.3U), 300 flowering plants in dry woodland,
presumably the same locality as Tunbridge Wells Common, K. E. Bull (Wallace 1954). Mount
Sion (TQ/5.3Z), in a wood (Forster 1816); probably built on. *Platt (TQ/616.562), D. |
McClintock, c. 1960, patch in woodland and in a hedge nearby, probably escaped from garden. |
Palmers Farm, wood (TQ/6.3E), about 100 flowering and more vegetative plants in ancient

CARDAMINE BULBIFERA IN THE BRITISH ISLES 241

woodland. High Wood (TQ/6.3E), scattered plants in five areas, with 60 flowering plants on
verge, 100 in cleared strip, and 100 in woodland, and 300 plants on dry sandy trackside in ancient
woodland, and 130 in deep grass on cleared edge of wood. Mouseden (TQ/6.3E), on stream banks
in fragments of ancient woodland, three groups with 17, 12 and 75 plants, extending into tetrad J.
Pembury, wood east of Larkfield Hall (TQ/6.3J), twelve plants by stream. N.W. of Lamberhurst
(TQ/661.377T), E. G. Philp, last seen about 1957, not refound, area spoiled by pheasant
management. TQ/7.2J, Philp (1982), further details not traced. Kent Bridge Farm (TQ/7.2P),
two sites: 1120 plants on bank of ditch in wood; 40+ plants on shaded road verge to north.
Winch’s Plantation (TQ/7.2P), 80 flowering plants in ancient woodland, well above stream level.
Peagle Wood (TQ/7.2P), a few plants on Kent side of Kent Ditch. Merriments Shaw (TQ/7.2P),
several plants on Kent side of Kent Ditch (see also v.c. 14). Goudhurst (TQ/708.374D), near
A262 west of, F. Rose, last seen 1950s; site now destroyed. Little Pix Hall Farm (TQ/7.3F), wood
north of, only two flowering plants seen, possibly more present. Furnace Wood (TQ/7.3H), south
side, one dense patch with 30 flowering plants and a few stragglers in woodland by track. Wet
Wood (TQ/7.3H), several plants near stream in replanted, wet ancient woodland; extinct at
another site at TQ/733.358. N of Hawkhurst, (TQ/757.313K), last seen 1950s (E. G. Philp, pers.
comm., 1991); possibly same as Highgate (Hanbury & Marshall 1899). Hedgingford Wood (TQ/
7.3M), two sites; north side, two patches in woodland near stream, one with 600, the other to
south with 650 flowering plants, and N.E. corner, 23 plants by stream in Hornbeam wood.
Hartley, Forestry Commission wood (TQ/7.3M) west of, 42 plants on edge of ride just clear of
conifers, an old site to east has been lost due to conifer planting. Another site at TQ/750.384M has
disappeared under conifers.

V.c. 17, Surrey: *Hog’s Back (TQ/0.5), demesne of ‘Greyfriars’, south-facing Beech wood, one
patch, J. R. Akeroyd, May 1968. *Box Hill (TQ/1.5Q), by the River Mole, Miss B. M. C.
Morgan, 1966, probably planted (Lousley 1976). About 250 plants seen in 1989, though few were
flowering. Grove Copse (TQ/1.3T), many plants in 1949, only one visible in 1966, D. P. Young.
Timber Gill (TQ/1.3T), about 100 plants in wooded ravine. Lower Gages Farm, ravine of Fylls
Brook (TQ/184.378Y), numerous small plants seen on edge of stream, Surrey Flora Committee
meeting, 1967; extinct by 1989 (and probably long before) due to activities of pigs. In deep ravine
near Garrett Farm, Ockley (TQ/1.3; precise farm not traced), both on Sussex and Surrey banks,
E. Straker, 15 May 1880 (BM); presumed different from above sites and now extinct. (The record
for Waddington (Brewer 1863) is an error for Lathraea squamaria (Salmon 1931).)

*V.c. 18, S. Essex: Warley Place (TQ/5.9V), still to be found in 1992 after 55 years in the derelict
garden of A. E. Wilmott’s home, where it was introduced (forma ptarmicifolia), D. C.
Bloomfield (Jermyn 1974).

V.c. 20, Herts.: Baldwins Wood (TQ/0.9E), L. J. Stearn, pre-1967; not seen 1989-1991 but possibly
overlooked. Bottom Wood (TQ/0.9G), one patch with 15 flowering stems. Chorleywood, Great
Greenhills Wood, west of M25 (TQ/0.9H), 200 plants (see also tetrad M). Chorleywood, west of
Dell Farm (TQ/0.91), thousands of plants; also E.N.E. of Dell Farm, over 2000+ plants on south
and west side of wood. *Chorleywood Common (TQ/0.91), 640 plants in secondary woodland
N.E. of pond; probably introduced. Bullscroft Spring (TQ/0.91), patch about 4 m7”, G. Salisbury
& J. Saunders, 1990. Sarrattmill Bridge, N.W. of (TQ/0.9J), 250 flowering plants in thin strip of
woodland. Limeshill Wood (TQ/0.9J), ‘‘a scattering of plants amongst Hyacinthoides on a steep
bank’’, G. Salisbury, 1991. Hanging Lane Wood (TQ/0.9J), over 2000 plants widespread in
woodland. Springwell, Garrett Wood (= High Wood near Rickmansworth (Webb & Coleman
1849)) (TQ/0.9L), numerous plants remaining in Beech wood, though half of the wood (mainly
the Middlesex part, where it was once abundant) has been lost to a quarry. Chorleywood, Great
Greenhills Wood east of M25 (TQ/0.9M), 3000-4000 in west half, and many thousands in the east
half, one of the largest, continuously covered areas seen. Beechengrove Wood (TQ/0.9N),
“masses Over quite a large area”, G. Salisbury, 1991. Loudwater (TO/0.9N), road verge
bordering large house, 150 plants possibly relict from Loudwater Wood (Webb & Coleman 1849),
and possibly the same as “larch plantation above Loudwater to the north, but adjoining the road
from Croxley Green to Solesbridge’’ (Jackson 1887). Another Jackson (1887) record “in a wood a
little further towards Solesbridge Wood” probably refers to Long Spring (TQ/052.970N) (T. J.
James, pers. comm., 1991). Scrubbs Wood (TQ/0.9, N/P), L. J. Stearn, presence now doubtful,
and probably the same locality as Blunt Wood, 20 April 1947, R. A. Graham, also not seen

242 A. J. SHOWLER AND T. C. G. RICH

recently. Sawpit Spring (TQ/059.982P), P. J. Ellinson, 1992; a small wood mostly destroyed
during construction of the M25. Harrocks Wood (TQ/0.9T), 400 plants in scattered colonies.
Assumed to be the same site as “wood at Red Heath’, Mrs E. Shute (Webb & Coleman 1849).
Whippendell Wood (TQ/0.9T), many, mostly small, scattered colonies (including Rousebarn
Lane, P. J. Ellison), over 3000 plants. Lees Wood (TQ/0.9U), S. & P. Blackmore, 2 June 1978. A
B.R.C. 40-column record card from ‘wood opposite Chandlers Cross, May 1922, P. W.
Richards” (NMW), presumably referring to Harrocks Wood or Whippendell Wood, was
allocated the grid reference TL/0.0 for the B.S.B.I. Maps Scheme (Perring & Walters 1962)
rather than TQ/0.9; this appears to be an error and there are no other records for TL/0.0. There is
also a record for “Brickett Wood, West Watford” (OXF). Bricket Wood (TL/1.0F) was in the
19th century an ancient wood (now the Building Research Station) and could conceivably have
supported the species as it was species-rich, calcareous and damp, but there are no other records
for this site (T. J. James, pers. comm., 1991).

woodland. Park Wood (TQ/0.9K), 100+ plants along west end of footpath in ancient wood. Jacks
Lock (TQ/0.9K), copse N.E. of, last seen by R. M. Hamilton in 1972 who described it as
“abundant”; wood now over-grown and not refound in 1989. Springwell, Garrett Wood (TQ/
0.9L); see v.c. 20.

V.c. 22, Berks.: Last recorded in Park Wood, Bisham (SU/8.8M) (presumably same as Bisham
Wood; Druce 1926), 29 April 1944, A. J. M. Bailey. A specimen in RNG labelled ‘““Newbury, C.
Grant, 1966’, has a grid reference referring to the University Campus in Reading.

V.c. 23, Oxon: Recorded from Oakley Hill, near Chinnor (SU/7.9P) but probably in error (R.
Fitter, pers. comm., 1990). |

V.c. 24, Bucks.: Bloom Wood (SU/8.8U), about 3000 plants in ancient woodland. Horton Wood
(SU/8.8U), thousands of plants in ancient Beech wood. Warren Wood (SU/871.899U), no other
details available. *Cliveden, National Trust (SU/9.8C), in the gardens near the pond. Booker
Common and Spring Coppice (SU/8.9F), a large population with about 800 plants flowering in
ancient Beech wood. Yewtree Hill Plantation (SU/8.91), 13 plants on edge of ancient woodland.
The Coppice (SU/8.91), 500+ plants in ancient Beech wood. Park Wood (SU/8.9J), Buckingham-
shire tetrad record collected 1965-1985 (R. Maycock, pers. comm., 1988), but not refound 1991—
1992. Common Wood (SU/8.9M), several thousand plants in ancient woodland, plus a few young
plants on disturbed land 200 m W.S.W. Piggott’s Wood (SU/8.9P), three populations with 300,
660 and 630 plants in main ancient wood, and one small colony with 20 plants on west side.
Deangarden Wood, near Keep Hill (SU/8.9Q), about 500+ plants in ancient Ash-Beech
woodland. The Rye (SU/8.9R), 30+ plants scattered in woodland (probably ancient). Millfield
Wood (SU/8.9S), about 500 flowering plants in ancient Beech wood. Hanging Wood (SU/8.9S),
2000-3000 plants, ancient woodland. Green Wood (SU/8.9S), about 1700 plants in four small
patches in ancient Beech woodland. Gomm’s Wood (SU/8.9T), about 5000 plants in ancient
Beech woodland. Hughenden Valley, ancient Beech woods north of Boss Lane Farm (SU/8.9T),
several thousand plants. Longrove Plantations (SU/8.9T), about 500 plants in replanted
woodland. Hughenden Valley, Citers Wood (SU/8.9U), Buckinghamshire tetrad record, 1978
(R. Maycock, pers. comm., 1988), not refound 1991-1992. Fennell’s Wood (SU/8.9V), 400+
plants in ancient Beech wood. Deangarden Wood, near Abbey Barn Farm (SU/8.9V), about 360
flowering and more vegetative plants in ancient Ash-Beech woodland. Barrowcroft Wood (SU/
8.9V), 20+ plants in ancient Ash-Beech woodland. Highfield Wood (SU/8.9W), several thousand
flowering and numerous vegetative plants scattered through ancient Beech woodland. Gomm
Valley (SU/8.9W), two small patches, old woodland bank in Nature Reserve, M. Young, May
1991. Kings Wood (SU/8.9W) about 800 plants in ancient Beech wood. Woolman’s Wood (SU/
9.8D), about 1000 plants, edge of clearing in replanted ancient woodland. Lower Pyebushes (SU/
9.8U), scattered in replanted ancient Oak-Ash wood. Cut-throat Wood (SU/9.9A), scattered in
ancient Beech wood, about 3000 plants. Hogback Wood (SU/9.9F), about 2400 plants in ancient
woodland. Seagrave’s Farm (SU/9.9G), wood N.W. of, 500 plants in ancient Beech woodland,
and 120 plants to east in narrow strip of ancient woodland. Ash Grove (SU/9.9J), 3500 plants near
railway in replanted ancient woodland. Keeper’s Wood (SU/9.9J), about 1000 plants in new
plantation of Beech and Oak. Great Beards Wood (SU/9.9L), scattered populations totalling
about 6000 plants in replanted ancient woodland (some under conifers), and 100 plants in wood to

2 a 5

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CARDAMINE BULBIFERA IN THE BRITISH ISLES 243

north. Blue Close Wood (SU/9.9L), thousands of plants, on edge of pine plantation and likely to
disappear. Owlsears Wood (SU/9.9L), 315 plants on edge of ancient wood and bank of old hedge.
Bottom Wood and Starveacre Wood (SU/9.9L), 180 flowering plants in replanted ancient
woodland. Stockings Farm (SU/9.9M), ancient woodland west of, three large populations with
3500, 300 and 210 plants. Wheatsheaf Wood (SU/9.9Q), 35 plants in recent Beech wood. Great
Legs Wood (SU/9.9, Q & V), 2000-3000 plants in four main areas in ancient Beech wood. SU/
9.9R, Buckinghamshire tetrad record collected 1965-1985, site details unavailable (R. Maycock,
pers. comm., 1988). Ongar Hill Farm (SU/9.9S), wood west of, J. Pitt, 1977; not found in 1989,
now overgrown. Parsonage Wood (SU/9.9T), 50+ plants in ancient Beech woodland. The Ash
Beds (SU/9.9V), 370 plants by track in mixed, recent woodland. *Stowe, National Trust (SP/
6.3Y), in the Japanese gardens, R. Maycock; several thousand plants of a variant approaching
forma ptarmicifolia were seen in 1992. SP/8.0Q, Buckinghamshire tetrad record collected 1965-
1985, site details unavailable (R. Maycock, pers. comm., 1988). Hyde Heath, Hyde House Wood
(SP/9.0F), D. Ferguson, 1986. White’s Wood (SP/9.0F), 200 plants, all vegetative, under dense
Spruce canopy, and 250, also all vegetative, under Beech. White House Farm (SP/9.0K),
Gracelets Wood north of, 3000 plants in old game covert. Monk’s Wood (SP/9.0K), numerous
plants in ancient wood and on bared strip. Elvidge Wood (SP/9.0K), ancient Beech wood west of,
500+ plants. Lower Bois, Hodds and Hilsbury Woods (SP/9.0Q); good numbers of plants spread
along N. edge of ancient wood, and a few plants in N.W. corner of Lower Bois Cemetery. The
Larches (SP/9.0X), only one plant seen in neglected, ancient, deciduous woodland. Newland
Park (TQ/0.9B, possibly also in tetrad C), C. J. Smith, 1979, recent woodland. Carpenters Wood
(TQ/0.9D), 120 plants in ancient woodland. Chenies (TQ/0.9E), 500 plants in wood adjoining
ancient woodland. Chenies Bottom (TQ/0.9E), in two main groups with 52 and 60 plants plus a
scattering of vegetative ones elsewhere. West Wood (TQ/0.9E), a very dense patch with 1350
plants in replanted ancient woodland. Turveylane Wood (TQ/0.9J); two small populations, one
with about ten plants, the other with about 20 plants, replanted ancient woodland. Mount Wood
(TQ/0.9J), three large groups plus scattered plants elsewhere in ancient woodland, about 650
plants. Denham Green, Ranston Covert (TL/0.8P), about 30 plants in ancient Beech Wood at
foot of very steep slope. -

V.c. 39, Staffs.: Pendeford, ‘“‘hedge sides on this farm” (SJ/8.0), Pitt (1796). Blithefield, grove by the

churchyard (SK/0.2), Mr Stanmers (Garner 1844); this site was thought to be a probable

introduction by Perring & Walters (1962), though on what basis is not known; Pendeford is at
the west end of Needwood Forest and if accepted as native at the eastern end, could well be
native here too. Needwood Forest (SK/0.1 or 0.2), Garner (1844). Ridge (1922-1929) ‘‘failed
to find the plant anywhere in Needwood Forest”, and others have followed his caution in
accepting the record. Yoxall (SK/1.2L), 40+ flowering plants on wooded road verge, replanted
ancient woodland. Edees (1972) also gives ‘“‘abundantly in a copse between Newchurch and
Scotch Hill . .. and in the wood, formerly known as Coalpit Slade, on the east side of the
road near Darleyoak Farm’’. *Ash Green, Trentham (SK/8.4, ?P/Q), wood of park, about 3000
plants. .

*V.c. 40, Salop: Broncroft Castle (SO/5.8), Miss M. B. Fuller, 1978-1992 (Sinker et al. 1985).

*V.c. 53, S. Lincs.: Wellingore Hail Park (SK/9.5Y), 1962, Miss E. J. Gibbons. Small Hornbeam
copse, with Mercurialis perennis and Adoxa moschatellina, probably escaped from the Hall
gardens (Mrs I. Weston, pers. comm., 1988).

*V.c. 55, Leics.: Glenfield, Leicester (SK/5.01) “shady border in gardens of the Gynsills’’, 1971

- (Primavesi & Evans 1988); this site is now partly under a hotel.

*V.c. 57, Derbys.: Between Marple and Strines, near the River Goyt (SJ/9.8), T. Barker (Linton
1903). Long Eaton (SK/4.3J) (Clapham 1969). Sheffield (SK/3.8L), in woodland in Graves Park,
first noted 12 April 1938, but no longer known (M. Shaw, pers. comm, 1992).

*V.c. 60, W. Lancs.: Silverdale (SD/4.7S), several patches in woodland, escaped from adjoining
Nursery. During a visit in 1988, Mr Kaye described C. bulbifera as more or less a weed, which he
had dug out in masses.

*V.c. 62, N.E. Yorks.: Scalby (TA/0.9A), “long established; the Natural History of the Scarborough
District lists it as ‘Rare. Scalby Churchyard, 1964 et seq ...’. It was then in a much more
restricted area than now. I assume it was originally planted ona grave. . . since it reached the wall
of the churchyard it has not looked back” (T. F. Medd, pers. comm., 1988). This colony even

244 A. J. SHOWLER AND T. C. G. RICH

received a mention in The Dalesman, January 1900 “‘— it thrives throughout the village”’ (an over-
statement).

*V.c. 69, Westmorland: Bowness (SD/4.9), W. Clitheroe, 1940 (forma ptarmicifolia).

*V.c. 72, Dumfriess.: Dardarroch, Dunscore (NX/8.8N), roadside verge, C. Rogers, 14 May 1981;
Mrs M. E. R. Martin, 1988 (forma ptarmicifolia). The status of another record from Maxwelton
House requires confirmation. Hooker (1870) listed C. bulbifera as ‘‘doubtfully native” in Ayr
(v.c. 75), Lanarks (v.c. 77) and Perth (v.cc. 87-89) but no specimens or further details have been
traced. Watson (1873) attributes the Ayr record to a specimen from a James Wilson.

*V.c. 91, Kincardines.: Glenbervie House (NO/7.8Q), very widely distributed in the garden, known
since at least 1934. The biggest introduced colony.

*V.c. 95, Moray: Dyke (NH/9.5Z), one plant, garden of Rose Cottage, M. McC. Webster, 1972,
introduced with plants from Blackhills (see next record). Blackhills House (NJ/2.5U), woodland
policies, M. McC. Webster, 1963 (Webster 1978).

*V.c. H9, Co. Clare: Ballyvaughan (M/2.0), outside P. O’Kelly’s house in a field and adjacent

hedge, 1987.
*V.c. H21, Co. Dublin: Malahide Castle (O/1.3), in the demesne, 1980, J. G. D. Lamb (Lamb 1983).
Rathfarnham (O/1.3), Marley Park, 1981, P. J. Jackson (Jackson 1983) (forma ptarmicifolia).

CURRENT STATUS

Cardamine bulbifera has thus been recorded in about 200 native sites; it is probably still present in

about 155 sites, has been lost from about 30 sites, and about 17 are untraceable or unchecked. Most-

of the native sites have disappeared due to woodland clearance, but replanting, especially with
conifers, is currently the main threat to the remaining populations. It has also been recorded from
about 45 introduced sites of which it is still present in 27.

ACKNOWLEDGMENTS

We wish to thank Margret Baecker for help with translation, Peter Flint for checking invertebrate
nomenclature, the Nature Conservancy Council for providing the Ancient Woodland Inventories,

all those who have provided information: J. R. Akeroyd, D. C. Bloomfield, Lady A. B. M. Brewis, |

Mrs M. Briggs, I. Brown, Mrs L. B. Burt, R. M. Burton, P. Casselden, Mrs P. Donovan, D. P. Earl,
J. Edelsten, P. J. Ellison, R. Fitter, B. R. Fowler, Miss M. B. Fuller, Mrs E. Gibb, P. Green, P. C.
Hall, P. A. Harmes, W. Havers, D. Hughes, S. L. Jury, T. J. James, R. Kaye, A. G. Knapp, D. H.
Kent, Miss R. G. Laidlaw, J. Lambert, A. C. Leslie, L. A. & Mrs P. D. Livermore, R. T. Mabey,
Mrs M. E. R. Martin, H. W. Matcham, R. Maycock, D. McClintock, Mrs E. M. J. Macphie, T. F.

Medd, E. C. Nelson, Ms E. M. Nic Lughadha, Mrs P. Nicholson, E. G. Philp, F. R. Philps, C. I.

Pogson, A. L. Primavesi, R. D. Randall, Miss E. J. Rich, F. Rose, G. Salisbury, Ms M. E. Scruby,
Miss M. R. Shaw, Mrs C. Smith, C. J. Smith, Mrs J. E. Smith, Mrs A. H. Sommerville, A. McG.
Stirling, N. J. H. Sturt, R. Thomas, W. H. Tucker, W. Turner, Mrs M. Vincent-Smith, Mr & Mrs C.
Webb, Mrs I. Weston, Mrs E. Wiltshire and M. G. Young; and the Keepers of BEL, BM, DBN, K,
OXF, LANC, RNG, TCD and YRK for access to or loan of specimens.

A.J.S. would like to thank the Botanical Research Fund and Buckinghamshire College of Higher
Education for help with travelling expenses.

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SS ee ee ee ee

— at

{i et > ee > ae ed ee

CARDAMINE BULBIFERA IN THE BRITISH ISLES 245

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Lams, J. G. D. (1983). Cardamine bulbifera in Co. Dublin. Ir. Nat. J. 21: 136.

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TOWNSEND, F. (1904). Flora of Hampshire. London.

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| WHITEBREAD, A., Barton, J. & Hutron, D. (1989a). East Sussex inventory of ancient woodland. Nature

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(Accepted October 1992)

Watsonia, 19, 247-257 (1993) 247

Orobanche reticulata Wallr. populations in Yorkshire
(north-east England)

Medio Y¥e-FOLEY

Division of Biological Sciences, University of Lancaster, Lancaster, LAI 4YQ

ABSTRACT

In the British Isles, Orobanche reticulata Wallr. (Orobanchaceae) has only been recorded with certainty from
Yorkshire. Its distribution and abundance within this area are reviewed, and the circumstances surrounding its
original discovery and those of other early records are described. The plant, a protected species, has been
recorded from at least 68 separate populations in ten 10-km squares, and 52 of these populations are extant,
although many are very small; it is particularly frequent in two separate but limited riverside areas.

INTRODUCTION

Orobanche reticulata Wallr. (Orobanchaceae) occurs in the British Isles as a root parasite of thistles,
especially Cirsium species (Asteraceae). All confirmed records are from Yorkshire (v.cc. 61, 62, 64,
65) where it occurs mainly, but not exclusively, on the magnesian limestone. In Europe its principal
area of distribution is in the alpine region but there are also many outlying, isolated localities in
northern Europe, including Scandinavia, the Baltic states, Holland, and north Germany (Hegi
1965; Chater & Webb 1972; Meusel et al. 1978). The plant also occurs in Russia, the Balkans, North
Africa, and extends eastwards into western Asia.

Its restriction to such areas of Yorkshire is unique amongst British native plants, and perhaps
because of this, doubts have been raised about its status (e.g. Pugsley 1926). Roman roads and
settlements occur near some of the earliest known localities and so an early introduction is a
possibility. However, when its British distribution is considered in the context of its overall
European pattern, where similar fragmented occurrences are frequent, especially near the limits of
its range, it seems most likely that the plant is genuinely native in Britain. Variability witnessed in
some populations (Foley 1992) supports this view, whilst an apparent reliance on a limited host
range suggests a persistent, isolated group of native populations.

O. reticulata is confirmed from 68 separate populations in ten 10-km squares, mainly in central
Yorkshire (v.c. 64) where there are two principal areas of distribution: between Leeds and
Wetherby and near Ripon. It also occurs at an isolated locality in v.c. 61, as well as on the extreme
south-western borders of v.c. 62 and v.c. 65, but has not been recorded from v.c. 63. In the few large
populations, the number of plants varies from one year to the next, but many populations are very
small, often with less than ten flowering plants annually. Owing to its rarity and very limited
distribution, it is a scheduled species protected under the Wildlife and Countryside Act, 1981 and
has been given a Red Data Book rating of 11E — Endangered (Perring & Farrell 1983).
Unfortunately, the host species are a great nuisance to farmers and landowners, and consequently
are frequently destroyed. O. reticulata therefore comes under an additional threat to that from
which other plants suffer.

In the field, confusion with other Orobanche species is unlikely although this may occur with
robust O. minor Sm. which occasionally shares the same host. Nevertheless, O. reticulata is quite
distinctive, being tall (up to c. 70 cm) and robust, with a fairly dense inflorescence, especially in the
upper part, and with cream-coloured corollas marked distally with blue-purple glands. The corolla
is cylindrical-campanulate and has a curved dorsal line which is especially characteristic amongst
British broomrapes. The stigma lobes are mauve-purple and the filaments glabrous to sparsely hairy
proximally, but can be somewhat glandular above. Even as a herbarium specimen, the plant is still

248 MJ. Y. FOLEY

distinctive, particularly with regard to its corolla shape, and in this state confusion is really only
likely to occur with O. rapum-genistae Thuill., or with O. elatior Sutton.

Many infraspecific taxa have been described from populations in continental Europe, and these
are summarised by Beck-Mannagetta (1930) and Gilli (1966), although none was recognised by
Chater & Webb (1972). British plants have been referred to O. reticulata var. procera (Koch) Beck,
which was differentiated from other varieties by its shorter, curved (not nearly straight) corollas.
However, further work may reveal that var. procera cannot be maintained even at varietal level.
Colour variants and forms of O. reticulata are much less frequent in Yorkshire than in Europe, but
some records have been noted recently (Jones 1989; Foley 1992).

In Britain, typical habitats are those of its hosts, rough pastures, road verges, and especially river
banks, river flood plains and associated light scrub. Cirsium arvense (L.) Scop. appears to be the
most frequent host, but C. vulgare (Savi) Ten., C. eriophorum (L.) Scop., C. heterophyllum (L.)
Hill and Carduus nutans L. are recorded, and also occasionally Cirsium palustre (L.) Scop. (e.g.
1932, Vachell, NAW). Scabiosa, Knautia (Dipsacaceae) and Cistus (Cistaceae) species are also said
to be parasitised in Europe. As with other Orobanche species, some claims for host can be
erroneous, since actual attachment to the roots is difficult to establish in the field. O. reticulata
seems to be at least partially perennial in Britain, flowering in early July, with the dead flower spikes
persisting for several months afterwards.

The account given below is based upon field work carried out over several years during which all
extant populations were visited by the author, many on several occasions. Estimates of plant
numbers were made and the various habitats assessed. Many herbarium specimens have also been
examined and if necessary re-determined by the author or by F. J. Rumsey. Relevant archival
records have also been consulted.

THE ORIGINAL DISCOVERY AS A BRITISH PLANT

In August 1907 H. E. Craven, a pharmaceutical chemist of Roundhay, Leeds, found an Orobanche
at Hetchell Crags, south-west of Wetherby, which he was unable to name. It appeared to be
parasitic on Cirsium eriophorum and was closest to O. minor. The following year, on seeing a letter
from G. C. Druce in the Pharmaceutical Journal in which he referred to the Botanical Exchange
Club, Craven wrote to Druce, asking for further details and enquiring whether membership of the
Club would enable him to get doubtful plants named as he had a few puzzling plants, one of which
was an Orobanche. Craven offered to send Druce some fresh specimens and this he did on 31 July
1908, having kept two plants in water, which had apparently become rather shrivelled. He also
offered to send a spike pickled in formalin, as well as duplicates from his previous year’s (1907)
collection, for Druce to forward to Beck-Mannagetta of Prague, the authority on European
Orobanche. A detailed description of the fresh plant and some duplicates were sent to Druce in
August 1908, and Druce exhibited the Orobanche at the Linnean Society that autumn, convinced
that it was not a known British species. During the summer of 1908, Craven also sent plants to E. M.
Holmes for naming. Holmes agreed that it was not previously known in Britain but suggested it
might be a hybrid; he also showed a specimen to the herbarium curator at Kew who was unable to
identify the plant. However, some months later, apologising for the delay due to illness, Beck-
Mannagetta replied to Druce (9 January 1909) naming the plant Orobanche reticulata f. procera
(Koch) Beck (= O. procera Koch). An original specimen sent to Beck-Mannagetta is in his
herbarium (PRC) and Beck-Mannagetta’s letter is appended to a sheet in Druce’s herbarium (OXF)
on which the latter mounted a specimen described “‘First as British” — presumably one of Craven’s
original specimens. Other early collections by Craven are in BM and MANCH. During the summer
following Beck-Mannagetta’s determination of the Hetchell plants as O. reticulata, Craven (17 July
1909) despatched most of his remaining duplicates for Druce’s retention, along with a map and a
detailed sketch of the habitat at Hetchell stating that the Orobanche grew there and nowhere else.
He also remarked that the tenant farmer had mown down all the thistles during the previous
summer (1908) and that he was concerned for the future of the plant. (This point is also made in
subsequent annotations to another sheet (1909, Lees & Pickard, YRK).) ~

Another Yorkshire botanist involved in the early recognition of the plant in Britain was F. A.
Lees. On receiving Beck-Mannagetta’s determination from Druce, Craven informed Lees and

OROBANCHE RETICULATA IN NORTH-EAST ENGLAND 249

presumably sent him a specimen, since one is preserved in Lees’ herbarium (July 1908, Craven,
BM); on the label, in Lees’ handwriting is the comment ““Orobanche procera, Koch (teste Dr Beck,
per Druce ad Craven 12.1.09)”. Also attached to the sheet are three photographs of the plant at
Hetchell Crags taken by J. H. Gough on 19 July 1913. On 29 July 1909, Lees, with J. F. Pickard,
went to see the plant at Hetchell, and one of the three specimens which they collected is preserved
(29 July 1909, Lees & Pickard, YRK). Writing to Druce on the following day, Lees commented that
the parasitised Cirsium eriophorum appeared to be very unhealthy, and that he had established the
connection from the Orobanche to the roots of the thistle. In this same letter Lees asked Druce to
publish a description of the plant which he subsequently did (Druce 1909a), and also offered to send
him specimens. Despite this, and Craven’s offer of a map on 17 July, Druce, along with A. H.
Evans, had already visited the site the previous month since a specimen exists which he collected at
the time (30 June 1909, Druce, OXF). This is added to a sheet of material collected by Craven in
1908, although Druce (1909a) stated that his visit was in July. From his correspondence with Druce,
it appears that Lees was the first person to raise the suggestion that the plant could be a Roman
introduction. He was aware that close to Hetchell Crags is the Pompocali, where the Romans had
their summer camp, and that there were also Roman roads and earthworks nearby.

Whilst the credit for the discovery of O. reticulata in Britain (and at Hetchell Crags) goes to
Craven, this is perhaps not strictly correct. The plant was first found there on 16 August 1902 by J. F.
Pickard (Lees’ companion in July 1909) but the specimen which he collected was erroneously
identified as O. elatior by Pickard and Lees (see notes on herbarium sheets: 1909, Pickard, BM;
1909, Lees & Pickard, YRK; 1910, Pickard, BM), and considered to be ‘“‘O. major” by Lees (quoted
by Cheetham & Sledge (1941)). Shortly afterwards, Pickard was away on business for two years (see
annotation, 1909, Lees & Pickard, YRK) and presumably did not question this identification
further. Craven subsequently re-found the plant at Hetchell, and not satisfied with these diagnoses
(Lees 1909), had it re-determined as described above. Pickard’s 1902 specimen has so far not been
traced. Although evidence points to the contrary, Druce appears to have indicated that the first
correct determination of the plant was his own, given directly on receipt of Craven’s original
specimens (Druce 1909a, 1911). However, in another statement, Druce (1909b) stated only that he
was convinced it was not a British plant, and had therefore sent a specimen to Beck-Mannagetta for
identification (see also Druce’s letter in herb. Beck (PRC)). Beck-Mannagetta’s reply to Druce
(9 January 1909) reads as a determination rather than a confirmation, and that is probably the case.

Shortly after Craven’s letter to Druce on 17 July 1909, it was thought that the Orobanche had been
exterminated at Hetchell. This resulted from a report in the Yorkshire Weekly Post of an address
given by J. G. Wilkinson, who was blind, to a Leeds Naturalist Club meeting. To disprove this, on 25
August 1909, Pickard and Wilkinson went to the place where Pickard saw the plant in 1902 (and also
in July 1909 with Lees) and found two in flower and eleven in fruit. Pickard stated that Wilkinson
considered that this locality was a different one “higher up on the slope and further on than
Craven’s’’, and Pickard was of the opinion that Craven’s population had been exterminated. Even
though Wilkinson was blind, he was considered reliable, and it is likely that there had been two
separate populations, with Craven’s being lost shortly after discovery. Examination of the general
area around the present site at Hetchell, together with the sketch which Craven sent to Druce,
contemporary notes on the flora, and Druce’s description (1909a) of Craven’s site — ‘‘a grassy slope
with bushes” — all support this view. Ironically, immediately after his visit in 1909, Druce had
received assurances from the landlord at Hetchell, that the thistles would be preserved.

OTHER EARLY RECORDS

Although the acceptance of O. reticulata as a British plant dates from the Pickard/Craven find,
specimens exist from other localities well before this date. These were, however, erroneously
referred to other Orobanche species. In the Leeds/Wetherby area a specimen was collected in 1868
from Roundhay Lime Hills (Lees & Abbott, CMM); it was identified as O. major (= O. rapum-
genistae) and thought to be growing on whin (Ulex europaeus), but is undoubtedly O. reticulata.
Roundhay is only a few kms from Hetchell Crags. Shortly after the Hetchell discovery, the plant was
also recognised from the large and already known broomrape colony at Hook Moor, Aberford (a
few kms south-east of Hetchell) whence it had variously been recorded as O. major and O. minor.

250 M. J. Y. FOLEY

Several specimens exist from here which are thus mistakenly labelled, with one collected as O.
major as early as 1878 (Webster, CMM). Despite assertions about the host for this specimen — ‘“‘One
root especially was only one or two inches from the gnarled stem of Ulex being far advanced to
ripeness. I did not trace any actually to the root of Ulex, altho’ I have no doubt” — and another
identified as O. minor “on Trifolium’ (1883, Williamson, YRK), both are nevertheless O.
reticulata. This illustrates the difficulty in being positive about which plant is actually parasitised.
The first contemporarily confirmed record at Hook Moor may be that of F. Ashwell (1916, LES) and
there is also a specimen from the previous year (1915, Cockerline, LES), but which may not have
had contemporary confirmation.

Another early record for O. reticulata is from Linton Common (again quite near Hetchell Crags)
where it was described by the finder as O. major. The date on the sheet (Pickard, CMM) is unclear,
being either 1901 or 1907. By 1907, Pickard might have been wary of this plant, following his
involvement at Hitchell in 1902 with what he thought was O.elatior, and so 1901 is perhaps the more
likely date. There is also a later annotation to a herbarium sheet (1909, Lees & Pickard, YRK)
stating that Pickard had also found and pressed the plant at Linton Common, where it was rare, but
that it was still there in 1936.

A second main centre of distribution is in the Ripon area of v.c. 64. In contrast to the Leeds/
Wetherby localities for which many early records can be confirmed by herbarium specimens, there is
little similar from here. The only specimen of O. reticulata so far traced which approximates to the
date of the Pickard/Craven find at Hetchell is one described as O. major “‘on gorse?’’, collected by
C. A. Cheetham on 18 July 1903 (C.A.C., BM). Unfortunately the label simply states ‘““Ripon”’ with
no further details of the locality. This specimen is erroneously referred to O. rapum-genistae by Lees
(Cheetham & Sledge 1941). It was not until 1939 that a localised Ripon area specimen was preserved
— from Bridge Hewick (Rob, YKN), although possible evidence for the plant’s early presence in the
area can be gained from the old Floras. In the Ripon area, nearly all modern records for O. reticulata
have been from riverside habitats, and the plant is now especially frequent there along an 8 km
stretch of the Ure, where at least 38 separate populations are extant. This lends credence to the
suspicion that some of the nineteenth century riverside Flora localities for other Orobanche species
in this area were misidentifications for O. reticulata. Those given for O. minor near Nunwick, at
Sleningford, Howe-upon-Swale, Ure Banks, Burnistone, and Bishopton (Baker & Nowell 1854;
Baker 1863; Slater 1883/4; Lees 1888), and for O. rapum-genistae below Tanfield (Cheetham &
Sledge 1941) are mostly close to extant O. reticulata populations, and may well have been mistaken
for that plant. Misidentifications for O. reticulata have already been shown to occur for some

nineteenth century specimens collected elsewhere as O. major and O. minor. Flora records from |

Roundhay and from Lotherton Moor (Hook Moor) both given by Lees (1888) must, in fact, have
been O. reticulata, and it is very likely that the same applied near Ripon.

Unfortunately it is rare for specimens of Orobanche to have been preserved from localities near
Ripon; ironically one of the few traced is undated (but prior to 1922) and was collected close to the
Ure between Ripon and Hutton Conyers (Lees, CMM). Lees finally determined this specimen as O.
reticulata but it is in fact O. elatior. An isolated occurrence lying to the east of the main area of
distribution was at Backhouse’s Nurseries, York (1886, Backhouse, K) where it was found in a
garden on Cirsium heterophyllum, presumably a casual occurrence. This is probably the first correct
identification of the host plant in Britain.

The first record from south-east Yorkshire (v.c. 61) was made in 1953 at North Grimston (Sledge
1954), at which site the plant is extant. In the same year it was also found at a nearby quarry (W. A.
Sledge, pers. comm.), but was not seen there again until 1992 during this survey; it is not known
elsewhere in v.c. 61. There are apparently only three instances of the plant being definitely recorded
from v.c. 62, and two of these are rather obscure. The first is an 1852 specimen (the earliest so far
traced from Yorkshire) of atypical O. reticulata collected from near Thirsk as O. elatior (Fowler,
OXF). Interestingly, there is also a collection (two sheets as O. minor) made in the previous year
from Sowerby, also near Thirsk (Baker, 1851, herb. Boswell-Syme, BM); the specimens are
intermediate between O. reticulata and O. minor and may possibly represent the hybrid. This record
deserves further investigation since natural hybrids of O. reticulata are not recorded in Britain
although they have recently been deliberately cultivated (Jones 1989). Whilst Fowler’s 1852 plant is
distinct from those collected by Baker, it is quite possible that they could be from the same
population. (Dead flower spikes, similar to the Sowerby plants, and again possibly of hybrid origin,

OROBANCHE RETICULATA IN NORTH-EAST ENGLAND 251

were seen near the Ure in 1992.) The second vice-county collection of O. reticulata is slightly more
recent and is typical, although labelled O. minor (1881, Webster, BM) but is unfortunately
unlocalised simply as ‘““N.E. Yorkshire” (see also Bennett 1917). Thereafter, it was not until 1984
that the plant was found in a riverside habitat in the extreme south-west of the vice-county, where a
flourishing colony survives. Before about 1980 there were apparently no records from v.c. 65, but
nine populations are now known, all on river margins, where only the past vagaries of the river
channel have resulted in their inclusion within that vice-county.

Outside Yorkshire, there are several dubious or erroneous records for O. reticulata. One which
may be significant is that given as O. major from Nottinghamshire (1835, Cooper, in herb. H. C.
Watson, K). The magnesian limestone does extend southwards into that county, but some of
Cooper’s specimens are considered to be of dubious provenance (Rumsey & Jury 1991) and this one
would benefit from independent verification. A record from Cheshire (v.c. 58) (Druce 1918; Perring
& Walters 1962; Perring & Farrell 1983) is erroneous since the specimen is O. minor (1918,
Dallman, OXF), as is an earlier one from Guildford (v.c. 17) quoted by Druce (1918) (1916,
Kennedy, OXF). Another error is that for Falmouth Dock (v.c. 1) in 1917 which again was O. minor
(F. J. Rumsey, pers. comm.). A record for var. pallidiflora (Wimm. & Grab.) Beck based on a
specimen collected from Brecon (v.c. 42) in 1905 (Knight, BM) has also been shown to be an error
(Pugsley 1940; Graham 1957; Rumsey & Jury 1991); a duplicate from this collection is in PRC.
Reports from E. Norfolk (v.c. 27) (Jones 1989) including one of an Orobanche parasitising Cirsium
vulgare are thought to be errors for other species (A. L. Bull, pers. comm.).

DISTRIBUTION AND HABITAT

The magnesian limestone formation occurs as a narrow band running north-south from Durham to
Nottinghamshire. In the area of distribution of O. reticulata it is rarely more than 10 km wide, and is
crossed in a roughly eastwards direction by the rivers Swale, Ure, Nidd, Wharfe and Aire.
Magnesian limestone is well-known for :ts species-rich flora.

10-km squares and tetrads (2 xX 2-km squares) for which there are confirmed records for O.
reticulata are mapped in Fig. 1 and Fig. 2 respectively, and fall within an area approximately 50 km
square. The two principal centres of distribution in the vicinity of Leeds/Wetherby and Ripon, the
disposition of the magnesian limestone formation and the principal rivers of the area, can all be
clearly seen.

Following the recognition of O. reticulata as a British plant there was a lull in the discovery of new
localities until the 1930s find by the river at Bridge Hewick, Ripon. Since then, other riverside ones
have been found some distance upstream from this site, many of which survive, and the plant is
particularly frequent in that area. Downstream also, new localities have occasionally come to light,
including one on the Ouse (formed by the confluence of the Ure and the Swale) found in 1984, the
first confirmed modern record for v.c. 62; there are, however, no definite records near the Swale
itself. Apart from a small population just south of Ripon and away from the Ure, all other post-1930
colonies in this northern (Ripon) area are on river banks or flood plains, and it seems clear that
rivers play an important role in dispersing seed. Also, plants often occur close to river banks and
eventually may be dislodged by water erosion. It is quite possible that these together with their host
will sometimes become established elsewhere. Favoured habitats for O. reticulata are near the
upper flood limit by the river bank, and especially the flat pasture borders nearby; occasionally they
grow some distance from the river but in places where winter flooding occurs. These are also the
- areas where the host species are plentiful.

In the southern (Leeds/Wetherby) area most newly-found localities are again in similar habitats
on the banks of the Wharfe, where propagation or dispersal has presumably occurred in a similar
manner. Population sizes can vary, but some appear to persist for many years with just a few plants
annually. The Linton Common locality is such an example. Here, even at the beginning of this
century only a few plants were recorded, and the same is the case today. Similarly at Hetchell Crags
where the plants occur on the magnesian overlay of the gritstone substrate and well away from the
river, numbers have apparently never been high. Cheetham & Sledge (1941) quoted a contempor-
ary account of F. A. Lees in which he stated that an increase in thistles had been observed at
Hetchell Crags between 1870 and 1879 but that he was certain that the broomrape was not present

252 M. J. Y. FOLEY

Ficure 1. The British distribution of O. reticulata shown on a 10-km square basis for which there are confirmed
records: @ 1991 onwards; © 1901-1990; x up to 1900.

during that time; this suggests a relatively recent colonisation just prior to its discovery there in
1902. During the 1920s about 20 plants were known at Langwith Scrub (W. A. Sledge, pers. comm.)
and today similarly small numbers still persist close by.

At Hook Moor, the classic site was unfortunately ploughed out about 1940, but surprisingly, and
despite reservations expressed at the time (Sledge 1942), what is still the largest British population
occurs on road embankments within the vicinity. Earth from the former site may have been
removed and used in construction of the present embankments and thus led to the plant’s presence
there, although natural recolonisation cannot be ruled out. Being on steep slopes, this is very well
drained and is also multi-aspect. It is rather different from the moister riverside localities favoured
elsewhere, and as the plant thrives here, it suggests that it may be a preferred habitat to the more
opportunistic riverside ones. Close to this re-colonised area, there is at least one, more natural, yet
very small population on an arable field border, probably a remnant of the former pre-1940 colony.

OROBANCHE RETICULATA IN NORTH-EAST ENGLAND 253

40 50

Ficure 2. The British distribution of O. reticulata showing tetrads from which there are confirmed records: @
1991 onwards; © 1901-1990; x up to 1900; the major river pattern and the disposition of the magnesian
limestone (hatched) are also indicated.

Roundhay limehills are mainly commercially developed nowadays and the plant is presumably
long extinct. Craven, the original discoverer in Britain, lived at Roundhay at the turn of the century
and would presumably have recorded it if he had known it there, and so it had perhaps gone even in
his day. Nothing is known of the plant’s habitat or frequency there, but Roundhay and Hook Moor
are both well away from major rivers, and could be remnants of an earlier area of distribution, from
which it has subsequently expanded.

All the localities mentioned occur on, or just eastwards of, the magnesian limestone formation.
Those to the east of it are usually close to major rivers and their occurrence there is likely to be a
result of downstream dispersal by water into habitats rich in alluvial deposits. No records for O.
reticulata have been traced to the west (upstream) of the magnesian limestone, and this reinforces
the argument for the downstream river dispersal through and away from this formation. However, a
_ magnesian limestone soil is not an essential requirement as shown by the plant’s occurrence in v.c.
61 on the chalk at North Grimston, where it formerly thrived (W. A. Sledge, pers. comm.) but is
now much less abundant. This locality is separated from all others and distant from any rivers, and
its presence there could possibly be from accidental introduction, since it has only been known since
1953.

Within the last 60 years or so, O. reticulata appears to have become more widely distributed in
Yorkshire than formerly, but whether this is due to a genuine expansion of range or results from
being previously overlooked, is uncertain. Some populations are highly stable, appearing on the
same area of thistle year after year, but others are more dynamic. These latter may change in size
and spatial distribution, apparently dying out (at least temporarily) in one part of the population,
only to appear a short distance away. Whether this depends on the vigour of the host or on some

254 M. J. Y. FOLEY

other local factor, is not clear. Observations in the field indicate that it is not the most dense stands
or the most robust thistles which are parasitised, but often smaller plants in scattered groups,
although it is not clear whether this state is caused by, rather than the cause of the parasite’s
distribution. However, even in the most favoured localities O. reticulata can remain highly
localised, with vast numbers of the host plant remaining un-parasitised, and it is assumed that
critical ecological factors control its ultimate viability.

SUMMARY

Of the 68 recorded populations of O. reticulata, at least 52 survive. The majority are on the
magnesian limestone formation and whilst this might not be an essential ecological requirement,
calcareous soils are favoured. Many populations have been found recently, but it is uncertain
whether these are newly-formed, or are much older and had been overlooked or even misidentified.
Many are small in numbers, but this is no indication of length of colonisation, since some colonies
are known to have persisted for over 70 years whilst consisting of only a few plants.

Of the extant populations, more than 85% are within the influence of river flood water and
dispersal and population formation by such means must clearly take place. Interestingly, these are
mostly restricted to two short reaches of the Ure and Wharfe and it 1s difficult to explain the absence
of similar records from the Nidd, which crosses the magnesian limestone between these two rivers.
Maybe a population has never developed sufficiently close to the Nidd to allow other riverside ones
to become established by water-borne means? Two other Yorkshire rivers which cross the
magnesian limestone also lack confirmed records in their vicinity, but the Swale does so over a very
narrow exposure and well to the north of the plant’s restricted area of distribution, whilst the Aire
although closer, is heavily industrialised. It is interesting that Potamogeton X suecicus K. Richter is
apparently also restricted in England (excluding the River Tweed on the Scottish border) to largely
similar reaches of the Ure and Wharfe, and is again not recorded from other rivers within the area.
New populations of O. reticulata will doubtless continue to form in favoured riverside localities but
colonisation elsewhere is likely to be rare, and if it occurs off the limestone, may originate from
inadvertent introduction.

The main threats to its survival result from the deliberate destruction of its hosts, from adverse
agricultural practices, and through engineering operations such as road construction. Some
populations may be lost when river banks are eroded, whilst gravel extraction is yet another threat.
Nevertheless it appears to be relatively resilient, occasionally reappearing at sites from which it was
thought to be extinct. There is much scope for investigation of the autecology and population
dynamics of O. reticulata.

RECORDED LOCALITIES AND RELEVANT HERBARIUM SPECIMENS

Whilst it is recognised that the number of flowering spikes can fluctuate annually and that two or
more may sometimes represent just a single plant, an attempt has been made to estimate a typical
size for each population. This is indicated using the following notation: A = 1—10 plants; B = 11-50;
C = 51-200; D = 201-1000; E = 1000+; X = probably extinct. Even if in fairly close proximity to
others, populations are considered to be separate if they are clearly observed as such in the field
even when consisting of just one or two plants. All populations shown as extant (A—E) have been
checked since 1991.

Localities which are not on the magnesian limestone formation are indicated by (*); those well
away from the influence of river flood water by (#).

Being a protected species, precise details of the populations are not given here, but have been
deposited at English Nature, York, and the Biological Records Centre, Monks Wood.

Herbarium specimens quoted earlier in the text are not usually repeated here, but their existence
is indicated by the code (Hb) inserted after the locality name. Others of relevance are listed after
each locality entry and although not exhaustive, they represent the majority of the more interesting,
historical specimens held in major national, or locally important herbaria. The fact that O. reticulata
is a Red Data Book plant is stressed and therefore further collections should not be made, but the

OROBANCHE RETICULATA IN NORTH-EAST ENGLAND 255

specimens quoted used for reference or examination. Where specimens have been redetermined
these are shown in this section as follows: (i) by F. J. Rumsey, (ii) by M. J. Y. Foley.

Notts., v.c. 56: J
Based upon the single unlocalised (Hb) record of a specimen of doubtful provenance.

Seo Vorks.. V.c.-ol:
SE/8.6, North Grimston (* #), found in 1953 (Sledge 1954) on chalk, well away from magnesian
limestone and river influence. Recently decreased, but still survives (B); herbarium specimens:
1953, Sledge (CMM, RNG). In 1953 also recorded at a nearby quarry but not seen again until 1992
(B). It is possible that there has been a recent introduction nearby at Wharram.

N.E. Yorks., v.c. 62:
SE/4.6, Linton-on-Ouse (*), first found in 1984, probably formed from a population further
upstream (B). There is also an unlocalised herbarium specimen from north-east Yorkshire (Hb)
collected as O. minor (ii).
SE/4.8, near Thirsk (* #) in 1852, based upon atypical specimens (Fowler, OXF). Two other sets
of specimens collected near here (at Sowerby) in 1851 (Baker, BM) may be of hybrid origin and
are possibly from the same population. Apparently not recorded since (X).

Mid-W. Yorks., v.c. 64:
SE/3.3 Roundhay Lime Hills (#), the early specimen as O. major (ii), (1868, Lees & Abbott,
CMM) is the only record traced. The site is now mostly developed and industrialised (X).
SE/3.4, East Keswick A, first noted in 1990 in a rough pasture (C). The following year several
interesting variants were observed (Foley 1992); East Keswick B, overgrown scrub, known since
about 1985 (B); Hetchell Crags (Hb), calcareous grassland above the gritstone escarpment, the
locality from which the first identified British O. reticulata specimens were collected by Craven in
1907 (B). It is probable that two populations formerly existed here, one becoming extinct in 1909
(X); herbarium specimens from Hetchell include: 1907, Craven (MANCH); 1908, Craven (OXF);
1908, Druce ex Craven (PRC); 1908, Craven (BM, CGE); 1908, Craven (OXF); 1909, Lees &
Pickard (CMM); 1909, Pickard (LES, CGE); 1910, Palmer (BM); 1913, Horrell (LES); 1919,
Cockerline (LES); 1951, Boniface (NMW); 1966, Hodgson (LANC); Whitwell, north bank of
river, first found in 1991 by P. P. Abbott (A); Langwith Scrub, known from this riverside locality
since the 1920s (W. A. Sledge, pers. comm.) and thought extinct, but a few plants were found
close by in 1991 (A). Lees’ (1888) record for O. major at Langwith Woods is very likely an error
for O. reticulata and there are also herbarium specimens probably from here: 1917, Horrell (OXF,
LES); Linton Common (Hb), recorded just after the beginning of this century, and still present in
small numbers (A).
SE/3.6, South Ripon (#), an area of limestone grassland, where occasional plants have been
found since 1989 including a single plant in 1992 (A); Roecliffe (*), a rough riverside pasture, first
recorded in 1969 in small numbers. Thought to be extinct, but four plants were found close by in
1992 (A).
SE/3.7, near Queen Mary’s Dubbs, five populations found in 1992, but a further one lost to
ploughing some years earlier (A, A, A, A, B, X); Ripon Parks, formerly several populations with
five still remaining on the flood plain of the Ure on Ministry of Defence land. Two others were lost
when the ground was ploughed in the early 1980s. Fasciated plants have been recorded from this
locality (A, A, A, A, C, X, X); Norton Conyers A, east bank of the Ure, ten populations mostly
found in 1992, one of which possibly contained hybrid piants (x O. minor) (A, A, A, A, A, A,B,
B, B, B); near Nunwick, four populations found recently (A, A, B, B); north of Ure Bank found
in 1992 (A); Bridge Hewick (Hb) where two separate populations flourished about 1940-1950 on
opposite banks of the Ure. Rob (1953) commented that the plant had decreased here, but there
were about 70 plants at one population in 1954 (H. J. M. Bowen, pers. comm.). Apparently
extinct at one of the old localities but four populations were found near the other in 1992 (A, A,

_ A,B, X). Herbarium specimens from Bridge Hewick include: 1952, Bangerter (BM); 1954, Rob
(YKN); 1954, Bowen (RNG). Other specimens only partially localised and probably from near
here are: Ripon, as O. major (i), 1903, C.A.C. (BM); near Ripon, 1956, Branson (E, RNG).
SE/4.3: Hook Moor (#) (Hb), one of the classic sites from which specimens can be found in most
major herbaria (often under the alternative locality names of Micklefield or Lotherton Moor),
one such early specimen dating from 1878. Formerly widéspread prior to the site being ploughed

256 M. J. Y. FOLEY

in about 1940. Since then good populations have developed on road embankments within the
vicinity, plants or seed possibly having been brought in with the soil during road construction, or
even formed by natural re-colonisation. Probably the strongest British population (D, possibly
E); there are two other nearby localities: a probable relict population on an arable field border.
(A), and another on a road embankment to the north (A). Three other records in the period
1952-1972 from other areas of Hook Moor (#) have not been recently confirmed (X, X, X).
Herbarium specimens include: as O. eu-minor (ii), 1881, Webster (BM); as O. minor (i), 1881,
Webster (E); 1892, Foggitt (BM); 1919, Sledge (CMM); 1922, Barnett (LES); 1922, Roper (LDS);
1929, Jones (LDS); 1930, Foggitt (BM); 1932, Vachell (NMW); 1932, Vachell & Knowling
(NMW); 1936, Sledge (LIV, RNG); 1936, Jackson (K); 1937, Lousley (LIV, RNG, K); 1937,
Woodhead (LANC); 1938, Chapple (OXF); 1939, Lousley (NMW); 1946, Lousley (E, RNG);
1946, Libbey (LTR); 1949, Raven (LTR); 1953, Frankland (three sheets) (LIV); 1988 (photo-
graphs only), Rumsey (RNG). There is an unlocalised specimen probably from here: near Leeds,
as O. rubra (i), 1922, Rogers (CGE).
SE/4.4, near Tadcaster, the sole record is based upon a semi-localised herbarium specimen
collected in 1935 (W.R.W., K) and now presumed extinct (X); near the Dunsforths, rumoured to
have been found here in the mid-1980s, but no definite records traced.
SE/5.5: The Nurseries near York (* #), based on Backhouse’s 1886 specimen (K). At this date,
Backhouse’s nurseries were in what are now the south-western suburbs of the city. Presumably a
casual introduction with no records since (X).

N. W. Yorks., v.c. 65:
SE/3.7, Badger Bank, five separate populations on the east bank of the Ure (A, A, A, A, B);
Norton Conyers B, east bank of the Ure, first found in 1988, including one broken, fasciated
plant, Norris & Lloyd-Evans (LES) (A); North Stainley, on a nature reserve on the west side of
the Ure. Recorded here in 1984, this population has apparently died out, but a second one
appeared close by in 1992 (A). There is a further population a short way downstream (A); Low
Batts, discovered here about 1980 (B).

ACKNOWLEDGMENTS

In the compilation of information for this paper, I am very grateful to many people for their help.
These include the herbarium curators at BM, CGE, CMM, E, HAMU, K, LANC, LDS, LES, LIV,
LTR, MANCH, NMW, OXF, PRC, RNG, YKN, YRK, as well as the following who have assisted
with details for some of the localities or with other information: P. P. Abbott, J. Deane, D. L.
Grant, D. E. Haythornthwaite, T. F. Medd, B. Molesworth, A. Norris, C. D. Preston, F. J.
Rumsey, C. Slater, W. A. Sledge, M. Thallon and S. Warwick. I am also grateful to M. S. Porter for
help in the field and to English Nature for sponsoring part of the work.

REFERENCES

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BENNETT, A. (1917). Orobanche reticulata Wallroth. Naturalist 42: 165.

CuaTer, A. O. & Wess, D. A. (1972). Orobanche L., in Tutin, T. G. et al., eds. Flora Europaea 3: 290.
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(Accepted January 1993)

Watsonia, 19, 259-263 (1993) 259

Skunkweed (Navarretia squarrosa (Eschsch.) Hook. & Arn.)
and other Polemoniaceae in Britain

E. J. CLEMENT
54 Anglesey Road, Gosport, Hants, PO12 2EQ
and
M. F. GARDNER and S. G. KNEES

Royal Botanic Garden, Edinburgh, EH3 5LR

ABSTRACT

The discovery of a large population of Skunkweed, Navarretia squarrosa (Eschsch.) Hook. & Arn. (Polemoni-
aceae) in Windsor Great Park, Berks. (v.c. 22) has prompted further investigation into this little known alien,
which is described, illustrated and discussed in conjunction with related species. A key is given to separate four
relevant genera.

INTRODUCTION

Skunkweed, Navarretia squarrosa (Eschsch.) Hook. & Arn. (Polemoniaceae), has a long history of
periodic occurrence in Britain but is not treated in recent Floras (Clapham et al. 1987; Stace 1991).
In early July 1990, M.F.G. was informed that a large population of N. squarrosa had been
discovered on private land within Windsor Great Park (v.c. 22, Berks.). On investigation it was
found that the colony of several thousand plants was thickly scattered on more or less bare sandy
ground which had until recently been a rubbish dump. This site had been sown the previous autumn
with a grass and wild-flower amenity mixture, the grass seed of which had originated from Oregon.
Germination of the sown mixture had been almost non-existent and instead the area was sparsely
covered by mainly mosses and several early colonising species such as Spergularia rubra (L.) J. & C.
Presl and Gnaphalium uliginosum L. It was interesting to note that grasses were almost absent. The
whole area was almost flat with a few slight hollows which presumably held water during the winter.
In June these were covered with the annual White Forget-me-not Plagiobothrys scouleri (Hook. &
Arn.) I. M. Johnst., a member of the Boraginaceae which is also a native of western North America
(Stace 1991).

In 1991 both N. squarrosa and P. scouleri reappeared in considerable numbers suggesting that
seed had been successfully set the previous year. However, long-term persistence is unlikely, as the
ground will probably become overgrown again. In mid-June only small seedlings of N. squarrosa
were present and on walking through these plants their foetid smell, resembling petrol or rotten
eggs, was very prominent.

DESCRIPTION OF NAVARRETIA SQUARROSA

The genus Navarretia (Polemoniaceae) contains 30 annual species of which 29 are native to western
N. America and one is native to Chile and Argentina (Mabberley 1987). Many species of Navarretia
look somewhat similar and the following description defines N. squarrosa.

Navarretia squarrosa (Eschsch.) Hook. & Arn., Bot. Beechey’s Voy. 8: 368 (1839). Fig. 1.
Synonyms: Hoitzia squarrosa Eschsch., Mém. Acad. Sci. St Pétersb. 10: 283 (1826); Gilia pungens

260 E. J. CLEMENT, M. F. GARDNER AND S. G. KNEES

No?
! RE
e

~W gh Ooms

NAVARRETIA SQUARROSA ca

Ficure 1. Navarretia squarrosa (Eschsch.) Hook. & Arn., Skunkweed.

Douglas ex Hook., Bot. Mag. 57: t. 2977 (1830); Gilia squarrosa (Eschsch.) Hook. & Arn., Bot.
Beechey’s Voy. 4: 151 (Oct. 1833); Aegochloa pungens (Douglas ex Hook.) Benth., Edward’s Bot.
Reg. 19: sub pl. 1622 (1 Oct. 1833); Navarretia pungens (Douglas ex Hook.) Hook., Fl. Bor. Am. 2:
75 (1838).

Erect annual 5—35 cm, with usually one main stem, rather stout and rigid, the lateral branches more
or less well developed producing a top-heavy plant narrowing to a slender base; all parts except
flowers glandular-viscid, with a strong, unpleasant odour (hence the vernacular name). Leaves
alternate, sessile, 1-5—3-5 x 1-2-5 cm, varying from bipinnate to pinnately and palmately dissected,

|
c
:

|
|
|
|
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—_——~

SKUNKWEED IN BRITAIN 261

the linear lobes spinescent. Flowers in capitate, bracteate clusters; calyx tube 7-10 mm, scarious
between the ribs, with unequal, subulate, spinescent lobes; corolla 7-13 mm, infundibuliform, blue
to whitish, barely exceeding the calyx; stamens 5, slightly unequal, inserted in corolla throat;
stigmas 3. Capsule ellipsoid, 3-locular, each loculus with 6-8 or more ovules. Seeds 0-6-0-8 mm,
dark brown, irregularly angled-ovoid, rugose. Native to western N. America, from Vancouver
Island in the north, south to California (Hitchcock et al. 1959).

Cantua pungens Torrey, Ann. Lyc. Nat. Hist. N. York 2: 221 (1828) was described on the basis of
a specimen collected by Edwin P. James in 1820, in the valley of the Loup Fork (Nebraska). This is
obviously not the same specimen as the type of Gilia pungens Douglas ex Hook. (collected by David
Douglas on the Multnahomack River, California); because the respective type descriptions of C.
pungens and G. pungens differ in several important points (e.g. C. pungens: flowers solitary,
axillary; G. pungens: flowers in capitate clusters), C. pungens is not considered to be synonymous
with Navarretia squarrosa, although Index Kewensis considered it to be so (R. R. Mill, pers.
comm.). The type locality of C. pungens is also far outside the geographical range of N. squarrosa.

Voucher specimens of Navarretia squarrosa and Plagiobothrys scouleri have been deposited in
RNG, E, herb. E.J.C. and herb. M.F.G. Specimens of N. squarrosa were also collected and
distributed by the University of Reading through the Société pour |’Echange des Plantes
Vasculaires de l Europe et du Bassin Méditerranéen. It is interesting to note that the oils within the
plant are volatile and pressed specimens soon totally lose their foetid odour. N. squarrosa seems to
be very poorly represented in the literature and in herbaria as a British adventive. For example,
there are no specimens held by BM, CGG and K.

HISTORY OF OCCURRENCE IN BRITAIN

1828-1830

The species was probably first cultivated in Britain in 1829 or 1830, when plants were raised by the
Horticultural Society of London from seed collected by David Douglas in North America. He
collected seed from plants growing on moist ground in mountain valleys near to the sources of the
Multnahomack river, one of the southern branches of the Columbia river in western North
America. Plants from this collection were subsequently illustrated in the Botanical Magazine, tab.
2977 (Hooker 1830) and although not garden worthy, references to Navarretia squarrosa have
persisted in horticultural literature to the present day (Huxley 1992).

1915-1933

The species was first collected as an alien by Miss I. M. Hayward in August 1915 under Ladhope
Bridge, Galashiels, Selkirks., v.c. 79, where it was growing on shingle beside Gala Water and had
presumably been introduced with wool shoddy (Druce 1916; Hayward & Druce 1919). It was
determined by A. Thellung as Gilia pungens Douglas ex Hook. & Arn. Some eight years later it was
collected from Sleaford, S. Lincs., v.c. 53, by Miss Landon where it was thought to have been
introduced with chicken corn (Druce 1924); again determined by Thellung, this time as G.
squarrosa Hook. & Arn. (OXF). The same year it was also collected from Hythe Quay, Colchester,
N. Essex, v.c. 19, where it was thought to have been introduced with malting refuse. Plants were still
being recorded from the same locality the following year (Brown 1930, OXF). The next recorded
occurrence was reported by Miss C. M. Rob from Topcliffe, near Thirsk, N. E. Yorks., v.c. 62,
where it came up in a newly made lawn. In this case the plant may have been introduced with sawn.
wood which originated from California and was used for constructing the doors of the house
(Pearsall 1934).

1978-1979

Nearly half a century passed before the species was seen again in Britain, this time as a weed in a
rose bed by Mrs M. Baecker at Carr Bank, Milnthorpe, Westmorland, v.c. 69, in 1978 and 1979
(Clement 1979, herb. E.J.C.).

262 E. J. CLEMENT, M. F. GARDNER AND S. G. KNEES

OCCURRENCE ELSEWHERE

Navarretia squarrosa has also been recorded as a bird seed alien in the Netherlands (Ooststroom &
Reichgelt 1963) and as a ruderal in Denmark (Hansen 1979). Outside Europe it is recorded in the
floras of New Zealand and Australia, having first been noted early in the century by Black
(Robertson 1957); it persists there to the present day (Toelken 1986; Webb et al. 1988).

OTHER POLEMONIACEAE IN BRITAIN

There is only one member of the Polemoniaceae native to Britain and Ireland. Jacob’s Ladder,
Polemonium caeruleum L., occurs in limestone grassland, on screes, on rock ledges and on the
borders of woodland in northern England and is reported (Stace 1991) as a sporadic garden escape
elsewhere. However, B. Wurzell (pers. comm.) warns that since a number of similar Polemonium
species, varieties and hybrids are grown in gardens, no wild plant of such provenance should be
attributed to P. caeruleum without careful verification. Stace (1991) also lists Phlox paniculata L..,
another North American species, as sporadically naturalised on rough and waste ground in
England. At the last published tally (Druce 1928), twelve alien Polemoniaceae were listed, the
genus Gilia contributing most taxa. None of these was included in the two more recent lists of British
plants (Dandy 1958; Kent 1992). Phlox drummondii Hook. and P. subulata L. have been observed
as adventives on urban wasteland (B. Wurzell, pers. comm.)

KEY TO THE GENERA OF POLEMONIACEAE OCCURRING IN BRITAIN

The following key may aid the identification of three of the commonest genera most likely to be
confused with Navarretia:

ft... Calyx green'and herbaceous throughout 2.3.20 .i00 5... eects oe eg ee eee 1. Polemonium
1. Calyx with prominent scarious or hyaline intervals between the green and more herbaceous

a) 9 1S) 2: | gee cain a Sr aa RRND Ke saleell UNS RUMI ARP Fi). MMMM, NEMO ME NN G50. dann omooaeeseoasuce oes 2
2. Staminal filaments very unequally inserted; leaves opposite ............... cece ceeeee eee 2. Phlox
2. Staminal filaments equally or almost equally inserted; leaves alternate ........................... 3
3. Calyx-lobes obviously unequal; leaf segments spine-tipped .................... petals 3. Navarretia
3. Calyx-lobes equal or nearly so; leaf-segments not spine-tipped .....................eee ee ee 4. Gilia

ACKNOWLEDGMENTS

The authors are grateful to the following for their assistance in the accumulation of information for

Mr M. C. Tebbitt and Dr P. F. Yeo. Thanks are also due to Dr J. R. Akeroyd and Dr R. R. Mill for
their helpful comments on earlier drafts of the paper and Mr B. Wurzell for providing further useful |
information on other alien Polemoniaceae. Mrs O. M. Stewart is especially acknowledged for
preparing the drawing.

this paper: Miss S. Andrews, Mrs M. C. Foster, Dr S. L. Jury, Miss S. K. Marner, Miss A. M. Paul, |

REFERENCES

Brown, G. C. (1930). The alien plants. of Essex. The Essex Naturalist 22: 41.

CLapHaM, A. R., Tutin, T. G. & Moore, D. M. (1987). Flora of the British Isles, 3rd ed. Cambridge.
CLEMENT, E. J. (1979). Adventive news 15. B.S.B.I. News 23: 12.

Danpy, J. E. (1958). List of British vascular plants. London.

Druce, G. C. (1916). Plant notes, etc., for 1915. Rep. botl Soc. Exch. Club Br. e 4: 203.

Druce, G. C. (1924). Plant notes, etc., for 1923. Rep. botl Soc. Exch. Club Br. Isl. 7: 46.

Druce, G. C. (1928). British plant list, 2nd ed. Arbroath.

SKUNKWEED IN BRITAIN 263

Haywarp, I. M. & Druce, G. C. (1919). The adventive flora of Tweedside. Arbroath.

Hansen, A. (1979). Floristiske meddelelser. Urt 4: 122-124.

Hitcucock, C. L., Cronquist, A., OwnBEY, M. & THompson, J. W. (1959). Vascular plants of the Pacific
Northwest, 4: Ericaceae through Campanulaceae. Seattle.

Hooker, W. J. (1830). Gilia pungens. Sharp-leaved Gilia. Curtis’s Botanical Magazine 57 (n.s. 4): plate 2977.

Huxtey, A. J. ed. (1992). Navarretia, The new Royal Horticultural Society Dictionary of Gardening 3: 296.
London.

Kent, D. H. (1992). List of vascular plants of the British Isles. London.

MaBBERLEY, D. J. (1987). The plant book. Cambridge.

OoststTRooM, S. J’ & REICHGELT, T. J. (1963). Aanwinsten voor de Nederlandse adventief-flora, 5. Gorteria 1:
141-143.

PEARSALL, W. H. (1934). New county and other records, 1933. Rep. botl Soc. Exch. Club Br. Isl. 10: 533.

RoseErtson, E. L. (1957). Polemoniaceae. Flora of South Australia 4: 707. Adelaide.

Stace, C. A. (1991). New Flora of the British Isles. Cambridge.

ToELKEN, H. R. (1986). Polemoniaceae. Flora of South Australia 3: 1133. Adelaide.

Wess, C. J., SYKES, W. R. & GARNOCK-JoNnES, P. J. (1988). Polemoniaceae. Flora of New Zealand 4: 957-961.
Christchurch.

(Accepted January 1993)

Waisonia, 19, 265—267 (1993) 265

Who was the author of Montbretia crocosmiiflora?

E. C. NELSON

National Botanic Gardens, Glasnevin, Dublin 9, Ireland

ABSTRACT

Montbretia crocosmiiflora, the basionym of Crocosmia X crocosmiiflora (Iridaceae) was published by Victor
Lemoine in The Garden, 1880, not by E. Morren; the correct citation of the current name is Crocosmia X
crocosmiifiora (Lemoine) N.E.Br.

INTRODUCTION

Montbretia crocosmiifiora is the basionym of the currently accepted name for the well-known
garden plant, Crocosmia X crocosmiiflora (Iridaceae), which has escaped and become widely
naturalized in coastal parts of the British Isles (Stace 1991; Nelson 1993). Although the generic
name Montbretia has long since been abandoned, it remains in everyday use as the vernacular name
especially for the naturalized plant. C. X crocosmiifiora is an artificial hybrid created by Victor
Lemoine of Nancy, France, who pollinated C. pottsii (Baker) N. E. Br. with pollen from C. aurea
(Hook.) Planch.; the seedlings first bloomed in August 1880.

For many years standard botanical accounts (e.g. de Vos 1984) credited publication of Montbretia
crocosmiiflora to C. J. Edouard Morren (1881), and the usual citation of the hybrid’s name was
Crocosmia X crocosmiiflora (Lemoine ex E. Morren) N. E. Br. (cf. de Vos 1984). Kostelijk (1984)
pointed out that Morren’s account, published in La Belgique Horticole 31 late in 1881 to accompany
plate 472, was predated by a note printed in the October 1881 number (118) of The Floral Magazine,
edited by Richard Dean, but Kostelijk (1984) and Wijnands (1986) omitted to note that the
September issue of the same periodical had an earlier account, reading as follows

**.. . Montbretia crocosmaefiora, a novelty sent by Mons. Lemoine, Nancy, France; much the

same in colour as M. Pottsii, but perhaps a little more yellow, and the flowers larger.”
Kostelijk (1984) proposed altering the citation to C. x crocosmiiflora (Lemoine ex Burb. & Dean)
N. E. Br.; this was noted by Wijnands (1986), and taken up by Trehane (1989), Stace (1991) and
Kent (1992) among others.

EARLY DESCRIPTIONS OF LEMOINE’S HYBRID

Kostelijk’s bibliographic search was not exhaustive. The following are even earlier accounts in
which the binomial Montbretia crocosmiiflora was used:

The Garden 21 August 1880 (p. 188)

~ “New Hybrid Montbretia. — Mons. V. Lemoine, of Nancy, sends us a new bulbous plant, which
he has obtained by fertilising Montbretia Pottsi . . . with Tritonia (Crocosma [sic]) aurea. The
progeny Mons. Lemoine proposes to name Montbretia crocosmaeflora. The flowers, he says, are
four or five times the size of M. Pottsi; and this successful cross he considers to be the starting
point for the production of a race of beautiful hardy varieties. The flowers sent are borne in the
same manner on the spike as those of M. Pottsi, but they are much larger and of a deeper colour.
The cross seems to be precisely intermediate between the two parents.”

The Garden 30 July 1881 (p. 125) :
“Montbretia crocosmaeflora, similar in every respect to M. Pottsi [sic] . . . but more robust in
habit, and having larger flower-spikes.”’

266 E. C. NELSON

The Gardener’s Chronicle 30 July 1881 (p. 153)
“Mr Barron also showed Montbretia crocosmaefiora [sic], one of Mons. Lemoine’s novelties,
much the same colour as M. Potsii [sic], perhaps a little more yellow, but the flowers larger — a
very good plant...”

The Garden 27 August 1881 (p. 203)
“MONTBRETIA CROCOSMAEFLORA - A very promising plant with orange and red
flowers, very bright, but somewhat withered owing to its journey from Nancy . .. From M.
Lemoine.”

DISCUSSION

The first quotation from The Garden (18: 188, dated 21 August 1880) must have been written very
shortly after the seedlings bloomed for the first time; it contains a diagnosis that is adequate under

the International Code of Botanical Nomenclature (Greuter et al. 1988; Art. 32.2) to validate the —

binomial Montbretia crocosmiiflora. It must be stressed that both the binomial and at least part of
the description are explicitly attributed to Lemoine, and the passage is written in the present tense.

But, by including the clause ““Mons. Lemoine proposes to name . . .”’, does the author perhaps fall
foul of Art. 34.1, that a name is “not validly published ... when it is merely proposed in
anticipation of . . . a particular circumscription’’?

The two subsequent notes, both published almost one year later on 30 July 1881, do not
contravene any articles of the current /.C.B.N. and thus the binomial was validly published as early
as 30 July 1881 — unless description in the issue of The Garden of 21 August 1880 is not ruled out.

Deciding which of the 30 July 1881 issues of the separate periodicals was the first published seems
a pointless exercise, but they predate The Floral Magazine notices by at least one month, so that the
protologue of Montbretia crocosmiiflora may be credited to one or other of these, or both. At this
period The Gardener’s Chronicle was edited by M. T. Masters, and The Garden by its founder,
William Robinson, and thus possible citations might include ‘Lemoine ex Masters’, and ‘Lemoine
ex W. Robinson’.

The publication of names in reports of horticultural shows is a matter that is not addressed by the
I.C.B.N. There is no reason to reject names, accompanied by diagnostic statements, when included
in such reports, except when they contravene particular articles of the code. Under the J.C. B.N.
(Art. 29, Berlin 1988), publication in non-scientific newspapers was forbidden after 1 January 1953;
thereby there is the implication that before that date publication of binomials in such periodicals is
valid. The descriptions of Montbretia crocosmiiflora quoted above appeared in reports of the Royal
Horticultural Society’s exhibition on 26 July 1881, and it is possible that London newspapers of 27
July contained reports in which there were descriptions — I have made no attempt to trace any such
reports. It is even possible that French periodicals and newspapers contained even earlier reports of
the first flowers in August 1880, for example. .

The consequences of this conundrum is that it is impossible unambiguously to assign Montbretia
crocosmiiflora to a single author. Both Masters and Robinson were describing the same plant,
having seen the same specimens on the same day, although it cannot be established that Masters or
Robinson personally wrote the news items concerned; they were the editors of the respective
periodicals, and one of their journalists could have contributed the show reports. Furthermore,
because of the real possibility that newspapers printed in August 1880 and July 1881 carried
descriptions and the binomial, it may be impossible to determine if the original place of publication
was really The Gardener’s Chronicle or The Garden.

CONCLUSIONS

Each of the quoted descriptions is of the same, indeed the original hybrid, so the best solution to this
conundrum is to accept that the name was first devised and used by Victor Lemoine, that it was
published validly in The Garden on 21 August 1880, and that because the clause “proposes to name”
is in the present tense this diagnosis need not be regarded as provisional and thus contrary to Art.
34.1. Thus the citation should read:

‘
Hl
ri
‘
it
ae

|
|
|

THE AUTHOR OF MONTBRETIA CROCOSMIIFLORA 267

Crocosmia X crocosmiiflora (Lemoine) N. E. Br., Transactions of the Royal Society of South Africa

20: 264 (1932).

Basionym: Montbretia crocosmiiflora Lemoine, The Garden 18: 188 (21 August 1880) [as
‘crocosmaeflora |; The Garden 20: 125 (30 July 1881); The Gardener's Chronicle 16 (n.s.): 153 (30
July 1881); The Garden 20: 203 (27 August 1881); The Floral Magazine no. 117 (September 1881);
The Floral Magazine no. 118, tab. 472 (October 1881); La Belgique Horticole 31: 229, tab. 14
(1881) [with formula ‘x Montbretia aureo-pottsi’’|.

TYPIFICATION OF MONTBRETIA CROCOSMIIFLORA

de Vos (1984) designated the illustration published in La Belgique Horticole 31 (tab. 14) as the
lectotype of Montbretia crocosmiifiora Lemoine ex Morren, but this is not acceptable (see I. C. B.N.
Art. 7) and must be rejected. In this instance the published illustration cannot be a lectotype but
could be selected as a neotype, as long as there are no herbarium specimens preserved of the original
materials received from Lemoine by Robinson in August 1880. However, better candidates for
selection as neotypes would be herbarium specimens prepared from the material displayed at the
Royal Horticultural Society in July 1881.

ORTHOGRAPHY

The earliest notes used either ‘crocosmiaeflora’ or ‘crocosmaeflora’. These are improperly formed
compounds; under the J.C. B.N. (Art. 73), the epithet should be corrected to ‘crocosmiiflora’, as is
standard practice.

ACKNOWLEDGMENT

My thanks are due to Dr R. K. Brummitt, for his helpful comments.

REFERENCES

DE Vos, M. (1984). The African genus Crocosmia Planchon. JI S. Afr. bot. 50: 463-502.

GREUTER, W. et al. (1988). International code of botanical nomenclature adopted by the Fourteenth International
Botanical Congress, Berlin, July-August 1987. K6nigstein.

KENT, D. H. (1992). List of vascular plants in the British Isles. London.

KosTELuK, P. J. (1984). Crocosmia in gardens. The Plantsman 5: 246-253

MorreNn, C. J. E. (1881). Notice sur le Montbretia crocosmiaeflora (hybrida) de M. V. Lemoine. X Montbretia
aureo-pottsi. La Belgique Horticole 31: 299-303, tab. 14.

NELson, E. C. (1993). Ergasiophygophytes in the British Isles — plants that jumped the garden wall, in ELus, R.
G. & Perry, A. R., eds. Plants wild and garden. Cardiff.

Stace, C. A. (1991). New Flora of the British Isles. Cambridge.

TREHANE, P. (1989). Index hortensis. Vol. 1, Perennials. Wimborne.

WunNanps, D. O. (1986). The correct citation of Montbretia crocosmiiflora. Bothalia 16 (1986): 51.

(Accepted February 1993)

Watsonia, 19, 269-278 (1993) 269

Short Notes

CONTRIBUTIONS TO A CYTOLOGICAL CATALOGUE OF THE BRITISH AND IRISH
FLORA, 3

This note continues our series presenting the results of a cytological survey of British and Irish
vascular plants (Hollingsworth et al. 1992). Here we report the chromosome numbers of 88 species
from 95 populations. A suffix ‘S’ denotes supernumerary chromosomes. Only one plant per
population was counted, except where stated. All counts were made on squashes of root-tips.
Voucher specimens have been deposited in LTR.

Ajuga reptans L., 2n = 32: Leics., v.c. 55, Swithland Wood, SK/53.12.

Alchemilla glaucescens Wallr., 2n = c. 110: Westmorland, v.c. 69, Crosby Gill, NY/61.11.

Alchemilla xanthochlora Rothm., 2n =c. 107: W. Lancs., v.c. 60, near Carnforth, SD/507.706.

Allium scorodoprasum L., 2n = 16: W. Lancs., v.c. 60, Potts Corner, SD/41.57.

Andromeda polifolia L., 2n = 48: Westmorland, v.c. 69, Meathop Moss, 3km N.E. of Lindale, SD/
44.81.

Apium inundatum (L.) Reichb.f., 2n = 22: Caerns., v.c. 49, Cilan, near Abersoch, SH/29.24.

Barbarea vulgaris R.Br., 2n= 16: W. Lancs., v.c. 60, River Lune, near Lancaster, SD/488.640.

Brassica napus L. subsp. oleifera (DC.) Metzger, 2n = 38: Dorset, v.c. 9, c. 1-5 km W. of Burton
Bradstock, SY/47.89.

Calystegia soldanella (L.) R.Br., 2n=22: Caerns., v.c. 49, Pwllheli, SH/37.34.

Cardamine amara L., 2n = 16: W. Lancs., v.c. 60, near Carnforth, SD/507.706.

Carex acutiformis Ehrh., 2n = 38: W. Lancs., v.c. 60, Lancaster University campus, SD/48.57.

Chrysosplenium oppositifolium L., 2n=42: W. Lancs., v.c. 60, near Millbeck footbridge, SD/
648.638.

Circaea lutetiana L., 2n=22: Dorset, v.c. 9, Kingcombe Meadows, c. 1:5 km N.E. of Toller
Porcorum, SY/55.99; W. Lancs., v.c. 60, near Millbeck footbridge, SD/648.638.

Crataegus monogyna Jacq. subsp. nordica Franco, 2n = 34: Caerns., v.c. 49, Penarwel valley, near
Llanbedrog, SH/32.32.

Crithmum maritimum L., 2n = 20: Dorset, v.c. 9, Weymouth, landward side of Fleet Lagoon, SY/
66.76.

Dactylorhiza purpurella (T. & T. A. Stephenson) So6, 2n = 80: W. Lancs., v.c. 60, SD/5.7.

Daphne laureola L., 2n = 18: Dorset, v.c. 9, Todber, ST/79.19.

Dryopteris filix-mas (L.) Schott, 2n = 164: W. Lancs., v.c. 60, near Yealand Conyers, SD/509.745.

Equisetum telmateia Ehrh., 2n = c. 216: Caerns., v.c. 49, track from Carreg to Porth Oer (Whistling
Sands), SH/16.29.

Eryngium maritimum L., 2n = 16: Caerns., v.c. 49, Pwilheli, south beach, SH/37.34.

Euonymus europaeus L., 2n = 32: Dorset, v.c. 9, Winterborne Kingston, SY/86.97.

Euphorbia peplus L., 2n = 16: Dorset, v.c. 9, Portland, Church Ope Cove, SY/69.70.

-Fraxinus excelsior L., 2n = 46: Leics., v.c. 55, Outwoods, Jubilee Wood, SK/511.165; Leics., v.c.
55, Swithland Wood, SK/53.12; W. Lancs., v.c. 60, Gait Barrows N.N.R., near Silverdale, SD/
480.776.

Geum rivale L., 2n = 42: W. Lancs., v.c. 60, near Forton, SD/486.534.
Geum urbanum L., 2n = 42: W. Lancs., v.c. 60, near Lancaster, SD/473.594.
Glaucium flavum Crantz, 2n = 12: Dorset, v.c. 9, c. 1-5 km W. of Burton Bradstock, SY/47.89.

Hydrocotyle vulgaris L., 2n=96: W. Lancs., v.c. 60, Leighton Moss R.S.P.B. reserve, near
Silverdale, SD/488.757.

Hypericum androsaemum L., 2n = 40: W. Lancs., v.c. 60, near Lancaster, SD/473.594.

Hypericum elodes L., 2n = 16: S. Devon, v.c. 3, Dartmoor, SX/66.80.

270 SHORT NOTES

Kickxia elatine (L.) Dumort., 2n = 36: Dorset, v.c. 9, near Blandford Forum, ST/88.06.

Lamium album L., 2n = 18: W. Lancs., v.c. 60, W. of Heysham, SD/427.616.

Lemna gibba L., 2n = 40: Cambs., v.c. 29, River Delph, bridge c. 6 km E. of Chatteris, TL/470.858.

Lemna minor L.,2n = 40: Cambs., v.c. 29, River Delph, bridge c. 6 km E. of Chatteris, TL/470.858.

Ligustrum vulgare L., 2n= 46: W. Lancs., v.c. 60, Gait Barrows N.N.R., near Silverdale, SD/
483.775.

Lithospermum officinale L., 2n=28 + 1S: W. Lancs., v.c. 60, Gait Barrows N.N.R., near
Silverdale, SD/479.776.

Littorella uniflora (L.) Asch., 2n = 24: Caerns., v.c. 49, Mynydd Cilan, SH/29.24.

Lotus pedunculatus Cav., 2n = 12: W. Lancs., v.c. 60, River Lune, near Lancaster, SD/483.628;
Westmorland, v.c. 69, Lowick Common, SD/292.847.

Luzula pilosa (L.) Willd., 2n = 66: W. Lancs., v.c. 60, near Millbeck footbridge, SD/648.638.

Lysimachia nummularia L., 2n = 43: W. Lancs., v.c. 60, near Capernwray, SD/528.714.

Lysimachia vulgaris L., 2n = 84: W. Lancs., v.c. 60, River Lune, near Lancaster, SD/484.641.

Malva sylvestris L., 2n = 42: Dorset, v.c.9,c. 1-5 km W. of Burton Bradstock, SY/47. 89; W. Lancs.,
v.c. 60, near Bank Houses, W. of Cockerham, SD/430.531.

Marrubium vulgare L., 2n = 34: Dorset, v.c. 9, Bats Head, SY/79.80.

Medicago lupulina L.., ‘On = 16: Dorset, v.c. 9, c. 1-5 km Ww. of Burton Bradstock, SY/47.84 (two
plants); W. Lancs., v.c. 60, River Lune estuary, edge of new sports centre, near Lancaster, SD/
463.624.

Menyanthes trifoliata L., 2n =54: W. Lancs., v.c. 60, near Lancaster, SD/458.612.

Myosotis stolonifera (DC.) Gay ex Leresche & Levier, 2n = 24: Westmorland, v.c. 69, Lowick
Common, SD/292.847.

Myrica gale L., 2n = 48: Dorset, v.c. 9, Hartland Moor N.N.R. (W. side of road), c. 4kmS.E. of |

Wareham, SY/96.85.

Nepeta cataria L., 2n = 36: Dorset, v.c. 9, between Blandford Forum and Thornicombe, ST/87.04.

Nymphoides peltata Kuntze, 2n = 54: Cambs., v.c. 29, Block Fen gravel pit, near Chatteris, TL/
431.839.

Oenanthe crocata L., 2n = 22: W. Lancs., v.c. 60, estuary of River Lune, near Lancaster, SD/
460.621.

Ononis repens L. subsp. repens, 2n = 60: Caerns., v.c. 49, Abersoch, N. end of Porth Fawr, SH/
31:27:

Oxalis acetosella L., 2n=22: Dorset, v.c. 9, Kingcombe Meadows, c. 1:5 km N.E. of Toller
Porcorum, SY/55. 99.

Parietaria judaica L., 2n = 26: Dorset, v.c. 9, Portland Church Ope Cove, SY/69. 70.

Persicaria amphibia . ) Gray, 2n= 96: W. Lancs., v.c. 60, near Carnforth, SD/507.706.

Persicaria bistorta (L.) Samp., 2n = 48: S. Lancs., v. c. Dos Rochdale Norden, Lower Mancroft Gate
Farm, SD/841.148; Westmorland, v.c. 69, near Arnside, SD/457. 767.

Persicaria maculosa Gray, 2n = 22: W. Lancs., v.c. 60, near Lancaster, SD/465.624.

Petasites hybridus (L.) P. Gaertner, B. Mey. & Scherb., 2n=60: W. Lancs., v.c. 60, near
Quernmore, SD/508.599.

Pimpinella saxifraga L., 2n = 40: Dorset, v.c. 9, Fontmell Down, c. 5kmS.S.E. of Shaftesbury, ST/
88.18.

Polystichum aculeatum (L.) Roth, 2n = 164: W. Lancs., v.c. 60, near Lancaster, SD/473.594.

Populus nigra L. subsp. betulifolia (Pursh) W. Wettst., 2n = 38: W. Lancs., v.c. 60, near Lancaster,
SD/454.615.

Potentilla palustris (L.) Scop., 2n = 35: Westmorland, v.c. 69, Lowick Common, SD/292.847.

Primula veris L., 2n = 22: W. Lancs., v.c. 60, near Silverdale, SD/470.749.

Prunella vulgaris L. (white-flowered), 2n = 28: Westmorland, v.c. 69, Broughton Beck, SD/
285.825.

Ranunculus flammula L. subsp. flammula, 2n = 32: W. Lancs., v.c. 60, Heysham Moss, dismantled
sidings, SD/422.604.

Ranunculus penicillatus (Dumort.) Bab. subsp. pseudofluitans (Syme) S. Webster var. vertumnus C.
Cook, 2n = 48: Mid-W. Yorks., v.c. 64, Bishop Monkton, SE/32.66. |

Ribes nigrum L., 2n = 16: Westmorland, v.c. 69, near Arnside, SD/477.782.

Ribes rubrum Li. 2n = 16: W. Lancs., v.c. 60, near Lancaster, $D/473.594.

i
:
i

Tee a a

SHORT NOTES 271

Rosa arvensis Hudson, 2n = 14: W. Lancs., v.c. 60, near Lancaster, SD/459.613.

Rumex obtusifolius L. subsp. transiens (Simonkai) K. H. Rech., 2n=40: Surrey, v.c. 17,
Richmond, River Thames, TQ/1.7.

Ruppia maritima L., 2n = 20: W. Lancs., v.c. 60, near Carnforth, SD/482.702.

Salix viminalis L., 2n = 38: W. Lancs., v.c. 60, W. of Heysham, SD/428.612.

Samolus valerandi L., 2n = 26: W. Lancs., v.c. 60, near Carnforth, SD/483.704.

Sanguisorba minor Scop. subsp. minor, 2n = 28: W. Lancs., v.c. 60, Leighton Moss R.S.P.B.
Reserve, near Silverdale, SD/489.758.

Sanicula europaea L., 2n=16: W. Lancs., v.c. 60, Leighton Moss R.S.P.B. Reserve, near
Silverdale, SD/489.759.

Scirpus sylvaticus L., 2n = 62: W. Lancs., v.c. 60, River Lune, near Lancaster, SD/485.638.

Sedum forsterianum Smith, 2n =c. 90: Rads., v.c. 43, Stanner Rock, 4 km N.W. of Kington, SO/
26.58.

Silene vulgaris Garcke subsp. vulgaris, 2n = 24: W. Lancs., v.c. 60, River Lune, near Lancaster, SD/
488.640.

Sonchus oleraceus L., 2n = 32: Dorset, v.c. 9, c. 1:5 km W. of Burton Bradstock, SY/47.89.

Sorbus aucuparia L., 2n=34: W. Lancs., v.c. 60, Gait Barrows N.N.R., near Silverdale, SD/
481.776.

Spirodela polyrhiza (L.) Schleiden, 2n = 40: Cambs., v.c. 29, River Delph, bridge c. 6 km E. of
Chatteris, TL/470.858.

Stachys arvensis (L.) L., 2n = 10: Dorset, v.c. 9, Chamberlayne’s Heath, c. 8 km N. of Wool, SY/
83.91.

Stachys officinalis (L.) Trev., 2n = 16: W. Lancs., v.c. 60, Gait Barrows N.N.R., near Silverdale,
SD/478.773.

Taxus baccata L., 2n = 24: Dorset, v.c. 9, Hod Hill, near Biandford Forum, ST/85.10.

Thalictrum flavum L., 2n = 84: W. Lancs., v.c. 60, estuary, near Carnforth, SD/493.714.

Trifolium medium L., 2n= 80: W. Lancs., v.c. 60, Gait Barrows N.N.R., near Silverdale, SD/
478.772.

Valeriana dioica L., 2n = 16: W. Lancs., v.c. 60, above Saltmire Bridge, by canal, SD/519.754.

Veronica anagallis-aquatica L., 2n = 36: Dorset, v.c. 9, near Sydling St Nicholas, ST/63.00.

Veronica beccabunga L., 2n = 18: Westmorland, v.c. 69, near Arnside, SD/477.782.

Veronica officinalis L., 2n = 36: Caerns., v.c. 49, Garn Fadryn, SH/27.35.

ACKNOWLEDGMENTS

We should like to thank English Nature and the R.S.P.B. for permission to collect plant material
from their reserves.

REFERENCES

HOoLiincswortH, P. M., GorNALL, R. J. & Bamey, J. P. (1992). Contributions to a cytological catalogue of the
British and Irish flora, 2. Watsonia 19: 134-137.

A.-K. K. A. AL-BERMANI, K. I. A. AL-SHAMMARY, J. P. BAILEY & R. J. GORNALL
Botany Department, The University, Leicester, LE] 7RH

POLYGONUM MARITIMUM L. IN EAST SUSSEX (V.C. 14)

On 18 June 1992 a number of plants of Polygonum maritimum L. (Polygonaceae), Sea Knotgrass,
were discovered growing on the beach at Brighton, East Sussex by one of us (A. S.). This was an
exciting find as this very rare plant has never before, to our knowledge, been recorded from East
Sussex (v.c. 14) and was last recorded in West Sussex (v.c. 13) by W. Borrer, although no station is
given and there is apparently no specimen in his herbarium. A previous record from Bognor in 1854

272 SHORT NOTES

by Professor I. B. Balfour was thought by H. C. Watson to be Polygonum oxyspermum C. A. Meyer
& Bunge ex Ledeb. subsp. raii (Bab.) D. A. Webb & Chater, and consultation of the herbarium of
the Royal Botanic Garden, Edinburgh (E) confirms this.

Following notification to English Nature, under the terms of the Wildlife & Countryside Act
1981, as a fully protected species under Schedule 8, a specimen was collected (herb. P. A. Harmes)
and confirmed by Dr J. R. Akeroyd. About 14 plants were found, of which eleven or so were
growing together in a clump, seeming at first glance to be a single plant. They ranged in size from
seedlings, with stems no more than 8 cm, up to mature plants with stems 60 cm in length. The plants
were procumbent and were in flower and fruit, which continued throughout the summer and they
were still flowering in late November. They grew well above the high tide mark and seemed to have
a preference for areas of fine shingle. The presence of a number of plants growing in a clump
suggests that this species may have been flowering here un-noticed in previous years, and had
seeded itself.

The small shingle beach where the plants were found was already well-known for its botanical
interest and supports a number of plants found nowhere else along this coast. Species recorded here
already include Atriplex glabriuscula, A. littoralis, A. portulacoides (Halimione portulacoides),
Crambe maritima, Cakile maritima, Raphanus raphanistrum subsp. maritimus, Beta vulgaris subsp.
maritima, Honkenya peploides, Spergularia media, Calystegia soldanella, Crithmum maritimum,
Glaucium flavum, Tripleurospermum maritimum, Parapholis strigosa, Elymus atherica (E. pycnan-
thus) and Catapodium marinum. Nomenclature follows Stace (1991).

The beach is largely composed of flint pebbles with patches of fine shingle or sand, as well as areas
with some humus cover. It is only about 110 m in length and, at its widest, about 50-60 m down to
the high tide mark, with vegetation restricted to about half of the area. For such a small site it has a
surprisingly rich maritime flora. The beach is enclosed on three sides by a groyne, the sea wall and
the western breakwater of Brighton Marina. This breakwater is about 640 m long and was
completed in about 1972. The beach in its present form and position probably dates from this time
and if this is the case, then all the plants here are recent colonists.

A build-up of shingle above the high tide mark has probably occurred here as the western
breakwater prevents its eastern drift. It seems also that this breakwater traps great amounts of
jetsam which is washed or blown on to the beach above the shingle ridge, where it tends to
accumulate. The jetsam consists of plastic, polystyrene, wood, ropes, nets, tar and cans as well as
plant material, including Fucus vesiculosus L. (Bladder-wrack), fruit and seeds, and accumulates in
lower-lying parts of the beach to form a rich mulch in which seeds can be seen germinating. As the
Channel current (North Atlantic Drift) and the prevailing wind through the summer come from the
south-west, it seems quite possible that seed of P. maritimum could have originated from sites in
Cornwall or S. Hants., or from the Channel Isles or Northern France. Less likely is the possibility
that seeds were brought in by birds; Greenfinches are the only seed-eating species (apart from
Sparrows) which frequent the beach.

P. maritimum is a characteristic species of the coasts of the Mediterranean region and is at the
northerly edge of its range in southern England. The recent confirmation of this plant at three other
stations in mainland Britain, together with records from S. E. Ireland and the Netherlands, suggest
that it may be extending, or at least consolidating, its range and this might be related to the hotter,
more Mediterranean-type summers that we have experienced in the south of England in recent
years (Akeroyd 1991). If this is the case, then we may see it turning up on other suitable beaches in
this country in the future. Table 1 shows the number of plants known in mainland Britain (excluding
the Channel Isles).

TABLE 1. POLYGONUM MARITIMUM L. IN BRITAIN

Location Number of plants in colony
W. Cornwall (v.c. 1), Gunwalloe 250
E. Cornwall (v.c. 2), Lantic Bay 1
S. Hants. (v.c. 11), Christchurch 49

E. Sussex (v.c. 14), Brighton 14

SHORT NOTES 273

There are other factors which have enabled coastal plants to become established at this locality.
Unlike many other shingle beaches in Brighton, this one is not regularly bulldozed, perhaps because
there is no access. It is not used much by the public and the amount of sea-borne rubbish as well as
the tar deter most people. Past threats to the beach have come from plans to extend Volk’s Electric
Railway, from Southern Water’s proposal to dump chalk, as well as from over-zealous beach
cleaning. These are hopefully no longer a threat, though beach parties and camp fires still occur.
Brighton Borough Council has, however, recently declared the beach a Site of Nature Conservation
Importance and there is a proposal to extend the adjacent S.S.S.I. to include this site. We hope
these measures will protect the plants here and we will be monitoring the site to see if P. maritimum
survives and increases.

ACKNOWLEDGMENTS

We should like to express our gratitude to the following for their assistance in the compilation of this
note: Dr J. R. Akeroyd, Mrs M. Briggs, P. Bowman, Miss R. Murphy, D. Pearman, Mrs O. M.
Stewart and R. Walls. Mrs C. J. Tatar kindly typed the manuscript.

REFERENCES

AKEROYD, J. R. (1991). Wildlife Reports. Higher Plants. British Wildlife 2: 182-183.
Stace, C. A. (1991). New Flora of the British Isles. Cambridge.

P. A. HARMES
10 Hillcroft, Mile Oak Road, Portslade, East Sussex, BN4 20D
A. SPIERS

69 Elm Grove, Brighton, East Sussex, BN2 3ET

WHITE-BLOSSOMED PINGUICULA GRANDIFLORA LAM. (LENTIBULARIACEAE) IN
THE BURREN, COUNTY CLARE, IRELAND

Pinguicula grandifiora Lam. (Leith Uisce, Kerry (or Large-flowered) Butterwort) has been
reported occasionally from Ireland with white flowers. There are late 19th century records, the
earliest by “Veronica’ (a nom-de-plume used by Frederick W. Burbidge, curator of Trinity College
Botanic Garden, Dublin (cf. Nelson 1987)) being published in The Garden on 11 September 1886:
“As peculiar to Ireland, or nearly so, we have . . . Pinguicula vulgaris var. grandiflora [i.e. P.
grandiflora|, in white, rosy lavender and dark violet forms’ ({Burbidge] 1886). Scully (1916)
recorded that in Co. Kerry while “colour variants are very rarely met with . . . forms with pure
white flowers are . . . occasionally seen and have been gathered in the Gap of Dunloe by Lady
Godfrey and on the east side of Caragh Lake by Capt. Creaghe-Haward . . . pale lilacforms. . . by
Mrs. Jenner from the Gap of Dunloe, and are recorded in the Irish Nat[uralist] 1906, p. 154, as
occurring also on the shores of the Lower Lake, Killarney.” No one has reported white-blossomed
Kerry Butterwort in more recent decades.

In 1903 P. grandiflora was discovered by Professor Ambrose Birmingham at Lisdoonvarna, Co.
Clare (Nelson & Walsh 1991), a habitat to the north of the long-known ones in counties Cork and
Kerry. In 1949, even further north, near Ballyvaughan, another colony was found (Heslop-Harrison
1949), and in 1973, at about the same latitude, yet another (Roden 1984). These latter colonies,
growing in the environs of springs issuing from Carboniferous limestone, are within the region
known as The Burren (Webb & Scannell 1983; Nelson & Walsh 1991).

In 1956, D. A. Webb collected a plant of P. grandiflora “‘with very pale, almost white flowers’”’
near Ballyvaughan and sent it to Steiger (1987) — nothing more is recorded about this plant,
although Webb & Scannell (1983) reported that “for several years around 1970, a small proportion
of the plants . . . had flowers of a very pale lilac colour, but searches in 1974 and 1975 failed to
rediscover them.”

274 SHORT NOTES

During the summers of 1989, 1990, and 1991 in one of The Burren populations, white-flowered
plants were seen and photographed by the present author, on the first occasion in company with Mr
and Mrs J. Leonard. The flowers were entirely white, without any purple or pink marks or tints;
however in 1990 a very young bud of one particular plant, just as it began to rise above the rosette,
had a light pink flush but this was not apparent on the fully open flowers at anthesis. In 1989 and 1990
plenty of seed was produced and shed by the white-blossomed plants.

Steiger (1987) published a photograph of a series of flowers of P. grandiflora variants, including
examples named P. grandiflora f. pallida (Gaudin) Casper and P. grandiflora subsp. rosea (Mutel)
Casper. In f. pallida the corolla throat was purple, and in subsp. rosea not only was the calyx purple
but the corolla throat was lined with darker pink. The white-blossomed Burren examples did not
have coloured markings on the corolla and the calyx is entirely devoid of red pigment. A search of
the literature (see e.g. Casper 1962, 1966) suggests that no entirely white variant of P. grandiflora
has been described hitherto. P. grandiflora subvar. albescens Rouy (1909: “‘corolle blanche lavée de
rose’) could be interpreted as including The Burren variant. (Schlauer (1986) listed ‘‘P. grandiflora
Lam. var. albescens Rouy” but I can find no trace of any legitimate publication of the combination at
varietal level.) However Rouy’s subvariety has been relegated to synonymy under P. grandiflora
subsp. rosea (Casper 1962, 1966) which certainly cannot encompass white-flowered plants
(‘‘corollae . . . fauce violacea vel pallida violacea-pilosa . . .”; Casper 1962: 85).

To designate the variant with a white corolla as a variety is extravagant, and thus I proposed
elsewhere (Nelson & Walsh 1991: 214, 318) that these occasional white-flowered plants should be
placed within a distinct form, P. grandiflora f. chionopetra. I suggest (without having any specimens
to confirm the proposition) that f. chionopetra may also still occur in Co. Kerry, having been
reported from that county before 1903 by Scully (1916).

Pinguicula grandifiora Lam. forma chionopetra E. C. Nelson forma nova.

Corolla candidissima, vel aliquando alabastrum novellum colore roseo suffusum; calyx flavovir-
ens nihil rufescens vel purpurascens.

Corolla pure white, or at most the very young unopened flowers tinted pink; calyx yellow-green
without red or purple tints.
Hototypus: 35 mm Kodachrome 64 colour transparency (no. 16, 25.05.90; accession number
1991.1) [precise locality withheld], County Clare, 12 May 1990, E. C. Nelson (DBN).

The epithet chionopetra is derived from chion (snow) and petra (rock), alluding to the white
flowers and the unique rocky habitat; it is equally appropriate for plants from Co.. Kerry should this
form be collected there again.

A colour photograph is designated as the holotype because, for conservation reasons, I
considered it was unacceptable to remove material from the solitary plant for preservation as an
herbarium specimen. This is permissible under the International Code of Botanical Nomenclature,
Art. 9, whereby an illustration may be a type of an infraspecific name.

REFERENCES

[BuRBIDGE, F. W.] ‘Veronica’ (1886). The Irish flora . . . The Garden 30: 239.

Casper, S. J. (1962). Pinguicula grandiflora, in Revision der Gattung Pinguicula in Eurasien. Feddes
Repertorium Specierum novarum 66: 74-87.

CaspER, S. J. (1966). Pinguicula grandiflora, in Monographie der Gattung Pinguicula. Bibliotheca botanica 127-
128: 163-168.

Hes.op-Harrison, J. (1949). Pinguicula grandiflora Lam. in N. Clare. Ir. Nat. J. 9: 311.

NELSON, E. C. (1987). Who was Veronica of The Garden? Garden History Society Newsletter 20: 3-4.

NEtson, E. C. & Watsu, W. F. (1991). The Burren. A companion to the wildflowers of an Irish limestone
wilderness. Aberystwyth & Kilkenny.

RopENn, C. M. (1984). New stations for Pinguicula grandiflora Lam. and Saxifraga X polita (Haw.) Link in the
west of Ireland. /r. Nat. J. 21: 369.

Rouvy, G. C. C. (1909). Pinguicula grandiflora, in Flore de France 11, 199-200. Paris.

SCHLAUER, J. (1986). Nomenclatural synopsis of carnivorous phanerogamous plants. A world carnivorous plant
list. Carnivorous Plant Newsletter 15: 59-117.

SHORT NOTES 275

ScuLLY, R. W. (1916). Flora of County Kerry. Dublin.

STEIGER, J. F. (1987). Pale-flowered varieties of Pinguicula grandiflora. Carnivorous Plant Newsletter 16: 104—
105.

Wess, D. A. & SCANNELL, M. J. P. (1983). Flora of Connemara and the Burren. Cambridge.

E. C. NELSON
National Botanic Gardens, Glasnevin, Dublin 9, Ireland

CORKSCREW RUSH (JUNCUS EFFUSUS L. FORMA SPIRALIS (J. McNAB) HEGI)
(JUNCACEAE) IN IRELAND AND BRITAIN

Henderson (1992) suggested that the spiral-stemmed variant of Juncus effusus L. (Juncaceae),
found in western Scotland and the Northern Isles of Scotland, represents a taxon different from the
“original”, long-cultivated one variously called, in common parlance, Corkscrew Rush, Irish Rush,
or Spiral Rush. He also stated, incorrectly, that the latter had only been reported once from the
wild.

The corkscrew rush (J. effusus var. spiralis J. McNab) was first collected by David Bishop (a Scot,
one-time curator of Belfast Botanic Garden — see Anon. 1849; Nelson 1984, 1987) in ‘“‘the wilds of
Connemara’’, western Ireland, before 1849 and not in “Northern Ireland. . . in 1869” as noted by
Henderson (1992) repeating McNab’s (1873a, 1873b) faulty recollection. The exact date of
discovery cannot now be ascertained but Bishop died in 1849, so he must have collected the
Corkscrew Rush some time earlier. Indeed, the first report of the variant appeared in an obituary of
Bishop (Anon. 1849):

“this extraordinary plant was exhibited at a meeting of the Botanical Society of Edinburgh, from

the collection at Dalkeith, by Mr. James M’Nab, and created great interest .. .”

I cannot trace a report of this particular exhibit. Several decades later, in April 1873, James McNab
read a brief account of the Corkscrew Rush to the Botanical Society in Edinburgh (McNab 1873a,
1873b), and indicated that it had been on display at the British Association for the Advancement of
Science’s 1871 meeting in Edinburgh; again I cannot trace any specific mention of the rush’s
appearance at the Association.

Henderson (1992) separated the western Scottish Spiral Rush from the Irish one by stating that
the latter has “quite erect [stems] . . . much more obviously spiral six to eight turns as against two or
three [in Scottish plants]’’, but this is not the case. I have examined living individuals of the Irish
Corkscrew Rush, as cultivated in the National Botanic Gardens, Glasnevin, and nursery-grown
plants (source unknown) (September 1992); these possessed stems with as many as 14 complete
rotations in the spiral, ranging to stems with a single attenuated rotation (the stem was merely
curved), as well as erect, untwisted stems. The largest proportion of the spiral stems (50%) had
fewer than five rotations; only 33% had 5—9 rotations. The stems projected at all angles, the less
spiralled ones tending to spread almost horizontally (i.e. these were suberect) because of the spiral.
Thus cultivated plants, propagated vegetatively from long-established Irish stock, do not have the
characteristics suggested by Henderson (1992); his variety is dubiously distinct from the cultivated
plant.

Several herbarium specimens in the Royal Botanic Garden, Edinburgh (see below), have
remarkable corkscrewing stems, but clearly these particular stems were selected because of their
form; they do not represent accurately the habit and range of spiralling in the original clone. The
- earliest illustration of the Corkscrew Rush from Ireland (The Gardeners’ Chronicle and Agricultural
Gazette, 10 May 1873: 647 (McNab 1873a; reprinted in McNab 1873b: 503)) showed a plant with
erect and spreading stems, some markedly spiralled but many merely curved (Fig. 1), exactly as in
those cultivated plants I have examined. A photograph published by Tutenberg (1905) also clearly
displays variation in habit. Those authors presumably had plants derived from Bishop’s original
collection.

As for Irish populations, Praeger (1934: 406), repeating records of this taxon from Inishturk (Co.
Mayo, v.c. H27) off the Connemara coast (Praeger 1907: 123 — ‘“‘the form with spreading stems was
several times observed’’), stated that it had

“‘spreading loosely spiral stems . . . The spiral rush just mentioned would appear to be an Atlantic

276 SHORT NOTES

VEN

)

Va)

(Sah
Ul
a

TST TA

Figure 1. Corkscrew Rush, Juncus effusus L. forma spiralis (McNab) Hegi; illustration published in The
Gardeners’ Chronicle and Agricultural Gazette, 10 May 1873: 647 (designated as neotype herein).

form: it is common on many of the Irish western islands, and is stated to be abundant in
Orkney...”

Praeger (1934: 424) also repeated records from Inishmurray (Co. Sligo, v.c. H28) —
‘Perhaps the most curious plant of the island was a diffuse form of Juncus conglomeratus [= J.
effusus; fide Praeger (1934)], the stems of which, instead of growing erect in a compact clump as
usual, spread out at every angle, from horizontal to vertical, giving the whole plant a very strange
appearance. . . this curious rush was abundant in damp places with the typical form.” (Praeger
1896: 178)

—and from Achill Island (Co. Mayo, v.c. H27; Praeger 1934: 408) —
“Near Sraheens village occurred a form with widely spreading stems, many of them spirally
curved, with several convolutions. I have gathered the same form from Inishmurray, Co. Sligo.
Mr Beeby informs me that in Orkney these spiral forms are frequent.”’ (Praeger 1904: 285)

Spence (1906, 1914, 1919) reported spiral-stemmed J. effusus from Orkney (v.c. 111), and Druce
(1922: 524) and Scott & Palmer (1987: 343) recorded this variant from Shetland (v.c. 112); the
latter authors suggested that the spiralling stems are deformed by wind —
“Juncus effusus var. spiralis McNab’, recorded from various localities by Druce [1922], was
surely no more than a wind-blown state with slightly curved stems, not the monstrosity with
corkscrew-like stems which James McNab grew .. .”

The Spiral Rush has been recorded in Surrey (v.c. 17) (see list of specimens below, and Leslie 1981),

and collected in Carmarthenshire (v.c. 44, see below).

SHORT NOTES 277

For at least a century, botanists have assigned to McNab’s variant all plants of J. effusus producing
spiral stems. Having examined herbarium material and living plants, I have been unable to detect
any difference between the variable Irish original and Henderson’s newly described taxon. Perhaps
a case can be made on the basis of habit — that western Scottish plants lack any erect stems — but the
range of variation within ‘populations’ of vegetatively propagated plants is so substantial that such
an argument could not be justified. I conclude, therefore, that all plants of this species with spiral
stems, irrespective of the degree of spiralling or angle of inclination, should be placed, as generally
done by previous authors, within the same taxon. Henderson’s varietal epithet is deemed
unnecessary and is here relegated to synonymy.

The status of the Spiral Rush is debatable. McNab (1873b) and Henderson (1992) reported that
seedlings have spiralling stems so it is a distinct genotype. While horticulturists have considered it
little more than a cultivar (J. effusus ‘Spiralis’ is then the valid name; cf. Nelson 1984), its occurrence
in the wild, especially in western Ireland and Scotland, suggests that it deserves recognition as a
botanical variety or form. As the variant is clearly distinguished from the common rush only by its
spiral stems, both erect and spreading, I suggest it is best treated as a botanical form; it does not
warrant recognition at varietal level.

_ Hegi (1909: Bd 2: 147, fig. 274) was the first author to employ the epithet ‘spiralis’ at form level
(Juncus effusus f. spiralis). Hegi did not attribute the epithet to any author and did not cite McNab’s
original paper, but it is unlikely that he was doing anything other than employing McNab’s well-
known epithet for the cultivated Irish plant at a revised rank; he was not describing a new taxon. To
argue that Hegi was describing a new taxon is unhelpful, because his epithet ‘spiralis’ becomes
illegitimate (Arts. 24. note 1: 64.4) and a new epithet becomes necessary for the form. Praeger
(1934: entry no. 406) also used J. effusus f. spiralis; he did not acknowledge Hegi but did refer to
Spence’s note (1919) which in turn referred to an earlier account (Spence 1906) that contains explicit
mention of McNab’s description (1873b). The synonymy may be summarised as follows:

Juncus effusus L. forma spiralis (J. McNab) Hegi, I/lustrierte Flora von Mittel-Europa, 1909: Bd.
2(16), 147.

Basionym: J. effusus var. spiralis J. McNab, Gard. Chron. 10 May 1873: 647, fig. 125; Trans. bot.
Soc. Edinb. 11 (1873): 502-504.

Neotype: [here designated] icon in Gard. Chron. 10 May 1873: 647, fig. 125.

Synonym: J. effusus var. suberectus D. M. Henderson, Watsonia 19 (1992): 133-134.

Holotype: Big Sand, Gairloch, W. Ross, v.c. 105, 6 December 1988, D. M. Henderson (E!).

Other specimens examined:

Sandy edge of site of Frensham Great Pond, Surrey, v.c. 17, [19 August 1943], A. J. Wilmott
19430819 (BM); ‘“‘grouse moor . . . Carmarthenshire Vans”, v.c. 44, 12 August 1904, E. Milner-
Jones (K); Orkney, v.c. 111, 1 September 1906, M. Spence (E); Deerness, Orkney, v.c. 111,
September 1908, M. Spence (BM); Rannsdale, Orphir, Mainland, Orkney, v.c. 111, 10 September
1923, H. H. Johnston 2474 (E, BM, K); Sutherland, Flotta, Orkney, v.c. 111, 15 August 1932, J.
Sinclair 762 (E); Mainland, Shetland, v.c. 112, June 1890, R. M. Barrington (DBN); roadside, Lax
Firth, Mainland, Shetland, v.c. 112, 27 July 1950, J. E. Lousley (K); by burn of Sandibanks,
Scalloway, Mainland, Shetland, v.c. 112, 25 July 1950, J. E. Lousley (K); near Sraheens, Achill
Island, v.c. H27, 30 July 1904, R. Li. Praeger (DBN); boggy place centre of island [Inishtrahull, Co.
Donegal], v.c. H34, 16 August 1939, D. J. Sullivan (DBN).

Cultivated specimens:

Edin[burgh] Bot. Garden, September 1876, F.M.W. (E); Botanic Garden [Edinburgh], 1886, (E);
Wakehurst Place, Sussex (acc. no. 000-69-19251), 20 July 1980, S. Andrews (K); sine loc. (‘Herb.
Hort. Kew’), Aug. 1881 (K).

ACKNOWLEDGMENTS

I must express my gratitude to Arthur Chater, Dr R. K. Brummitt, C. D. Preston and Dr John
Edmondson for their helpful comments on a draft of this paper, and to the Royal Botanic Garden,

278 SHORT NOTES

Edinburgh, the National Museums & Galleries on Merseyside, Liverpool, the Royal Botanic |
Gardens, Kew, and the Natural History Museum, London, for loans of herbarium specimens.

REFERENCES

Anonymous (1849). ‘“‘Mr David Bishop . . . [obituary]’’. Cottage Gardener 2: 306-307.

Druce, G. C. (1922). Flora zetlandica. Rep. botl Soc. Exch. Club Brit. Isl. 6: 457-526.

Heci, G. (1909). [Juncus effusus f. spiralis]. Illustrierte Flora von Mittel-Europa. Bd. 2(16): 147, fig. 274.
Munich.

HENDERSON, D. M. (1992). A new spiral variant of Juncus effusus L. (Juncaceae). Watsonia 19: 133-134.

LesuiE, A. C. (1981). The flora of Wisley and Ockham Commons, with Chantley Heath, Surrey. London
Naturalist 60: 21-26.

McNas, J. (1873a). Remarks on Juncus effusus spiralis and the varieties of ferns. Gard. Chron. 10 May 1873:
647.

McNas, J. (1873b). Remarks on Juncus effusus spiralis and the varieties of ferns. Trans. bot. Soc. Edinb. 11:
502-504.

NELSON, E. C. (1984). [Juncus effusus ‘Spiralis’| in An Irish flower garden, pp. 23-24. Kilkenny.

NELSON, E. C. (1987). Scottish connections in Irish botany and horticulture. Scottish Nat. 1987: 3-31.

PRAEGER, R. L. (1896). The plants of Inishmurray, Co. Sligo. Irish Nat. 5: 177-178.

PRAEGER, R. L. (1904). The flora of Achill Island. Irish Nat. 13: 265-289.

PRAEGER, R. L. (1907). The flora of Inishturk. Irish Nat. 16: 113-125.

PRAEGER, R. L. (1934). The botanist in Ireland, entry number 406. Dublin.

Scott, W. & PALMER, R. (1987). The flowering plants and ferns of the Shetland Islands. Lerwick.

SPENCE, M. (1906). Note on Juncus effusus, var. spiralis. Trans. bot. Soc. Edinb. 23: 233.

SPENCE, M. (1914). Flora orcadensis, p. 78. Kirkwall.

SPENCE, M. (1919). Juncus effusus spiralis. J. Bot., Lond. 57: 69.

TUTTENBERG, F. (1905). Juncus effusus var. spiralis. Gartenflora 54: 406-407, abb. 53.

E. C. NELSON
National Botanic Gardens, Glasnevin, Dublin 9, Ireland

Watsonia, 19, 279-295 (1993) 279

Plant Records

Records for publication must be submitted to the appropriate Vice-county Recorder (see B.S.B.I. Year Book
for 1993), and not the Editors. The records must normally be of species, hybrids or subspecies of native or
naturalized 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 vascular plants of the British Isles by D. H. Kent (1992),
from which the species’ numbers, taxonomy and nomenclature are taken. The B.S.B.I. is to set up a procedure
to register changes to this list, and a number of records of additional species have been held over until the new
arrangements are in place. The Ordnance Survey national grid reference follows the habitat and locality. 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 are field records if no other source is stated.

Records from the following vice-counties are included in the text below: 2, 4-6, 9, 11, 14, 17, 21, 22, 24-30, 35,
38, 39, 41-53, 57-59, 64, 67-73, 75, 77-81, 83, 85-87, 89, 93, 98-103, 108.

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.

4/1.1. EQUISETUM HYEMALE 42, Brecs.: Stream-bank near Hay-on-Wye, SO/243.404. W. J. H.
Price, 1992. 2nd record. 98, Main Argyll: Stony flush, Coire Dearg, Fraochaidh, NN/043.518.
R. Leishman et al., 1992. 2nd record. 108, W. Sutherland: Banks of burn, Allt na h-Airbhe, NC/
12.23. P. A. & I. M. Evans, 1992, herb. I.M.E. 2nd confirmed post-1930 record.

4/1.8 X 9. EQUISETUM PALUSTRE X E. TELMATEIA (E. X FONT-QUERI) *11,S. Hants.: Roadside,
South Baddesley, Boldre, SZ/349.966. M. W. Rowe, 1991, E, det. C. N. Page.

9/1.1. PILULARIA GLOBULIFERA *108, W. Sutherland: Loch Borralan, NC/26.10. E. Charter,
1983. Shallow water, Cam Loch, NC/22.12. P. A. & I. M. Evans, 1992, herb. I.M.E. 1st and 2nd
records.

11/1.1 x 2. PoOLYPODIUM VULGARE X P. INTERJECTUM (P. X MANTONIAE) *64, Mid-W. Yorks.:
Hackfall Wood, Grewelthorpe, SE/23.77. D. J. & J.S. Tennant, 1991, det. R. H. Roberts. wey
Ayrs.: Cliffs, Ailsa Craig, NX/02.99. B. Zonfrillo, 1992, det. A. McG. Stirling.

11/1.2. POLYPODIUM INTERJECTUM *64, Mid-W. Yorks.: Near Beezley Falls, Ingleton, SD/
70.74. D. J. Tennant, 1979. Scotton Banks, Knaresborough, SE/33.57. D. J. Tennant, 1985. 1st and
2nd records, both det. R. H. Roberts.

15/2.5c. ASPLENIUM TRICHOMANES subsp. PACHYRACHIS *67, S. Northumb.: Calcareous rock
wall, Hareshaw Linn near Bellingham, NY/842.854. J. M. Ide, 1992, BM, det. A. C. Jermy.

15/2.6. ASPLENIUM TRICHOMANES-RAMOSUM *46, Cards.: Wheelpit and shafts of leadmine,
Esgair-Fraith, SN/741.912. S. P. Chambers & J. A. Martin, 1992.

17/1.1. POLYSTICHUM SETIFERUM 59, S. Lancs.: Base of wall near Chorlton Brook, Chorlton,
SJ/813.931. D. Bishop, 1991. 1st post-1930 record.

17/3.2 X 3. DRYOPTERIS FILIX-MAS X D. AFFINIS (D. X COMPLEXA) *58, Cheshire: Trackside,
Roman Lakes, Marple, SJ/968.876. B. Porter, 1992, NMW, det. H. Corley.

280 PLANT RECORDS

17/3.3b. DRYOPTERIS AFFINIS Subsp. CAMBRENSIS *83, Midlothian: Rocky moorland slope N. of |

White Craig, NT/072.543. D. R. McKean, 1992, det. A. C. Jermy.

17/3.8 X 9. DRYOPTERIS CARTHUSIANA X D. DILATATA (D. X DEWEVERI) *83, Midlothian:
Marshy area in wood, Hermands Birchwood S.W.T. Reserve near West Calder, NT/0.6. A. F.
Dyer, 1986, E, conf. C. N. Page.

26/2.2. NUPHAR PUMILA *73, Kirkcudbrights.: Kirriereoch Loch, NX/364.866. O. M. Stewart,
1991. B:

+28/6.1 X var. ACONITUM NAPELLUS X A. VARIEGATUM (A. X CAMMARUM) *44, Carms.:
Roadside verge, Llangunnor Road, Carmarthen, SN/422.196. G. Hutchinson, 1990, NMW.
*50, Denbs.: Roadside, Trefnant, SJ/056.712. G. Battershall, 1992.

28/13.7. RANUNCULUS PARVIFLORUS 24, Bucks.: Grazed turf by old gravel pit S. of Ritchings
Park, TQ/038.782. D. Green, 1992. 1st post-1930 record.

28/13.14. RANUNCULUS REPTANS *98, Main Argyll: Sandy silt on loch shore, Loch Awe near
Collaig, NN/022.204. B. H. Thompson, 1992, LTR, det. R. J. Gornall.

28/17.3. THALICTRUM FLAVUM 45, Pembs.: Tall fen, Castlemartin Corse, SR/899.997. F. A.
Abraham, 1992. 2nd record.

729/2.1. MAHONIA AQUIFOLIUM *77, Lanarks.: Roadside bank between Biggar and Coulter,
NT/0.3. A. C. & P. Macpherson, 1992.

+30/1.1. PAPAVER ORIENTALE *73, Kirkcudbrights.: Disused quarry E. of Haugh of Urr, NX/
805.673. O. M. Stewart, 1992.

+31/2.2 CORYDALIS CAVA 6, N. Somerset: Dominant plant in ground flora, Terrace Wood,
Ston Easton, ST/630.540. S. Preddy, 1992. ist record since 1922 record from Ston Easton.

40/2.1 X +2. ALNUS GLUTINOSA X A. INCANA (A. X PUBESCENS) *46, Cards.: Wet alder wood
and adjacent waste ground, Llanilar railway station, SN/628.752. A. O. Chater, 1992, NMW.

+40/2.2. ALNUSINCANA *50, Denbs.: Roadside wood, Llanfair Dyffryn Clwyd, SJ/169.506. J. A.
Green, 1992.

+43/3.1. ATRIPLEX HORTENSIS *58, Cheshire: Shore just above tideline, Hoylake, §J/210.890.
V. Gordon, 1992.

43/3.4. ATRIPLEX LONGIPES *77, Lanarks.: Tidal shore of R. Clyde, Linthouse, Glasgow, NS/
53.66. P. Macpherson, 1985, herb. P. M., det. J. M. Mullin. Not refound.

43/4.1a. BETA VULGARIS subsp. MARITIMA 75, Ayrs.: Shingle near mouth of Pinbain Burn,
Lendalfoot, NX/138.916. A. McG. Stirling, 1992. Only extant locality.

43/6.4. SALICORNIA OBSCURA *6, N. Somerset: Bare mud of lower salt marsh, St George’s
Wharf, Easton-in-Gordano, ST/495.779. M. A. R. & C. Kitchen, 1991, det. F. Rose.
43/6.6. SALICORNIA FRAGILIS *6, N. Somerset: Pebbly, sandy bare area in lower salt marsh, St

George’s Wharf, Easton-in-Gordano, ST/492.775. M. A. R. & C. Kitchen, 1991, det. F. Rose.

+46/1.4. ARENARIA BALEARICA 43, Rads.: Silty concrete top of unfinished dam, Dolymynach
Reservoir, SN/909.619. R. G. Woods, 1992. 2nd record.

46/7.11. CERASTIUM PUMILUM 50, Denbs.: Limestone grassland, Rhyd y Foel, SN/917.777,
and Colwyn Bay, SN/830.799. Both G. Battershall, 1992. 1st and 2nd post-1930 records.

46/9.1. MOENCHIA ERECTA *41, Glam.: By paths through Agrostis curtisii heath on S.-facing
hillside E. of Nottill, SS/537.885. Q. O. N. Kay, 1992, NMW. 44, Carms.: Dry, S.-facing bank
near Cynghordy, SN/820.390. I. K. Morgan, 1992, NMW, conf. G. Hutchinson. 2nd post-1930
record.

46/17.3. SPERGULARIA MARINA *80, Roxburghs.: Edge of A6088 road near Southdeanrig, NT/
646.085. Edge of A68 road near Mossburnford, NT/663.155. Both M. E. Braithwaite, 1992, herb.

PLANT RECORDS 281

R. W. M. Corner. ist and 2nd records. 81, Berwicks.: Edge of A697 road near Carfraemill, NT/
517.526. M. E. Braithwaite, 1992, herb. M.E.B. 1st inland record.

+46/18.1. LYCHNIS CORONARIA *51, Flints.: Track leading to disused sand pit near Ddél Uchaf
N.R., Ysceifiog, SS/140.715. J. Hughes, 1992.

746/22.2. SAPONARIA OCYMOIDES *28, W. Norfolk: Steep wooded bank, King’s Lynn, TF/
625.189. R. M. Payne, 1992.

46/25.5. DIANTHUS DELTOIDES *79, Selkirks.: Rocky basic pasture, Tinnis Top, Old Tinnis,
NT/383.292. D. J. Methven, 1992, conf. R. W. M. Corner.

747/1.7. PERSICARIA AMPLEXICAULIS *39, Staffs.: Rough grassland by A515, Draycott in the
Clay, SK/159.295. J. Clarke, 1992, herb. B. R. Fowler.

47/1.11 X 14. PERSICARIA MACULOSA X P. HYDROPIPER (P. X INTERCEDENS) *30, Beds.: Bank of
R. Great Ouse, Great Barford, TL/13.52. J. G. & C. M. Dony, 1987, LTN, conf. J. Timson.

47/1.12. PERSICARIA LAPATHIFOLIA 80, Roxburghs.: Field margin, Clarilaw, Hawick, NT/
525.184. M. E. Braithwaite, 1992, herb. R. W. M. Corner, det. J. R. Akeroyd. 1st confirmed post-
1930 record.

47/1.16. PERSICARIA MINOR 44, Carms.: Edge of pond, Gwaith Go-Bach Pond, Dinefwr Castle
Woods, SN/619.221. I. K. Morgan, 1992, NMW, det. G. Hutchinson. 2nd record. 67, S.
Northumb.: Stony lake margin, Greenlee Lough, NY/772.695. G. A. & M. Swan, 1992, herb.
G.A.S., det. J. R. Akeroyd. 2nd record.

47/4.1. POLYGONUM MARITIMUM *14, E. Sussex: Stable shingle beach, W. side of Brighton
Marina, TQ/33.03. A. Spiers, 1992, herb. P. A. Harmes, conf. J. R. Akeroyd. 15 plants. (see pp.
271-273.)

47/4.5. POLYGONUM BOREALE *73, Kirkcudbrights.: Vegetable patch, New Abbey, West
Maryfield, NX/970.664. O. M. Stewart, 1991, conf. D. R. McKean. Still present in 1992. Turnip
field, Milnmark, NX/657.820. O. M. Stewart, 1992, E, conf. J. R. Akeroyd. 1st and 2nd
records. 103, Mid Ebudes: Ground trampled by cattle, The Reef, Tiree, NM/006.449. B. H.
Thompson, 1990, GLAM, det. B. T. Styles. 2nd record.

47/5.41 X +2. FALLOPIA JAPONICA X F. SACHALINENSIS (F. X BOHEMICA) *50, Denbs.: Waste
ground, Wrexham, SJ/338.490. K. Watson, 1992, conf. V. Gordon. *59, S. Lancs.: Waste
ground beneath Pier, Southport, SD/833.177. V. Gordon, 1992, det. A. P. Conolly.

47/8.13b. RUMEX CRISPUS subsp. LITTOREUS *58, Cheshire: Boulder clay sea-cliffs, Thurstas-
ton, SJ/23.83. G. M. Kay, 1992. *98, Main Argyll: Coastal shingle and rock, Loch Gilp, NR/
86.85. A. McG. Stirling & B. H. Thompson, 1992. Rocky shore, W. coast of Kerrera, NM/80.29.
B. H. Thompson, 1992. 1st and 2nd records.

47/8.13b X 18. RUMEX CRISPUS subsp. LITTOREUS X R. PULCHER (R. X PSEUDOPULCHER) 725,
E. Suffolk: Stable shingle, Aldeburgh, TM/46.56. G. D. Kitchener, 1992, herb. G. D. K., conf. J. R.
Akeroyd. Recorded here in 1976 by J. R. Akeroyd & C. D. Preston. Ist ever record of this hybrid
with subsp. littoreus as R. crispus parent.

— 47/8.13 xX 19. RUMEX cRISPUS X R. OBTUSIFOLIUS (R. X PRATENSIS) *103, Mid Ebudes:
‘Garden, Isle of Erraid, NM/30.20. J. W. Clark, 1992, E, det. D. R. McKean.

47/8.20. RUMEX PALUSTRIS *24, Bucks.: Dry, overgrown ditch between Colnebrook and
Sutton, TQ/025.781. D. Green, 1992, AYM.

48/1.10d. LIMONIUM BRITANNICUM subsp. CELTICUM *69, Westmorland: On limestone, Frith
Hall, estuary of R. Leven, SD/3.7. W. H. Pearsall, 1916, YRK. Stonework of Arnside railway
viaduct, SD/4.7. C. Webb, 1988, LANC. 1st and 2nd records of species, both det. M. Ingrouille.

51/1.7b. HYPERICUM MACULATUM subsp. OBTUSIUSCULUM *81, Berwicks.: Woodland edge near
Carfraemill, NT/510.529. M. E. Braithwaite, 1992, herb. M.E.B. 1st record since 1938 and only
extant locality.

282 PLANT RECORDS

51/1.12. HyPERICUM PULCHRUM 75, Ayrs.: Ailsa Craig, NS/01.00. B. Zonfrillo, 1992, GL. 1st
record from Ailsa Craig.

52/1.1. TILIA PLATYPHYLLOS +*50, Denbs.: Hedge, Colwyn Bay, SH/837.778. G. Battershall,
1992.

53/1.1. MALVA MOSCHATA +*98, Main Argyll: Area of hard-standing, Loch Ederline, NM/
870.029. A. McG. Stirling & B. H. Thompson, 1992.

753/1.2. MALVA ALCEA *41, Glam.: Waste ground, Sully, ST/163.679. S. G. Lambert, 1991,
NMW, det. D. McClintock. 1st Welsh record.

+53/1.6. MALVA PUSILLA *28, W. Norfolk: Waste ground by track in Oxborough Wood, TF/
722.012. J. E. Caffney, 1992, det. N. K. B. Robson.

53/3.1. ALTHAEA OFFICINALIS 28, W. Norfolk: Dikeside, Marshland St James, TF/503.103.
R. M. Payne, 1992. 2nd record.

57/1.4 X 6. VIOLA RIVINIANA X V. CANINA (V. X INTERSITA) *73, Kirkcudbrights.: Disused
railway, Waterside, NX/723.676. O. M. Stewart, 1992, det. A. J. Silverside.

57/1.6. VIOLA CANINA 83, Midlothian: Forest ride, Gladsmuir, NT/919.572. J. Muscott, 1992.
1st post-1930 record.

457/1.11 xX 12 X alt. VIOLA X WITTROCKIANA *71, Man: Dumped garden rubbish, Ramsey
Mooragh, SC/451.955. L. S. Garrad, 1989.

61/2.2 X 3. SALIX FRAGILIS X S. ALBA (S. X RUBENS) *47, Monts.: River bank, Cilcewydd,
Welshpool, SJ/227.034. I. C. Trueman, 1991, det. C. A. Sinker.

+61/2.6. SALIX DAPHNOIDES 44, Carms.: Coastal dunes, Pembrey Forest, SN/405.012. R. D.
Pryce et al., 1992, NMW, det. G. Hutchinson. /

61/2.9 X 12. SALIx VIMINALIS X S. AURITA (S. X FRUTICOSA) *47, Monts.: River bank,
Cilcewydd, Welshpool, SJ/226.033. I. C. Trueman, 1991, det. C. A. Sinker.

61/2.10 X 12. SALIX CAPREA X S. AURITA (S. X CAPREOLA) *46, Cards.: Scrub on riverbank,
Afon Rheidol by Glanyrafon Industrial Estate, SN/614.804. A. O. Chater, 1991, NMW, conf. R. D.
Meikle.

61/2.12 x 15. SALIX AURITA X S. PHYLICIFOLIA (S. X LUDIFICANS) *83, Midlothian: Marsh by
Gala Water, Heriot House, NT/40.54. R. Learmouth, 1992, herb. R.L., det. R. D. Meikle.

761/2.13. SALIX ERIOCEPHALA *46, Cards.: Amongst Molinia caerulea on bank of Afon
Ystwyth, Llanfarian, SN/588.778. S. P. Chambers & A. O. Chater, 1992, NMW, conf. R. D.
Meikle.

+62/7.1. ERYSIMUM CHEIRANTHOIDES *46, Cards.: Newly reconstructed verge of A44(T),
Lovesgrove, SN/631.811. A. O. Chater, 1992, NMW.

62/12.5. RORIPPA SYLVESTRIS *93, N. Aberdeen: Waste ground, Insch, NJ/629.278. D. Welch,
1992, ABD, conf. T. C. G. Rich.

+62/16.1. AUBRIETA DELTOIDEA 50, Denbs.: Disused quarry N. of Bryn Euryn, SH/8.8.
R. Lewis, 1992. 2nd record.

62/21.4. DRABA MURALIS +*52, Anglesey: Disused limestone quarry near Llangoed, SH/60.81.
D. F. Evans, 1991.

62/22.3. EROPHILA GLABRESCENS *79, Selkirks.: Rocks by R. Tweed below Yair Bridge, NT/
459.325. *80, Roxburghs.: Rocky pasture, Smailholm Craigs, NT/637.347. Both R. W. M.
Corner, 1991, herb. R.W.M.C., det. T. T. Elkington.

62/30.3. LEPIDIUM HETEROPHYLLUM 28, W. Norfolk: Field edge, Marham, TF/732.103.
J. Williamson, 1992, det. K. A. & G. Becket. 2nd record.

PLANT RECORDS 283

62/30.5. LEPIDIUM RUDERALE 70, Cumberland: Gullery, S. end of Rockcliffe Marsh, NY/
310.626. D. Hawker, 1992, LANC. Disturbed roadside verge, Brunstock, Carlisle, NY/426.592.
P. Burton, 1992, LANC. Ist records since 1908.

+62/30.8. LEPIDIUM DRABA 81, Berwicks.: Railway bank, Reston, NT/878.619. A. R. Jermyn,
1992. 1st record since 1892. Side of A6112 road between Duns and Grantshouse, NT/812.645. M. E.
& P. F. Braithwaite, 1992. 2nd extant locality.

762/30.8b. LEPIDIUM DRABA subsp. CHALEPENSE *6, N. Somerset: Salt marsh, St George’s
Wharf, Portbury, ST/500.780. M. A. R. & C. Kitchen, 1991, det. T. C. G. Rich. *29, Cambs.:
Waste area at corner of arable field, Morden Grange chalkpit, TL/295.401. A. Showler, 1992, det.
TFaGsG Rich:

+62/36.1. ERUCASTRUM GALLICUM *5, S. Somerset: Disturbed roadside bank, Bayford, ST/
728.290. I. P. Green, 1992. Gravel heap, Henstridge Marsh, ST/754.200. I. P. Green, 1992. 1st and
2nd records.

762/37.1b. CoINCYA MONENSIS subsp. RECURVATA 44, Carms.: Penrhyngwyn, Machynys,
Llanelli, SS/517.973. I. K. Morgan, 1992. NMW, det. G. Hutchinson. 2nd record.

+62/38.1. HIRSCHFELDIA INCANA *42, Brecs.: Roadside verge near Brecon, SO/067.278.
M. Porter, 1991. *71, Man: Tip S.W. of Point of Ayre, NX/460.040. B. A. Tregale, 1988.

63/1.3. RESEDA LUTEA *93, N. Aberdeen: Disused gravel pit, Memsie, NJ/985.620. D. Welch,
1991, ABD.

64/1.1b. EMPETRUM NIGRUM subsp. HERMAPHRODITICUM 73, Kirkcudbrights.: Rocky ground on
E. side of Carlin’s Cairn, Corserine, NX/498.882. R. W. M. Corner, 1992, E. 2nd record.

65/13.2. VACCINIUM MICROCARPUM *93, N. Aberdeen: Sphagnum flush, Buck of Cabrach, NJ/
417.238. R. W. M. Corner, 1991, herb. D. Welch. 2nd record.

66/1.3. PYROLA ROTUNDIFOLIA *93, N. Aberdeen: Acidic flush, Buck of Cabrach, NJ/413.239.
R. W. M. Corner, 1992. ist localised record.

67/1.1b. MONOTROPA HYPOPITYS subsp. HYPOPHEGEA +58, Cheshire: Large colony in old tree
nursery, Priory Gardens Reserve, Sale, SJ/802.926. A. & S. Bell, 1992. Only extant locality.

69/1.2. PRIMULA ELATIOR *24, Bucks.: Ancient woodland over calcareous clay, Bovington
Great Wood, TL/002.052. R. Mabey, 1977. Heather Grove, SP/994.055. R. Mabey, 1977-1989. 1st
and 2nd records.

69/1.3. PRIMULA VERIS 73, Kirkcudbrights.: Short turf near shore, Burnfoot, NX/742.446.
O. M. Stewart, 1991. Only extant locality.

+69/4.4. LYSIMACHIA CILIATA 98, Main Argyll: Damp ground by Loch Ederline, NM/870.029.
A. McG. Stirling & B. H. Thompson, 1992. 2nd record.

69/8.1. SAMOLUS VALERANDI *24, Bucks.: Margin of lake formed after gravel extraction,
Great Linford, SP/845.439. C. Machaddie, R. Maycock & A. Woods, 1991.

172/1.1. ESCALLONIA MACRANTHA *49, Caerns.: Woodland edge, Coed Tan yr Allt between
Pontwgan and Rowen, SH/761.717. R. Lewis, 1992, NMW, det. R. G. Ellis & G. Hutchinson.

+72/2.4. RIBES SANGUINEUM *98, Main Argyll: Forest roadside, Minard Castle, NR/970.951.
B. H. Thompson, 1992.

+73/1.3. CRASSULA HELMSII *30, Beds.: Pond near Podington, SP/937.628. M. Powell, 1988,
LTN, conf. C. R. Boon. *70, Cumberland: Water 70 cm deep, Lord’s Bay, Derwent Water,
NY/266.218. G. Halliday, 1992, LANC, det. C. D. Preston.

+73/5.7. SEDUM SPURIUM *28, W. Norfolk: On concrete, Ickburgh, TL/823.950. K. A.
Beckett, 1992.

284 PLANT RECORDS

+73/5.10. SEDUM RUPESTRE *98, Main Argyll: Steep bank between road and seashore near
Kilchoan, Loch Melfort, NM/803.131. B. H. Thompson, 1992. Churchyard wall between North
Connel and Ardchattan, NM/944.359. B. H. Thompson, 1992. Ist and 2nd records.

+74/4.1. DARMERA PELTATA *73, Kirkcudbrights.: Marshy ground, Carruchan, NX/949.734.
O. M. Stewart, 1992.

+75/3.1 X 3. SPIRAEA SALICIFOLIA X S. DOUGLASII (S. X PSEUDOSALICIFOLIA) *29, Cambs.: By
footpath leading from road between Swavesey and Over, TL/368.701. J. C. A. Rathwell, 1992,
CGE. 1st confirmed record.

+75/3.2. SPIRAEA ALBA 70, Cumberland: Roadside hedge, Aikshaw, Westnewton, NY/
124.463. E. Marper, 1992, LANC, det. A. J. Silverside. 1st record since 1876 collection from same
site.

+75/3.2 X 3. SPIRAEA ALBA X S. DOUGLASII (S. X BILLARDI) *75, Ayrs.: Side of A736 at Bloak
Moss, Kilwinning, NS/36.45. J. Flanagan & K. Cruikshank, 1976, herb. A. J. Silverside, det.
A.J. S. Overgrown river shingle, R. Stinchar near Colmonell, NX/1.8. A. McG. Stirling, 1991, E,
det. A. J. Silverside. Ist and 2nd records.

775/3.6. SPIRAEA CANESCENS *46, Cards.: Rocky scrub, Allt Goch quarries, Cwrtnewydd, SN/
491.483. A. O. Chater, 1991, NMW.

+75/8.6. RUBUS PARVIFLORUS *50, Denbs.: Deciduous wood, Gresford, SJ/340.542.
K. Watson, 1992, det. V. Gordon.

+75/8.8. RUBUS PHOENICOLASIUS *4, N. Devon: Sourton Quarry, SX/523.896. L. J. Margetts,
1992, herb. W. H. Tucker. *77, Lanarks.: Pebbly shore of Canting Basin, Old Princes Dock,
Glasgow, NS/56.65. P. Macpherson, 1991, herb. P.M., det. A. Newton.

75/8.16. RUBUS BRIGGSIANUS *5, S. Somerset: Forest ride, Staple Hill, ST/23.16. L. J.
Margetts, 1991, det. A. Newton.

75/8.45. RUBUS CRUDELIS *27, E. Norfolk: Hedgerow by lane near woodland, Saxthorpe, TG/
11.32. A. L. Bull, 1992, herb. A.L.B., det. A. Newton. Extension of range c. 160 km northwards.

+75/8.55. RUBUS LACINIATUS *4, N. Devon: Car park, Tiverton, SS/960.128. B. Benfield,
1992.

75/8.98. RUBUS CURVISPINOSUS *21, Middlesex: Bushy slope and open woodland, East Heath,
Hampstead. TQ/28.86. D. E. Allen, 1992, BM, conf. A. Newton.

75/8.117. RUBUS POLYANTHEMUS 75, Ayrs.: Shingle shore near the pier, Ailsa Craig, NX/
02.99. B. Zonfrillo, 1992, GL, det. A. McG. Stirling. 1st Ailsa Craig record.

78/8.123. RUBUS SEPTENTRIONALIS *11, S. Hants.: Under gorse, Rockford Common, SU/
17.08. D. E. Allen, 1986, herb. D.E.A., det. A. Newton. 75, Ayrs.: Ailsa Craig, NX/0.9.
B. Zonfrillo, 1992, GL, det. A. McG. Stirling. 1st Ailsa Craig record.

75/8.134. RUBUS ARMENIACUS *98, Main Argyll: Side of road to Bonawe Ferry, Taynuilt, NN/-
0.3. A. McG. Stirling, 1992.

75/8.136. RUBUS HYLOPHILUS *11, S. Hants.: Among gorse, Hamble Common, SU/484.059.
D.E. Allen, 1992, BM, det. A. Newton.

75/8.142. RUBUS ULMIFOLIUS 75, Ayrs.: Near the lighthouse, Ailsa Craig, NX/024.997.
B. Zonfrillo, 1992, GL, det. A. McG. Stirling. 1st Ailsa Craig record.

75/8.156. RUBUS LEUCOSTACHYS *25, E. Suffolk: By path through woodland, Martlesham,
TM/257.464. A. L. Bull, 1991, herb. A.L.B., conf. A. Newton. *27, E. Norfolk: Secondary
woodland, Pretty Corner, Sheringham, TG/15.41. A. L. Bull, 1992, herb. A.L.B., conf.
A. Newton.

75/8.172. RUBUS MUCRONATOIDES *42, Brecs.: Deciduous wood, Llandefaelog-fach, SO/
028.324. M. Porter, 1990, herb. M.P., conf. A. Newton.

PLANT RECORDS 285

75/8.206. RUBUS ANISACANTHOS *27, E. Norfolk: Tilia cordata wood, Hockering Wood, TG/
0.1. A. L. Bull, 1992, herb. A.L.B., det. A. Newton.

75/8.224. RUBUS BLOXAMII *42, Brecs.: Ride in plantation, Crychan Forest N. of Glan Dulas,
SN/87.42. M. Porter, 1991, herb. M.P., det. A. Newton.

75/8.270 RUBUS HYLOCHARIS *11, S. Hants.: Wood margin, Johnston’s Coppice, Purbrook,
SU/689.078. D. E. Allen, 1992, BM, det. A. Newton.

_ 75/8.305. RUBUS BRITANNICUS *11, S. Hants.: Gundymoor Wood, Havant, SU/69.07. D. E.
Allen, 1992, BM, det. A. Newton. *27, E. Norfolk: Wood margin, Wolferton Park, TG/16.31.
A. L. Bull, 1992, herb. A.L.B., det. A. Newton.

75/8.313. RUBUS NEMOROSUS *42, Brecs.: Riverbank, Llanhamlach, SO/078.274. M. Porter,
1991, herb. M.P., conf. A. Newton.

75/9.2. POTENTILLA PALUSTRIS 5, S. Somerset: Elworthy, ST/065.240. P. R. Green, 1992. 2nd
record.

75/9.13 X 15. POTENTILLA ERECTA X P. REPTANS (P. X ITALICA) *73, Kirkcudbrights.: Steep
grassy bank, Little Raebury, NX/704.436. O. M. Stewart, 1991, E, det. D. R. McKean.

+75/18.3. ACAENA OVALIFOLIA *98, Main Argyll: Forest roadside above Benmore Gardens,
NS/137.856. B.H. Thompson, 1990, conf. P. F. Yeo.

75/19.10b. ALCHEMILLA FILICAULIS subsp. VESTITA 46, Cards.: Neutral grassland in meadow by
Ddol-wen, S.E. of Aberaeron, SN/467.611. A. O. Chater, 1992, NMW. 2nd record.

75/21.4 X 11. ROSA ARVENSIS X R. STYLOSA (R. X PSEUDORUSTICANA) *29, Cambs.: Narrow
ride, Triangle Ride, Hayley Wood, TL/291.532. C. D. Preston & A. L. Primavesi, 1992, CGE, det.
A.L.P. |

75/21.4 X 12. ROSA ARVENSIS X R. CANINA (R. X VERTICILLACANTHA) *29, Cambs.: Parish
boundary hedge S.E. of Wood Farm, Hardwick, TL/365.571. C. D. Preston & A. L. Primavesi,
1992, CGE, det. A.L.P. *46, Cards.: Roadside bank N.W. of Blaen-twrch, SN/677.501. A. O.
Chater, 1990, NMW, det. A. L. Primavesi.

75/21.5. ROSA PIMPINELLIFOLIA *29, Cambs.: Swaffham Prior, TL/5.6. Dr Jermyn, 1826 &
25.6.1827, CGE, det. A. L. Primavesi. Confirmation of records hitherto thought dubious.

*75/21.7. ROSA “HOLLANDICA’ *46, Cards.: Scrub on waste ground near site of Llanilar
Station, SN/625.753. A. O. Chater, 1992, NMW.

75/21.11. ROSA STYLOSA 35, Mons.: Rough grassy common, Snowball Common, Portskewett,
ST/499.884. T. G. Evans, 1991, conf. G. G. Graham. Trackside, Minnetts Wood, ST/453.895. T. G.
Evans, 1992, herb. T.G.E., conf. G. G. Graham. Ist and 2nd post-1930 records.

75/21.11 X 12. RoSA STYLOSA X R. CANINA (R. X ANDEGAVENSIS) *29, Cambs.: Side of ditch,
Fen Road, Bassingbourn, TL/325.449. P. D. Sell, 1982, CGE, det. A. L. Primavesi. *35,
Mons.: Streamside and grassy bank, S. E. Carleon below Christchurch, ST/342.897. T. G. Evans,
1990, herb. T.G.E., det. G. G. Graham. Ist Welsh record.

75/21.11 X 14. ROSA STYLOSA X R. OBTUSIFOLIA *29, Cambs.: Hedge N.W. of Park Farm
- Cottages, Stetchworth, TL/660.565. C. D. Preston & S. E. Yates, 1992, herb. A. L. Primavesi, det.
Ale P..

75/21.12 X 13a. ROSA CANINA X R. CAESIA subsp. CAESIA (R. X DUMALIS) *46, Cards.: Hedge
S. of Blaenpistyll, Tremaen, SN/232.475. A. O. Chater, 1985, NMW, det. A. L. Primavesi.

75/21.12 X 13b. ROSA CANINA X R. CAESIA subsp. GLAUCA (R. X DUMALIS) *28, W. Norfolk:
Edge of track N. of Hockwold cum Wilton, TL/739.893. C. D. Preston, 1992, det. A. L.
Primavesi. *35, Mons.: Lower White Castle, SO/38.16. T. G. Evans, 1992, herb. T.G.E., det.
G. G. Graham. *47, Monts.: Hedge, Llanwnog, SO/04.94. A. Franks & J. Clarke, 1991, det.
A. L. Primavesi.

286 PLANT RECORDS

75/21.12 X 14. ROSA CANINA X R. OBTUSIFOLIA (R. X DUMETORUM) *29, Cambs.: Hedge along
footpath between Burton End and Leys Wood, TL/626.494. C. D. Preston & A. L. Primavesi, 1992,
CGE, det. A.L.P.

75/21.12 X 18. ROSA CANINA X R. RUBIGINOSA (R. X NITIDULA) *29, Cambs.: Milton Country
Park, TL/48.62. G. M. S. Easy, 1992, CGE, det. A. L. Primavesi.

75/21.13b. ROSA CAESIA subsp. GLAUCA *35, Mons.: Parkland, E. Blackwood, ST/17.96.
R. Fraser, 1990, herb. T. G. Evans, det. G. G. Graham.

75/21.15. ROSA TOMENTOSA *35, Mons.: Narrow wood, Llantarnam Abbey, ST/305.927. T. G.
Evans, 1992, herb. T.G.E., det. G. G. Graham. 2nd post-1930 record.

75/21.15 X 16. ROSA TOMENTOSA X R. SHERARDII (R. X SUBERECTIFORMIS) *46, Cards.: Hedge,
Derlwyn W.N.W. of Tregaron, SN/662.605. A. O. Chater, 1992, NMW, det. G. G. Graham.

75/21.16. ROSA SHERARDII 9, Dorset: By old track, Stalbridge, ST/74.17. L. J. Margetts & J.
Ounsted, 1992, det. G. G. Graham. 2nd record.

75/21.16 X 19. ROSA SHERARDH X R. MICRANTHA *46, Cards.: Hedge W. of Capel Maen-y-
groes, New Quay, SN/384.589. A. O. Chater, 1992, NMW, det. G. G. Graham.

+75/22.3. PRUNUS CERASIFERA *49, Caerns.: Copse between Tan-y-groes and Tal-y-cafn
Bridge, SH/783.718. R. Lewis, 1992, NMW, det. R. G. Ellis & G. Hutchinson.

+75/22.12. PRUNUS SEROTINA *26, W. Suffolk: Scrub on grass heath, Barnham Cross
Common, TL/867.817. K. A. Beckett, 1992.

+75/22.14. PRUNUS LAUROCERASUS *49, Caerns.: Edge of woodland opposite Maenan Abbey
Hotel, SH/79.65. R. Lewis, 1992. Small copse between. Tan-y-groes and Tal-y-cafn Bridge, SH/
78.71. R. Lewis, 1992. 1st and 2nd post-1930 records.

+75/28.21. SORBUS CROCEOCARPA *47, Monts.: Trackside hedge. Brooks, SO/146.996. P. J. M.
Nethercott, 1991.

$75/28.23. SORBUS LATIFOLIA *25, E. Suffolk: Between road and dyke, Wolsey Bridge,
Wangford, TM/47.76. Suffolk Wildlife Trust member, 1990, CGE, det. P. D. Sell.

+75/32.40. COTONEASTER STERNIANUS *11, S. Hants.: Foot of gravelly bank of A35, Markway
Hill, SU/244.026. R. P. Bowman, 1992, herb. R.P.B., det. J. Fryer. *83, Midlothian: Edge of
golf course, Levenhall, Musselburgh, NT/35.73. D. R. McKean, 1988, E, det. J. Fryer.

75/35.7 X 8. CRATAEGUS MONOGYNA X C. LAEVIGATA (C. X MACROCARPA) *77, Lanarks.:
Overgrown hedge, Sunnyside, S.E. of Hamilton High Parks, NS/74.51. A. McG. Stirling, 1979:
herb. P. Macpherson, conf. B. Wurzell.

177/7.1d. ANTHYLLIS VULNERARIA subsp. CARPATICA +*77, Lanarks.: Grassy waste ground, |
Garden Festival site, Glasgow, NS/56.65. P. Macpherson, 1992, herb. P.M., det. J. R. Akeroyd (as
var. pseudovulneraria).

77/7.1e. ANTHYLLIS VULNERARIA subsp. LAPPONICA *75, Ayrs.: Coast, Ardrossan, NS/2.4.
G. A. C. Macpherson, 1963, herb. P. Macpherson, det. J. R. Akeroyd.

77/14.1. VIcIA OROBUS 75, Ayrs.: Slown’s Cairn, Glenmuck, NS/510.033. E. L. Birse, 1958.
2nd record.

+77/15.6. LATHYRUS TUBEROSUS *58, Cheshire: Canal bank, Hassall Green, SJ/771.587. J. E.
Hawksford, 1992.

+77/15.7. LATHYRUS GRANDIFLORUS *58, Cheshire: Bank of old railway, Portwood, Stockport,
SJ/904.915. E. & O. Kearns, 1992, det. G. M. Kay.

T7/15.12. LATHYRUS NISSOLIA +42, Brecs.: Roadside verge near Brecon, SO/073.275.
M. Porter, 1991. 2nd record. +58, Cheshire: Railway embankment, Wardle, Nantwich, SJ/
603.580. G. M. Kay, 1992. Only persistent population.

PLANT RECORDS 287

77/18.2a. MEDICAGO SATIVA subsp. FALCATA +*50, Denbs.: Colwyn Bay by-pass, SH/823.787.
J. A. Green, 1990, NMW.

77/19.1. TRIFOLIUM ORNITHOPODIOIDES 44, Carms.: Colliery tip, Morlais Colliery, Llangen-
nech, SN/571.021. I. K. Morgan, 1992, NMW, conf. G. Hutchinson. 2nd record.

+77/19.19a. TRIFOLIUM INCARNATUM subsp. INCARNATUM 52, Anglesey: Disturbed soil above
shore E. of Amlwch Port, SH/46.93. J. Clarke, 1991. 2nd record. *80, Roxburghs.: Turnip field
N. of Synton Mossend, NT/478.213. M. E. Braithwaite, 1992, herb. R. W. M. Corner. Ist record
except for wool aliens.

; +77/22.2. LABURNUM ALPINUM 44, Carms.: Hedge along B4333 E. of Dangoilan, SN/342.332.
I. K. Morgan, 1992, NMW, det. A. O. Chater. 2nd record.

+77/23.2. CyTISUS MULTIFLORUS *81, Berwicks.: Rocky bank above railway, Penmanshiel,
NT/796.671. M. E. & P. F. Braithwaite, 1992, herb. M.E.B.

77/26.2. ULEX GALLI 80, Roxburghs.: Bank above Rule Burn E. of Ruletownhead, NT/
619.129. M. E. Braithwaite, 1992, herb. R. W. M. Corner. 1st confirmed record for over a century.

+79/2.2. MYRIOPHYLLUM AQUATICUM *29, Cambs.: Ditch, Willingham, TL/408.703. C. D.
Preston & S. E. Yates, 1992.

79/2.4. MYRIOPHYLLUM ALTERNIFLORUM 24, Bucks.: Recently cleared pond, Naphill Common,
SU/845.964. D. Showler, 1992. 1st post-1930 record.

84/1.1 <x 3. EPILOBIUM HIRSUTUM X E. MONTANUM (E. X ERRONEUM) *39, Staffs.: Rough
grassland slope, Fauld Crater, SK/183.277. I. Brown, 1986, det. T. D. Pennington. *$3,
Midlothian: Verge of A7 N. of Fountainhall, NT/41.52. R. Learmouth, 1990, E, det. T. D.
Pennington.

84/1.3 X +8. EPILOBIUM MONTANUM X E. CILIATUM *64, Mid-W. Yorks.: Trackside, Burley-
in-Wharfedale, SE/165.473. Yorkshire Naturalists’ Union excursion, 1992, det. T. D. Pennington.

84/1.4. EPILOBIUM LANCEOLATUM 728, W. Norfolk: Gravelly garden, Stradsett, TF/669.053.
R. M. Payne, 1992. 2nd record. 38, Warks.: Dry diorite dust at pool margin in quarry,
Atherstone, SP/306.958. J. W. Partridge, 1990, WAR, det. T. D. Pennington. 2nd record. *45,
Pembs.: Slate quarry spoil, Teifi Marshes, Cardigan, SN/189.451. A. O. Chater, 1992.

84/1.5. EPILOBIUM TETRAGONUM *68, Cheviot: Margin of lake, Nelly’s Moss Lakes, NU/
081.023. G. A. Swan, 1991, herb. G.A.S., det. T. D. Pennington. 1st confirmed record.

84/4.2. OENOTHERA FALLAX 25, E. Suffolk: Verge of Viewpoint Road, Landguard Common,
TM/28.32. A. Copping, 1992, herb. E. & M. Hyde. 2nd record. *46, Cards.: Disused railway,
Llanfarian, SN/591.778. S. P. Chambers, 1992, det. J. C. Bowra.

+85/2.1. AUCUBA JAPONICA *71, Man: Trollaby Lane, SC/352.784. L. S. Garrad, 1971. Hedge,
Ballachurry, Andreas, SC/402.968. M. Devereau, 1989. Ist and 2nd records.

88/1.1. EUONYMUS EUROPAEUS *71, Man: Deep river gorge between limestone and Manx Slate
rocks, Santon Gorge, SC/297.695. B. A. Tregale et al., 1990.

789/1.1 X per. ILEX AQUIFOLIUM X I. PERADO (I. X ALTACLERENSIS) *46, Cards.: Mixed
deciduous woodland, Penglais Woods, Aberystwyth, SN/590.821. A. O. Chater, 1992.

_ 91/2.9. EUPHORBIA LATHYRIS +*73, Kirkcudbrights.: Shingle on shore, Carsethorn, NX/
991.599. O. M. Stewart, 1992. Present for at least three years here.

91/2.12. EUPHORBIA PORTLANDICA *101, Kintyre: Coastal sand dunes, Carskey Bay, NR/
661.078. M. E. Mitchell & A. J. Silverside, 1991.

793/1.1. VITIS VINIFERA *46, Cards.: Bank of Afon Rheidol N.W. of Pont Pen-y-bont,
Penparcau, SN/590.806. A. O. Chater, 1992, NMW.

288 PLANT RECORDS

+102/1.4. OXALIS EXILIS *26, W. Suffolk: Garden paths, Bacton Green, TM/035.656. —
R. Addington, 1991, herb. E. & M. Hyde.

103/1.9. GERANIUM SANGUINEUM +*46, Cards.: Grassy roadside verge, Penparc, SN/201.479.
A. O. Chater, 1992, NMW.

+103/1.17. GERANIUM MACRORRHIZUM *77, Lanarks.: Grassy waste ground, Garden Festival
Site, Glasgow, NS/56.64. P. Macpherson, 1992, herb. P.M. Spreading.

103/1.20. GERANIUM PURPUREUM +*71, Man: Garden, The Grove Rural Life Museum,
N. Ramsey, SC/444.955. L. S. Garrad, 1990. Established for over 60 years.

+106/1.1. HEDERA COLCHICA *6, N. Somerset: Steep-sided combe, Combe Lane, Glaston-
bury, ST/507.391. P. R. Green, 1992. *43, Rads.: Wood, Garth Dingle Wood, Llowes, SU/
189.420. R. G. Woods, 1992.

+107/4.2. ERYNGIUM PLANUM *77, Lanarks.: Grassy bank, Garden Festival Site, Glasgow, NS/
56.65. P. Macpherson, 1991, herb. P.M., det. D. R. McKean. Spreading.

107/6.2. ANTHRISCUS CAUCALIS 81, Berwicks.: Sandy roadside bank, Pease Bay, NT/793.707.
M.E. & P. F. Braithwaite, 1992, herb. M.E.B. Ist record since 1893.

+107/14.1. AEGOPODIUM PODAGRARIA 75, Ayrs.: Near lighthouse, Ailsa Craig, NX/024.998.
B. Zonfrillo, 1992, GL. 1st Ailsa Craig record. :

107/17.1. CRITHMUM MARITIMUM *59,S. Lancs.: Artificial shingle beach derived from builders’
rubble, Hightown, SD/295.023. V. Gordon, 1992.

107/26.3. BUPLEURUM TENUISSIMUM 51, Flints.: Side of R. Dee, Shotton, SJ/311.693. G. R.
Sloman, 1992. 1st post-1930 record.

107/34.2. CARUM VERTICILLATUM *22, Berks.: Species-rich meadow, Winkfield, SU/911.721.
J. L. Francis & A. L. Morton, 1992, RNG, conf. S. L. Jury.

108/3.3. CENTAURIUM LITTORALE *51, Flints.: Dune slacks between Point of Ayr and the
Warren, Talacre, SJ/111.851. P. Frost, 1991, det. F. Ubsdell. 1st post-1930 record.

111/3.2 X +4. CALYSTEGIA SEPIUM X C. SILVATICA (C. X LUCANA) *48, Merioneth: Thicket,
Gloddfa Road, Barmouth, SH/615.157. P. M. Benoit, 1992, NMW. Ist confirmed record.

4111/3.3. CALYSTEGIA PULCHRA *28, W. Norfolk: Hedge, Litcham, TF/889.173. K. A.
Beckett, 1992.

113/2.1. NYMPHOIDES PELTATA +*46, Cards.: Ditch in marsh near Brimstone Wildlife Centre,
Penuwch, SN/609.630. I. S. Francis, 1992. Slow-flowing meanders of Afon Aeron at Winllan,
Trefilan, SN/567.573. A. O. Chater & I. S. Francis, 1992. Ist and 2nd records.

116/2.1. ECHIUM VULGARE 73, Kirkcudbrights.: Reclaimed land, Mutehill, NX/686.486.
O. M. Stewart & A. White, 1992. Only extant locality.

116/4.3. SYMPHYTUM TUBEROSUM +47, Monts.: Churchyard, Penstrowed, SO/070.916. C. A.
Small, 1991. 2nd record.

+116/7.1. CYNOGLOTTIS BARRELIERI *25, E. Suffolk: Well established in sandy waste ground,
Lewis Lane, Stutton, TM/138.348. E. M. Hyde, 1992, herb. E. & M. Hyde, det. E. J. Clement.

+116/10.1. TRACHYSTEMON ORIENTALIS 49, Caerns.: Under trees by R. Ro near Rowen, SH/
762.717. R. Lewis, 1992. 2nd record.

116/15.9. MyosoTIS RAMOSISSIMA 73, Kirkcudbrights.: Sandy ground near shore, Mossyard,
NX/549.515. O. M. Stewart, 1992. 2nd extant locality.

+116/16.1. OMPHALODES VERNA *71, Man: Overgrown cemetery, Braddan New Cemetery,
SC/365.772. L. S. Garrad, 1991.

PLANT RECORDS 289

118/1.5 X 6. STACHYS SYLVATICA X S. PALUSTRIS (S. X AMBIGUA) *50, Denbs.: Damp
grassland, Llay, SJ/325.553. K. Watson, 1992.

7118/4. 1c. LAMIASTRUM GALEOBDOLON subsp. ARGENTATUM *48, Merioneth: Near Pen-y-Cefn,
Dolgellau, SH/722.185. R. G. Ellis, 1987, NMW. *71, Man: Small wooded area by stream,
Ballavitchell Road, Crosby, SC/328.799. L. S. Garrad, 1982. Still present in 1991. Streamside,
Agneash, Lonan, SC/432.863. L. S. Garrad, 1988. 1st and 2nd records. *80, Roxburghs.: Bank
of R. Teviot at Hornshole Bridge, Hawick, NT/533.167. M. E. Braithwaite, 1992.

118/9.1. MARRUBIUM VULGARE 44, Carms.: Grey dunes, Pembrey Burrows, SN/435.002. I. K.
Morgan & R. D. Pryce, 1992. Only extant locality.

7118/13. rac X nep. NEPETA RACEMOSA X N. NEPETELLA (N. X FAASSENII) *50, Denbs.:
Railway bank, Llandulas, SH/9.7. G. Battershall, 1992.

118/23.4. MENTHA SUAVEOLENS +*59, S. Lancs.: Grassland, Martin Mere Wildfowl Trust
reserve, SD/42.14. P. J. Wisniewski, 1991.

118/25.4. SALVIA VERBENACA +75, Ayrs.: Banks, Ardstinchar Castle, NX/086.825. A. McG.
Stirling & A. Rutherford, 1992. Only extant locality, where first recorded before 1903.

120/1.2. CALLITRICHE TRUNCATA *9, Dorset: Ornamental lake, Forde Abbey Lake, ST/
359.049. H. J. M. Bowen, 1992.

120/1.4. CALLITRICHE PLATYCARPA 25, E. Suffolk: Ditch S. of New Dyke, Shipmeadow, TM/
388.907. C. D. Preston & N. F. Stewart, 1989. 1st post-1930 record.

7121/1.6. PLANTAGO ARENARIA *4, N. Devon: In great quantity on new roadside verges of
A399, Berrynarbor, SS/567.473. W. H. Tucker, 1992, conf. B. Wurzell.

124/1.2. VERBASCUM VIRGATUM +*59, S. Lanes.: Waste ground between Higginshaw and
Manchester, SD/934.057. A. Frank & P. Tolfree, 1988.

4124/1.5. VERBASCUM PHLOMOIDES *38, Warks.: Waste ground, Longford, Coventry, SP/
34.84. J. Robbins, 1981, WAR, det. I. K. Ferguson. Waste ground, Leamington Spa, SP/324.649.
J. W. Partridge, 1986, WAR, det. P. J. Copson.

124/1. {5 X 7. VERBASCUM PHLOMOIDES X V. THAPSUS (V. X KERNERI) *29, Cambs.: Waste
ground, Hauxton Gravel Pits, TL/437.521. G. M. S. Easy & P. H. Oswald, 1992, herb. G.M.S.E.
With both parents.

4124/4.1. MIMULUS MOSCHATUS 98, Main Argyll: Coastal flush, Achnacloich, Loch Etive,
NM/959.342. B. H. Thompson, 1992. 2nd record.

7124/11.2. CYMBALARIA PALLIDA *50, Denbs.: Old wall, Eglwys Bach, SH/823.665. B.S.B.I.
excursion, 1991.

7124/13.3. LINARIA PURPUREA 75, Ayrs.: Kilwinning, NS/3.4. B. Simpson, 1975. Rubbish tip,
Doon Bridge, Dalmellington, NS/461.059. A.McG. Stirling & O. M. Stewart, 1992. Ist and 2nd
records.

+124/16.18. VERONICA PEREGRINA 2, E. Cornwall: Base of old wall, Lostwithiel, SX/145.605.
Ee Griffiths, 1991. Ist record since 1923.

7124/16.25. VERONICA LONGIFOLIA 73, Kirkcudbrights.: Bank of R. Cree S. of Creebridge,
NX/412.653. O. M. Stewart, 1992. 2nd record.

124/18.1. SIBTHORPIA EUROPAEA *9, Dorset: Moist ground in streamside wood, Hewood
Bottom, ST/363.030. B. Edwards, 1992, conf. C. D. Preston.

124/20.5b. EUPHRASIA ARCTICA Subsp. BOREALIS 83, Midlothian: Heathy grassland near Stow,
NT/46.41. M. Little, 1985, E, det. A. J. Silverside. 1st post-1930 record.

124/20.19. EUPHRASIA SCOTTICA 81, Berwicks.: Base-rich flush, Whalplaw Burn, NT/548.568.
M. E. Braithwaite & D. G. Long, 1992, herb. M.E.B., det. A. J. Silverside. 2nd record.

290 PLANT RECORDS

124/24.1. RHINANTHUS ANGUSTIFOLIUS +*35, Mons.: Artificial grassy bank, Greenmeadow
Community Farm, ST/27.96. C. Titcombe et al., 1992, det. C. T. & T. G. Evans. 2nd Welsh record.

125/2.1. OROBANCHE PURPUREA *9, Dorset: On Achillea millefolium, Small Mouth, Portland,
SY/67.75. J. Pyatt, 1992, det. F. J. Rumsey. 1st confirmed record. +*17, Surrey: On Achillea
millefolium, Royal Botanic Gardens, Kew, TQ/1.7. T. A. Cope & B. R. Spooner, 1992. Not
deliberately introduced.

128/2.4. UTRICULARIA STYGIA *98, Main Argyll: Oligotrophic loch, Corran Lochan, NS/
216.953. A. McG. Stirling & B. H. Thompson, 1990, E. Edge of mire pool, Altan Airigh Mhic
Chainnich, NN/032.112. B. H. Thompson, 1992, det. A. McG. Stirling & B.H.T. Ist and 2nd
records.

+129/1.4. CAMPANULA PERSICIFOLIA *51, Flints.: Dune grassland, Talacre Dunes, Point of
Ayr, SJ/120.849. R. G. Ellis, 1988, NMW, det. G. Hutchinson.

129/1.12. CAMPANULA TRACHELIUM +*64, Mid-W. Yorks.: Stonework of old bell pit, Grass
Wood, Grassington, SD/988.655. M. Hanson, 1991. Ist confirmed record, previous reports being
errors for C. latifolia.

+129/9.1. DOWNINGIA ELEGANS *26, W. Suffolk: New golf course, Elveden, TL/8.8. P. Gi
Lawson, 1992, conf. E. J. Clement (from photograph).

+131/3. orb X mic. SYMPHORICARPOS ORBICULARIS X S. MICROPHYLLOS (S. X CHENAULTII) *46,
Cards.: Bank of Nant Ardal, Llanilar, SN/623.751. S. P. Chambers, 1992, NMW.

133/1.3. WALERIANELLA RIMOSA *27, E. Norfolk: Cornfield, High Kelling, TG/1.4. Mrs
Gomersal, 1939, det. G. Foggitt. Recorded here until 1945.

+135/1.2. ECHINOPS EXALTATUS *99, Dunbarton: Waste ground, Forth & Clyde Canal E. of
Temple Lock, Glasgow, NS/55.69. G. McGhee, 1989, E (photograph), det. D. R. McKean.

135/2.1. CARLINA VULGARIS *98, Main Argyll: Dry, S.-facing ledges on cliffs of raised shore
platform, Rubha Seanach, Kerrera, NM/80.25. B. H. Thompson, 1992, herb. B.H.T.

135/3.1. ARCTIUM LAPPA *46, Cards.: Rough grass and scrub, Teifi Marshes, Cardigan, SN/
182.455. A. O. Chater, 1992, NMW. :

135/3.2a. ARCTIUM MINUS subsp. PUBENS 46, Cards.: Edge of paddock E.. of St Dogmaels
Abbey, SN/168.459. A. O. Chater, 1992. 2nd record. *50, Denbs.: Roadside, Holt, SJ/401.538.
K. Watson, 1992. 1st confirmed record.

7135/11.2. CENTAUREA MONTANA 49, Caerns.: Overgrown cemetery, Llangelynin Church, SH/
771.735. R. Lewis, 1992. 2nd record and only extant locality.

135/16.3. LEONTODON SAXATILIS 98, Main Argyll: Dry, gravelly road verge, Poliphail, NR/
933.692. E. Stewart & B. H. Thompson, 1992, herb. B.H.T., conf. A. McG. Stirling. Only extant
locality.

135/22.1. LACTUCA SERRIOLA 44, Carms.: Newly built roadside verge, Machynys, SS/511.987.
I. K. Morgan, 1992, NMW, conf. G. Hutchinson. Ist record since 1912. 46, Cards.: Building site
by St Anne’s churchyard, Penparcau, SN/592.799. A. O. Chater, 1992, NMW. 2nd record.

135/26.2. CREPIS MOLLIS 81, Berwicks.: Base-rich rocky knowe, Brunta Burn, NT/595.505.
M. E. & P. F. Braithwaite, 1992, herb. M.E.B. Ist record since 1924.

135/28.25. HIERACIUM STEWARTII *93, N. Aberdeen: By R. Deveron, Ardmiddle, NJ/687.494.
D. Welch, 1991, herb. D.W., det. D. J. McCosh.

135/28.33. HIERACIUM LISSOLEPIUM *79, Selkirks.: Rocks by R. Tweed, Yair Bridge, NT/
45.32. D. J. McCosh, 1990, E.

135/28.83. HIERACIUM CAESIOMURORUM *93, N. Aberdeen: Shady rock ledge in gorge, Craig
Castle, NJ/472.247. D. Welch, 1990, herb. D.W., det. D. J. McCosh.

PLANT RECORDS 291

135/28.91. HIERACIUM OISTOPHYLLUM *48, Merioneth: Scree, Penant Dyfi, Upper Dovey
Valley, SH/9.2. P. H. Raven, 1961, BM, det. D. J. McCosh.

135/28.92. HIERACIUM SILVATICOIDES *89, E. Perth: Gorge, Allt Feith Lair, Fealar, NO/00.79.
D. J. McCosh, 1990, BM.

¥135/28.115. HIERACIUM SCOTOSTICTUM *24, Bucks.: Old beech wood, Lucas Wood, High
Wycombe, SU/874.934. A. Showler, 1992, det. J. Bevan.

135/28.142. HIERACIUM ORIMELES *86, Stirlings.: Rocks by waterfall, Loup of Fintry, NS/
66.88. D. J. McCosh, 1991, CGE, det. P. D. Sell. *87, W. Perth: Balquharn Glen, Alva, NS/
86.97. D. J. McCosh, 1989, E, det. P. D. Sell.

135/28.145. HIERACIUM SCOTICUM *102, S. Ebudes: Coastal rocks near Allt na Gile, Jura, NR/
47.78. D. J. McCosh, 1991, herb. D.J.MccC., det. P. D. Sell.

135/28.171. HIERACIUM EBUDICUM *102, S. Ebudes: Sea cliffs near Glen Trosdale, Jura, NM/
67.00. D. J. McCosh, 1991, CGE, det. P. D. Sell.

135/28.180. HIERACIUM BRITANNICIFORME *100, Clyde Is.: Base of felsite cliff, Creag Dubh,
Bennan Head, Arran, NR/997.207. A. R. Church, 1988, herb. D. J. McCosh, conf. P. D. Sell.

135/28.185. HIERACIUM EUSTOMON *46, Cards.: Base of sea cliff, Penbryn, SN/2.5. J. H.
Salter, 1928, NMW, det. J. Bevan.

135/28.191. HIERACIUM IRICUM 103, Mid Ebudes: Rough grassland, Cornaigbeg, Coll, NM/
234.630. A. Walker, 1991, herb. J. Clark, det. A. McG. Stirling. 1st Coll record.

135/28.196. HIERACIUM HEBRIDENSE *102, S. Ebudes: Waterfall in gorge, Allt nan Dearc,
Jura, NR/67.98. D. J. McCosh, 1991, herb. D.J.McC., det. P. D. Sell.

135/128.198. HIERACIUM SHOOLBREDII *72, Dumfriess.: Rock by Spoon Burn, Moffat, NT/
153.107. D. J. McCosh, 1988, CGE, conf. P. D. Sell.

135/28.209. HIERACIUM VENNICONTIUM *86, Stirlings.: Cliffs W. of Spout of Ballochleam, NS/
6.9. D. J. McCosh, 1991, herb. D.J.McC., det. P. D. Sell. *87, W. Perth: Main crags, Craig
Horn, Alva, NN/8.0. D. J. McCosh, 1988, PTH. *102, Mid Ebudes: Coastal rocks N. of Allt
Bun an Eas, Jura, NR/45.76. N. F. Stewart, 1991, E, det. P. D. Sell.

135/31.1. ANTENNARIA DIOICA *81, Berwicks.: Dry bank, Dye Water between Byrecleuch and
Trottingshaws, NT/63.58. K. Robeson & S. Williams, 1992. 2nd extant locality.

+135/40.4. SOLIDAGO GIGANTEA *$1, Berwicks.: Bank of R. Whiteadder below Paxton, NT/
93.53. M. E. Braithwaite, 1992, herb. M.E.B.

7135/41.4 x 5. ASTER NOVI-BELGII X A. LANCEOLATUS (A. X SALIGNUS) *39, Staffs.:
Grassland, Christian Fields N. of Lichfield, SK/111.113. B. R. Fowler, 1992. *73, Kirkcud-
brights.: Grassland near shore, Gillfoot, NX/976.554. O. M. Stewart, 1980, E, det. P. F. Yeo.
Banks of R. Cree below Creebridge, NX/4.6. O. M. Stewart, 1992. 1st and 2nd records.

7135/41.5. ASTER LANCEOLATUS *73, Kirkcudbrights.: Banks of R. Cree below Creebridge,
NX/4.6. O. M. Stewart, 1992.

- $135/44.1. CONYZA CANADENSIS *81, Berwicks.: Roadside verge by Chesterfield Caravan

Park, NT/772.700. M. E. Braithwaite, 1992, herb. M.E.B. 83, Midlothian: Flourishing colony
on waste ground near Victoria Dock, Leith Docks, NT/268.768. O. M. Stewart, 1989. 1st record
since 1905.

4135/44.2. CONYZA SUMATRENSIS *41, Glam.: Garden weed, Coed Glas Road, Llanishen,
- Cardiff, ST/168.817. D. Hart, 1991, NMW. list Welsh record.

4135/51.1. SANTOLINA CHAMAECYPARISSUS 41, Glam.: Roadside embankment, Penarth Road,
Leckwith, Cardiff, ST/169.738. J. P. Curtis, 1980, NMW, det. D. McClintock (as subsp. tomentosa
(Pers.) Arcangeli). 2nd record, ist since 1946.

292 PLANT RECORDS

+135/57.1. LEUCANTHEMELLA SEROTINA 25, E. Suffolk: Coastal dunes, Minsmere, TM/
478.661. E. Beaumont, 1991, det. R. M. Burton. 2nd record.

135/60.1 X 2. TRIPLEUROSPERMUM MARITIMUM X T. INODORUM *46, Cards.: Grassy slope by
sewage works, Aberystwyth harbour, SN/582.813. A. O. Chater, 1992, NMW.

+135/61.4. COTULA SQUALIDA *83, Midlothian: Weed in closely mown lawn, Royal Botanic
Garden, Edinburgh, ST/24.75. D. R. McKean, 1989, E, det. D.R.McK. & J. Cullen. A population
of female plants, apparently established for at least 30 years.

135/62.t1 X 10. SENECIO CINERARIA X S. JACOBAEA (S. X ALBESCENS) *50, Denbs.: Waste
ground, Colwyn Bay, SH/856.776. G. Battershall, 1992. *99, Dunbarton: Side of footpath,
John Street, Helensburgh, NS/29.82. A. Rutherford, 1992, E, conf. A. McG. Stirling.

135/62.10 X 11. SENECIO JACOBAEA X S. AQUATICUS (S. X OSTENFELDII) *81, Berwicks.: With
both parents on riverside, Abbey St Bathans, NT/763.617. M. E. Braithwaite, 1992, herb. M.E.B.

¥135/71.2. PETASITES JAPONICUS *75, Ayrs.: Grassy roadside bank near Knockdolia Castle,
Ballantrae, NX/12.85. A. McG. Stirling & A. Rutherford, 1992.

135/86.1. EUPATORIUM CANNABINUM 83, Midlothian: Steep, wet slope near Crichton Castle,
NT/379.613. D. R. McKean, 1992. 1st post-1930 record.

136/1.1. BUTOMUS UMBELLATUS *43, Rads.: Shallow water at edge of R. Wye, Cabalva, Clyro,
SO/233.457. D. R. Drewett & D. C. Boyce, 1992.

138/2.1. STRATIOTES ALOIDES +28, W. Norfolk: Pond in chalk pit, Wells-next-the-sea, TF/
929.429. P. R. Banham, 1986. Muddy stream, Caldecote, TF/73.02. M. Keene, 1992. Only extant
localities.

+138/3.1. EGERIA DENSA *41, Glam.: Amongst Typha latifolia in small pond, Fairwater Park
Pond, Cardiff, ST/143.778. M. E. Gillham, 1988, NMW, det. D. A. Simpson. 2nd Welsh record.

+138/4.2. ELODEA NUTTALLII *50, Denbs.: With E. canadensis in R. Dee near Rodens Hall,
SJ/416.489. K. Watson, 1990, NMW. *73, Kirkcudbrights.: In R. Dee near Threave Island, NX/
743.624. N. F. & O. M. Stewart, 1990.

142/1.5 X 9. POTAMOGETON LUCENS X P. PERFOLIATUS (P. X SALICIFOLIUS) *28, W. Norfolk:
Dike, Hilgay, TL/642.978. H. Williamson, 1991, det. C. D. Preston.

142/1.9 X 19. POTAMOGETON PERFOLIATUS X P. cRISPUS (P. X COOPER!) *70, Cumberland:
Shallow water at edge of R. Eden, Carlisle, NY/41.56. C. D. Preston, 1989, CGE, and G. Halliday,
1992, LANC, conf. C. D. Preston.

142/1.20 x 21. POTAMOGETON FILIFORMIS X P. PECTINATUS (P. X SUECICUS) *85, Fife:
Abundant at edge of bay, Loch Fitty, NT/126.913. P. Hollingsworth & C. D. Preston, 1992, CGE.

+147/2.1. LYSICHITON AMERICANUS 99, Dunbarton: Marshy ground, shore of Loch Lomond
near Boturich Castle, NS/38.84. J. Mitchell et al., 1992. 2nd record.

148/2.3. LEMNA TRISULCA *78, Peebless.: Ornamental pond, Baddinsgill, NT/1.5. D. J.
McCosh, 1992.

4148/2.4. LEMNA MINUTA *46, Cards.: Garden pond, Cenarth, SN/266.417. A. O. Chater,
1992, NMW, conf. A. C. Leslie.

151/1.6. JUNCUS FOLIOSUS *99, Dunbarton: Peaty ditch, Carman Muir, Cardross, NS/36.78.
A. McG. Stirling, 1992, E.

151/1.13 x 14. JUNCUS ARTICULATUS X J. ACUTIFLORUS (J. X SURREJANUS) _*80, Roxburghs.:
Fen margin, Branxholme Wester Loch, NT/419.108. R. W. M. Corner, 1991, herb. R.W.M.C., det.
C. A. Stace.

PLANT RECORDS 293

152/3.5. ELEOCHARIS QUINQUEFLORA 2, E. Cornwall: Small coastal stream between Doyden
Point and Port Quin, SW/968.805. M. G. C. Atkinson, 1992, det. L. J. Margetts & R. J. Murphy. 1st
post-1930 record.

152/4.1. BOLBOSCHOENUS MARITIMUS *108, W. Sutherland: Shallow brackish pool, Rubha
Ruadh, Loch Laxford, NC/168.571. R. E. C. Ferreira, 1984.

152/7.2. SCIRPUS TABERNAEMONTANI *108, W. Sutherland: Phragmites australis swamp in
shallow, brackish water, Upper Invernaver Saltings, NC/709.598. R. E. C. Ferreira, 1986.

152/8.2. ISOLEPIS CERNUA *70, Cumberland: Eroded bank at top of beach, Silecroft, SD/1.8.
M. M. Gill, 1992, LANC. 1st authenticated record.

4152/11.2. CYPERUS ERAGROSTIS *58, Cheshire: Shallow pool in old railway sidings, Crewe, SJ/
* 705.555. J. E. Hawksford, 1992, herb. G. M. Kay, det. E. J. Clement.

152/13.2. RHYNCHOSPORA FUSCA *108, W. Sutherland: Bog, Laxford Bridge, NC/23.46. R. E. C.
Ferreira, 1980, E.

152/16.5. CAREX OTRUBAE *108, W. Sutherland: Brackish pool, Loch a’ Mhuilinn N.N.R.,
NC/15.39. R. E. C. Ferreira, J. G. Roger & S. Angus, 1980.

152/16.10. CAREX ARENARIA 39, Staffs.: Disused railway sidings, Bushbury, Wolverhampton,
$J/916.022. B. R. Fowler, 1992, herb. B.R.F. 2nd record.

152/16.11. CAREX DISTICHA 43, Rads.: Damp peaty hollow, Newmead Farm, Llanelwedd, SO/
050.536. S. M. Gooch, 1992. 2nd record.

152/16.13. CAREX DIVISA *45, Pembs.: Damp roadside near the sea 0:5 km N. of Dale, SM/
808.066. J. W. Donovan, 1992, det. A. O. Chater.

152/16.20. CAREX ELONGATA *35, Mons.: Boggy ditch, Coed Robert Wood near Raglan, SO/
397.098. R. Fraser, 1992, herb. T. G. Evans, det. A. O. Chater & T.G.E.

152/16.23. CAREX HIRTA *108, W. Sutherland: Tall fen grassland, Eriboll Farm, NC/435.569.
R. E. C. Ferreira & J. G. Roger, 1978, E. Marsh at mouth of Strathy River, NC/835.656. R. E. C.
Ferreira, 1978. 1st and 2nd records.

152/16.24. CAREX LASIOCARPA 26, W. Suffolk: Hopton Fen N.R., TL/990.800. M. J. Ausden,
1991, conf. A. C. Jermy. Only extant locality.

152/16.25. CAREX ACUTIFORMIS 93, N. Aberdeen: Alder woodland by river, Glass, NJ/
463.403. D. Welch, 1992, ABD, det. A. O. Chater. 1st post-1930 record.

152/16.29. CAREX VESICARIA *108, W. Sutherland: Carex rostrata mire, mouth of Strathnaver
River, NC/451.503. R. E. C. Ferreira, 1980.

152/16.32. CAREX SYLVATICA 108, W. Sutherland: Edge of river, Loch Druim Suardalain, NC/
12.21. P. A. & I. M. Evans, 1992, herb. I.M.E. 2nd record.

152/16.39. CAREX LAEVIGATA $1, Berwicks.: Shaded flush by Bruntaburn Wood, NT/595.511.
M. E. Braithwaite, 1992, herb. M.E.B. ist record since 1892.

152/16.46c. CAREX VIRIDULA subsp. VIRIDULA 26, W. Suffolk: Lakenheath Poor’s Fen N.R..,
TL/702.827. M. Harding, 1992, conf. A. C. Jermy. Only extant locality.

152/16.64. CAREX AQUATILIS 108, W. Sutherland: Edge of Ledmore River, NC/22.12.P.A.&
I. M. Evans, 1992, herb. I.M.E. 2nd post-1930 record.

7153/3.1. SASA PALMATA *83, Midlothian: shady ditchside in woodland, Cramond, NT/
177.768. J. Cook & D. R. McKean, 1991, E.

153/12.4. FESTUCA ALTISSIMA *81, Berwicks.: Wooded burnside, Brunta Burn, NT/596.505.
M. E. Braithwaite, 1992, herb. M.E.B., det. D. R. McKean.

2904 PLANT RECORDS

153/12.6. FESTUCA ARENARIA *46, Cards.: Embryo dune, S. end of Ynys-las Dunes, SN/
605.928. A. O. Chater, 1992, NMW, conf. C. A. Stace.

153/12.9. FESTUCA VIVIPARA *83, Midlothian: Scree slope, Arthur’s Seat, Edinburgh, NT/
267.730. R. Saville, 1990, E, det. D. R. McKean. 1st record since 1824.

4153/12.15. FESTUCA BREVIPILA *39, Staffs.: Disused railway near Common Lane, Wom-
bourne, SO/870.924. C. B. Westall, 1991, herb. B. R. Fowler, det. M. A. Wilkinson.
153/12.1 X 413.2. FESTUCA PRATENSIS X LOLIUM MULTIFLORUM (X FESTULOLIUM BRAUNII) *69,

Westmorland: Wet area by path, Dalton-in-Furness, SD/224.739. P. Burton, 1989, LANC.

153/12.2 X 13.1. FESTUCA ARUNDINACEA X LOLIUM PERENNE (X FESTULOLIUM HOLMBERGI!) *99,
Dunbarton: Grassy pathside near Duchess Woods, Helensburgh, NS/286.833. A. McG. Stirling &
A. Rutherford, 1992, E.

153/13.1 < +2. LOLIUM PERENNE X L. MULTIFLORUM (L. X BOUCHEANUM) *2, E. Cornwall:
Arable field, Hudsland Farm E. of Kilhampton, SS/283.125. T. W. J. D. Dupree, & R. M. H.
Hodgson, 1992, det. P. J. O. Trist.

153/14.5. VULPIA UNILATERALIS *9, Dorset: Five Barrow Hill, SY/876.838. H. J. M. Bowen et
al., 1992.

153/16.2. PUCCINELLIA DISTANS 50, Denbs.: Edge of forest road, Llanfiangel Glyn Myfyr, SJ/
032.510. J. A. Green, 1989, det. G. Hutchinson. 2nd record. *80, Roxburghs.: With
Spergularia marina on gravel by A68 road, Carter Bar, NT/697.068. M. E. Braithwaite, 1992, herb.
R. W. M. Corner.

153/18.6. POA ANGUSTIFOLIA *69, Westmorland: Side of railway, Appleby-in-Westmorland,
NY/681.211 and NY/666.247. R. W. M. Corner, 1992, LANC, det. J. Edmondson. ist and 2nd
records. 70, Cumberland: Railway bank, Lazonby, NY/545.404. R. W. M. Corner, 1992,
LANC, det. J. Edmondson. *75, Ayrs.: Ailsa Craig, NX/0.9. B. Zonfrillo, 1992, GL, det.
A. McG. Stirling. *83, Midlothian: Dry grassland, Sunnybank Place, Edinburgh, NT/276.743.
A. J. Silverside & E. H. Jackson, 1989, herb. A.J.S. 99, Dunbarton: Grassy waste ground,
Erskine Bridge, NS/46.72. A. McG. Stirling, 1992, E. 2nd record.

153/20.1. CATABROSA AQUATICA *98, Main Argyll: Silty sand with freshwater seepage, bay
below Balure near Benderloch, NM/876.382. B. H. Thompson, 1992, CGE, conf. A. McG. Stirling.
Muddy, cattle-trodden ground with freshwater seepage, Bran Phuirt, Lynn of Lorn, NM/883.410.
B. H. Thompson, 1992, conf. C. D. Preston. 1st and 2nd records.

153/22.1. SESLERIA CAERULEA *57, Derbys.: Calcareous grassland on skeletal soil, Monk’s
Dale, SK/136.736. H. Buckingham, 1989, conf. R. Smith. Extension of range 80 km southwards.

153/31.2. KOELERIA MACRANTHA 75, Ayrs.: Ailsa Craig, NX/0.9. B. Zonfrillo, 1992, GL, det.
A. McG. Stirling. Ist Ailsa Craig record.

153/32.3. DESCHAMPSIA FLEXUOSA 29, Cambs.: Edge of woodland ride, Gamlingay Wood,
TL/243.536. T. C. E. Wells, 1991. Only extant locality. .

153/39.8. AGROSTIS VINEALIS 43, Rads.: Dry acidic grassland, Camnant, SO/080.835. R. G.
Woods, 1992. 2nd record.

138/40.2. CALAMAGROSTIS CANESCENS *24, Bucks.: Edge of ride through hazel coppice,
Leckhamstead Wood, SP/727.403. R. J. Hornby, L. M. Jones-Walters & J. Spencer, 1989.

153/50.2. BROMUS COMMUTATUS 42, Brecs.: Meadow N. of Llanwrtyd Wells, SN/860.487.
S. Gooch, 1990, herb. M. Porter, det. T. A. Cope. 2nd record.

153/51.2. BROMOPSIS BENEKENII 42, Brecs.: Limestone gorge N. of Cefn Coed, SO/024.087.
T. C. G. Rich, 1992. 2nd record, Ist since 1899. ;

+153/53.1. CERATOCHLOA CARINATA *2, E. Cornwall: Waste ground by track, Saltermill, SX/
430.637. A. Atkinson, 1991, conf. L. J. Margetts.

PLANT RECORDS 295

153/54.1. BRACHYPODIUM PINNATUM *49, Caerns.: Limestone grassland, Great Orme, SH/7.8.
P. W. Ball, 1959, LIV. 1st confirmed record. +83, Midlothian: Heath near limestone ballast
from railway, Cobbinshaw Loch, NT/019.591. R. Saville, 1992, E, det. D. R. McKean. 2nd record.

153/59.4. HORDEUM SECALINUM 46, Cards.: Roadside hedgebank W. of Llanfihangel-y-
Creuddyn, SN/658.761. A. O. Chater, 1992, NMW. 2nd record.

+153/66.1. CYNODON DACTYLON *39, Staffs.: Well established on former railway sidings,
Wolverhampton, SJ/916.022. B. R. Fowler, 1992, herb. B.R.F.

153/67.1 X +3. SPARTINA MARITIMA X S. ALTERNIFLORA (S. X TOWNSENDII) *98, Main Argyll:
Silty tidal inlet, An Sailean, Benderloch, NM/895.402. B. H. Thompson, 1990, det. T. A. Cope.
Plants recorded in 1984 at NM/888.418. and 902.418. as S. anglica were presumably this taxon.

154/1.1d. SPARGANIUM ERECTUM Subsp. OOCARPUM *46, Cards.: Ditch at edge of Typha swamp,
Pentwd Marshes, Cardigan, SN/184.449. A. O. Chater & A. P. Fowles, 1990.

158/7.1. COLCHICUM AUTUMNALE [73, Kirkcudbrights.: Delete record published in Watsonia
19: 151 (1992); plant is Crocus nudiflorus.| )

158/20.2 X +3. HYACINTHOIDES NON-SCRIPTA X H. HISPANICA 50, Denbs.: Grassland, Bryn
Euryn, Colwyn Bay, SH/832.798. G. Battershall, 1992.

4158/24.4. ALLIUM NEAPOLITANUM *27, E. Norfolk: Grassy hedgebank, Eaton Hill, Norwich,
TG/20.06. E. T. Daniels, 1991. Known here for 25 years. *71, Man: Hedgebank, Bride Hills,
NX/447.005. M. Devereau, 1991. Known here for several years.

4158/24.9. ALLIUM PARADOXUM 73, Kirkcudbrights.: Bank of R. Urr, W. of Haugh of Urr,
NX/801.660. O. M. Stewart, 1992. 2nd record.

158/24.11. ALLIUM OLERACEUM 58, Cheshire: River bank, Catton Hall near Frodsham, SJ/
545.773. G. M. Kay, 1992. 2nd record.

7158/24.12. ALLIUM CARINATUM *53, S. Lincs.: Saxilby Road, Skellingthorpe, SK/92.72.
M. Hill, 1992, conf. I. Weston. [58, Cheshire: Delete record published in Newton, A. (1991)
Supplement to Flora of Cheshire p. 39; plant is A. oleraceum. |

158/24.13. ALLIUM AMPELOPRASUM Var. BABINGTONII +*50, Denbs.: Railway bank, Llandulas,
SH/918.783. G. Battershall, 1992. 1st record of species.

158/24.14. ALLIUM SCORODOPRASUM 99, Dunbarton: Shore N. of Ardmore Point, NS/32.79.
A. Rutherford, 1990. 2nd record.

7159/5.2. IRIs SIBIRICA 77, Lanarks.: Damp wood, Busby, NS/58.55. P. Macpherson, 1991,
herb. P.M., det. B. Wurzell. 2nd record.

+159/5.4. IRIS VERSICOLOR *6, N. Somerset: Pond, Emborough, ST/613.507. P. R. & I. P.
Green, 1992.

7159/8.3. CROCUS NUDIFLORUS *70, Cumberland: Grass verge of path near Rose Castle, S. of
Dalston, NY/372.465. E. E. Marper, 1992, LANC. *73, Kirkcudbrights.: Mill Island, R. Cree
N. of Newton Stewart, NX/409.662. J. McCleary, 1989.

+159/13.1. CROCOSMIA PANICULATA *70, Cumberland: Muncaster Park, SD/090.958. M. M.
Gill, 1992, LANC, det. O. M. Stewart.

7159/13.3. CROCOSMIA POTTSII *98, Main Argyll: Streamside near Scout Cottage, Lunga, NM/
809.084. B. H. Thompson, 1992.

162/14.1. ANACAMPTIS PYRAMIDALIS 58, Cheshire: Roadside embankment, Ringway, SJ/
807.836. J. McHarry, 1992. Only extant locality.

162/18.3 xX 5. DACTYLORHIZA INCARNATA X D. PURPURELLA (D. X LATIRELLA) *80, Rox-
burghs.: With parents in calcareous flush, Murder Moss, N'T/503.286. R. W. M. Corner, 1991, det.
I. Denholm (from photographs).

Watsonia, 19, 297-303 (1993) 297

Book Reviews

Random-access guide to selected British Hawkweeds. J. Bevan, based on a program by C. J. Legg;
illustrated by K. J. Adams. Pp. 24 + disk. National Museum of Wales & Field Studies Council,
Cardiff & Preston Montfort. 1992. Price £15 (ISBN 0—7200—0367-9).

The development of computer-assisted keys to critical groups of British plants has accelerated in
recent years; see also the review in Watsonia 19: 199-200 (1993) of the FLORA program. Despite
the modest reference in its title to ‘selected’ British Hawkweeds, this program actually covers 251
taxa of British species of Hieracium (excluding Pilosella), three more than are recognised in Kent’s
List of vascular plants of the British Isles (see review in this issue). Colin Legg’s program, a version of
which also exists for the genus Carex, is fast becoming a general-purpose tool which can handle data
from different genera using standardised procedures.

Having obtained the review copy only a few days before helping to run a practical class for first-
year biology undergraduates, we were able to give it a baptism of fire by testing its effectiveness
among a number of first-time users. The results were reassuringly good; both the software and the
data scoring have evidently been thoroughly tested. Terminology is explained in a well-illustrated
leaflet which accompanies the program; both macroscopic and microscopic characters are used.
Glandular hairs, which are so effective for differentiating the species of Doronicum sect.
Macrophylla in south-west Asia, are equally valuable here in providing useful characters from both
phyllaries and peduncles. Geographical data are coded into twelve units, six of which cover
Scotland; 16 habitat types and three altitudinal zones are also covered.

Perhaps the most sophisticated feature of the program, and one which greatly contributes to its
effectiveness, is the system of the character weighting. Each of the 324 character states is rated
according to how well it differentiates between species, allowing the program to place more weight
on ‘good’ characters when computing a coefficient of similarity. Results are presented either in
descending order of number of simple matches or of weighted matches, the choice being left to the
user. Having obtained a list of provisional results, one can also display the characters which are most
effective to differentiate the top 20 species in the list.

Once one has entered data from an unknown specimen, there are several ways of interpreting the
results. These include: identifying and describing the species most similar to the unknown one;
tabulating characters for the top-ranking species; comparing pairs of species; and matching the
character states of the unknown species with those of the species chosen by the program as the one it
most closely resembles.

The booklet concludes with lists of subjective and unhelpful characters, and a short bibliography.
The key does assume that the user can recognise a member of the genus Hieracium, and provides no
means of excluding other Compositae other than Pilosella. The section on getting started ends with
the very sensible advice that it is essential to check identifications against specimens named by
experts in well-curated herbaria. No herbarium with a computer which runs MS-DOS should be

without this program. A version also exists for BBC computers.

J. R. EDMONDSON
The Orchid book: a guide to the identification of cultivated orchid species. Edited by J. Cullen. Pp.
xxxvi + 529; illustrated. Cambridge University Press, Cambridge. 1992. Price £24.95 (ISBN 0-521-
41856-9).

Basically an identification guide to the genera and species of orchid in general cultivation, this book
is based on the Orchidaceae account published in The European Garden Flora 2, but with updated

298 BOOK REVIEWS

text. It is essentially a work of reference with about 900 species briefly described. The selection of
which species to include has been governed by the criterion that on the whole they should be
representative of general European collections of cultivated orchids, and occasionally of others held
elsewhere in the world. Consequently only a small proportion of those which have at one time or
another been in cultivation has been covered.

An introductory section outlines the basic structure of the orchid flower, the growth habit of the
plant, and makes brief reference to cultivation techniques. Next is a set of keys which firstly divide
the family into five apparently separate morphological groups and subsequently into their respective
genera. The main section follows and comprises descriptive text of the selected genera and species.
It is arranged in the format of a Flora with a brief summary given for each genus including
information on geographical range, cultivation conditions and important literature references.
Diagnostic characters for each genus are usually briefly stated and may be followed by sub-keys
which break them down into their component species. These are then summarised under a series of
standard headings which cover descriptions of floral structures, habit, climatic hardiness, native
distribution range, synonyms, and also include a list of literature sources where illustrations may be
found. For some species, where floral description is difficult, there are small accompanying
sketches, but other illustrations in the book are limited to just 16 colour photographs.

Published so as to be available for the World Orchid Congress held in Glasgow in 1993, it is
primarily aimed at the orchid grower and will be of only limited interest to the British botanist. To.
the latter its main value lies in the insight it gives into the more exotic species occurring world-wide,
and in the most useful reference list to orchid illustrations. Its treatment of native British and
European orchids is very basic and brief; for example in Dactylorhiza, D. majalis is listed without
reference to any of the distinctive components of the aggregate, whilst D. sambucina is dismissed in
less than 50 words. Whilst the keys are no doubt capable of separating the selected species covered
within the book, they are very brief and generalised and likely to be of only limited use to practical
botanists. Nevertheless, it is a well-produced book in which great attention has been given to detail
both in the text and floral sketches. It should become an essential source of reference for both
amateur and specialist grower.

M. 3-Y =. FOLey

Stewart and Corry’s Flora of the North-east of Ireland, 3rd edition. Edited by P. Hackney. Pp. 419,
with 17 figures, nine black and white plates and 14 colour plates. Institute of Irish Studies, The
Queen’s University, Belfast. 1992. Price £17.50 (ISBN 0-85389-446-9).

The first edition of this Flora (1888) was mainly the work of Samuel Alexander Stewart. Thomas
Corry was tragically drowned while botanising on Lough Gill some years previously but since he had
jointly initiated work on the flora of the three North-east counties of Down, Antrim and
Londonderry, his name was added as joint author and the Flora published with financial help from
the Corry family. Stewart told the story of the tragic drowning in his introduction to the Flora, and
paid handsome tribute to the young man who was Babington’s assistant at Cambridge and who
seemed certain to make a significant contribution to both British and Irish botany. In the second
edition of the Flora (1938) Robert Lloyd Praeger paid tribute to Stewart’s critical and painstaking
approach to the work, noting that he had waged “‘ruthless war on imposters and claimants without
sufficient credentials” and provided a really reliable list of the local flora, “‘so accurate indeed that
the ensuing fifty years . . . has revealed very little that must now be withdrawn — a remarkable
tribute to the compiler’.

This third edition is firmly rooted in tradition. Prominence is given to the old records and the aim
has been to update, verify and assess the status of each species. 280 pages in 2-column format have
been given over to the catalogue, allowing for a generous inclusion of localities, annotations and
comments. Bryophytes have been omitted from this edition. The now standard practice of
recruiting specialists to write the accounts of critical groups or including only those records critically
examined by specialists, duly acknowledged, has been applied.

Part of Praeger’s ‘Botanist’s Guide’ to the area from the second edition of the Flora has been
retained. It is supplemented by further chapters by Paul Hackney on the topography of the area and

BOOK REVIEWS 299

its outstanding botanical features, as well as comparisons of the flora with that of the rest of Ireland
and with South-west Scotland (the Mull of Kintyre is only 20 km away from the Antrim coast).
There are chapters on the climate by Nicholas Betts and on the history of the vegetation by J. R.
Pilcher, each containing much that is of interest to the general reader.

The northerly position and relatively mild climate of the area are reflected in the flora. Atlantic
species such as Trichomanes speciosum and Hymenophyllum tunbrigense manage to survive while
Geranium pratense, Gymnocarpium dryopteris, Carex magellanica and Carex pauciflora are
confined to this north-east corner of Ireland. Among the surprising absentees from this large and
complex area is Gentianella amarella, common on the southern Irish dunes and extending to the far
north of Scotland.

The appendix of doubtful records, casuals and species without good credentials has been
dispensed with. All records are included in one list but with appropriate comments. Widely
cultivated species such as forestry and hedging species are included. Stewart might not have
approved of these departures. The almost smiling face of this tough little man, whose only fault
according to Praeger was his ““overweening modesty”’’, greets the reader on p. 8. He would, I feel
sure, have quickly overcome his earlier reservations and heartily approved of this splendid
production which is remarkably low-priced for the quality of paper, printing and binding and sets a
high standard for Irish regional Floras to follow.

D. SYNNOTT

Verbreitungsatlas der Farn- und Bliitenpflanzen Kdrntens. H. Hartl, G. Kniely, G. H. Leute, H.
Niklfeld & M. Perko. Pp. 451, 16 colour plates and c. 2450 maps. Verlag des Naturwissenschaft-
licher Verein fir Karnten, Klagenfurt. 1992. Price 360 Austrian Schillings (ISBN 3—85328-000-S5).

The history of botany in this most beautiful part of Austria goes back to, at least, the 1700s and the
time of Franz Xaver Freiherr v. Wulfen, commemorated by Jacquin in Wulfenia carinthiaca. Since
then, there has been continuous activity in the study of the plants of Carinthia’s high mountains, its
forests, lakes and valleys. With an area of about 10000 sq km and a flowering plant and fern total of
almost 2500 species it has, for its size, a very rich and varied flora. And a glance through the colour
photographs of this finely produced book shows how the combination of a scenic environment and
interesting or beautiful plants blend to make Carinthia a place of special attraction for the field
botanist. But this is not a coffee-table book to be browsed over and laid down. It is a scholarly work
stemming from the distillation of much scientific knowledge gleaned over more than two centuries;
its presentation has also been meticulously planned. The period from inception, gestation to
appearance covered almost 25 years and the impressive list of European botanists who were
involved testifies to its impeccable credentials.

The main aim of the book is to present by computerised dot-maps, together with complementary
symbols and notes, the present-day Carinthian distributions and status (i.e. endangered, synanthro-
pic, alien, doubtful status, etc.) of all its fern and flowering plant species. The province is divided
into c. 88 grid squares. The text covers c. 40 pages of introduction, botanical history, geography;
there are twelve pages of colour plates, twelve to a page, 307 pages of dot maps, arranged
alphabetically by genus (which makes for easy reference) and about 70 pages of very useful
supplementary information covering particular species, hybrids, literature and local names.

It is unlikely to be a best-seller among British botanists, but for those who do buy it or can
otherwise consult it, it will act as an incentive to get the reader out to Klagenfurt, the Worther See
and the Karawanken.

Any book devoted to an artificial area is likely to have a built-in likelihood of being insular in
concept and of limited readership appeal. The criticism could be made here, but nevertheless it is a
significant contribution to botanical knowledge in a floristically important part of S.E. Europe.

The Klagenfurt Natural History Society deserve congratulations for supporting the project and
seeing it through to such a satisfactory conclusion; at 360 Austrian Schillings (c. £23) it is not, by
present standards, expensive.

I. C. HEDGE

300 BOOK REVIEWS

An illustrated guide to Fungi on wood in New Zealand. 1. A. Hood. Pp. 424. Auckland University
Press, Auckland. 1992. (Distributed outside New Zealand by Oxford U.P.) Price £19.50 (ISBN 1-
86940-063-1). .

I write this review from the standpoint of a ‘consumer’. The book will be of particular interest to an
experienced amateur mycologist like myself who is about to visit New Zealand on a natural history
tour, because it is aimed at ‘‘amateur naturalists in general”. In this respect, it achieves its purpose
admirably.

The acknowledgment in the Foreword that there is a need for a much better appreciation of the
indispensable role played by fungi in the ecology of the New Zealand forests is pertinent, as this
matches a currently developing change of focus in the study of fungi in Europe. There is an excellent
Introduction to the book, comprising sections on historical background, biology and ecology,
amongst others. Very clear, concise accounts of the processes of reproduction and dispersal, growth
and nutrition, hosts and distribution are given with minimal resort to technical language. This is
most encouraging for the non-specialist reader.

The accounts incorporate appropriate references to particular species illustrated in the text. The
pen-and-ink drawn figures are delightful. Producers of other field guides might learn from these.
Coloured drawings are not always an advantage and may indeed be counterproductive for the
beginner. The figures are laid out spaciously with details of the fungi clearly identified; eight pages
of colour plates, each with six small (occasionally too small) images, are included. The text is equally
well presented and illuminating. The keys are adequately explained, and are easily followed. In
conclusion, this book can be highly recommended for the amateur naturalist.

R. Coox.

The botanizers. Amateur scientists in nineteenth-century America. E. B. Keeney. Pp. xii + 206, with
eleven black and white illustrations. University of North Carolina Press, Chapel Hill & London.
1992. Price $29.95 (ISBN 0-8078—2046-6).

No, this is not a history of the B.S.B.I.’s counterpart in America (not that there is any quite similar
body there): it is a general study of amateur botany in the United States during the period when, in
the author’s words, it “‘rapidly became the most popular science in America for recreational and
pedagogical purposes”’. Oddly, despite this, and despite the profusion of historians of science on the
other side of the Atlantic and the multiplicity of their doctoral theses (of which this book is itself the
outgrowth of one), the subject has never previously come in for the degree of scholarly attention it
deserves. This reflects the much greater emphasis long placed over there on professional status and
the proportionately weak survival of an amateur scholarly tradition — in striking contrast to Britain
and the Netherlands. As a result, as the author points out, American historians “have often
overlooked the nearly self-evident fact that amateurs and professionals pursue science for different
reasons .. . and the influence of those differences on the science that these professionals and
amateurs pursued.’’ People whose prime motive is self-cultivation are ipso facto not of much
interest to historians of science, who tend in any case to think that too few of them left writings to
make it feasible to study them as a collective entity. That this is a serious misconception Elizabeth
Keeney amply demonstrates: quite apart from the records and publications of societies there is “‘a
vast array of commonly untapped sources heavily laden with information on amateur science’,
ranging from popular journals to advice manuals and works of fiction. These she has energetically
quarried, with results that are eminently readable.

Despite the alien context, B.S.B.I. members will find much in these pages that is very familiar, for
amateur botany on the two continents seems to have run broadly in parallel. At the same time they
will be surprised to find what continue to be normal and everyday field botany practices portrayed as
quaint, bygone Victoriana. The author appears to have seen or talked to no present-day plant
taxonomists, either amateur or professional, and has consequently failed to realise that the world
she describes is by no means the irrecoverably antique one that she supposes.

The volume is attractively produced, with a number of well-chosen, out-of-the-way illustrations

BOOK REVIEWS 301

(two photographs of the Josselyn Botanical Society on an outing in Maine in 1896 are particularly
appealing), numerous source-notes and an exhaustive bibliography.

D. E. ALLEN

List of vascular plants of the British Isles. D. H. Kent. Pp. xvi + 384. Botanical Society of the British
Isles, London. 1992. Price £11.50 (ISBN 0—90115-821-4).

This is a numbered list of more than 4 200 taxa of vascular plants “known at the present time to be
native to or introduced and established in” the British Isles. For each taxon there is a tripartite
number for family, genus and species; subspecies are indicated by letters and hybrids by the
numbers of their parents. Many synonyms are given but the list is not comprehensive. The inclusion
of basionyms is a welcome innovation; it also provides some startling facts. Did you know, for
example, that Symphoricarpos albus was originally thought to be a Vaccinium? or that Lysimachia
terrestris was first described as a mistletoe? :

_ The arrival of Kent’s List was keenly anticipated and has fully lived up to expectations.
Admittedly, much of what is new, such as the classification and nomenclature, had already appeared
in Stace’s New Flora (reviewed in Watsonia 19: 161-163 (1992)), as had the use of tripartite numbers
rather than Dandy’s bipartite system, but it is only in Kent’s List that the full impact of these changes
becomes apparent. Rigorous application of the rules of priority and “‘the attempt . . . to take into
account taxonomic studies in Europe as a whole when determining the status of British and Irish
representatives” has resulted in wholesale name changes. Such changes, found in both Kent and
Stace, are rarely welcome but the majority must be accepted if we are ever to reach a stable
nomenclature. Others are still a matter of opinion — do you prefer to split Scirpus into seven genera.
as in Kent and Stace, or to leave them united as in Dandy?

There is no doubt that Kent’s List is an essential companion that must be on the bookshelf of all
botanists, amateur or professional, who have an interest in the Bnitish and Irish flora. Ido, however,
have a few quibbles. Why, for instance, are extinct aliens excluded? They are present in many
herbaria and will no doubt be included in many Floras and Checklists; if one of the purposes of the
List is to aid herbarium or Flora arrangement (as Dandy’s List was stated to be) then I would have
preferred to see them in. The omission of some important crop plants such as Hordeum vulgare is
especially puzzling. Given the choice, I would have included all the aliens numbered in Stace. This
would at least have resulted in a standard numbering system for both books rather than the
confusing double system we have now. As early as family 21, the numbering in Kent and Stace
diverges when Kent omits the Taxodiaceae, the Redwood family. Numbering of genera within
families and species within genera is similarily inconSistent and the decision by Kent not to give
unique numbers to any hybrids, although given to some of the commoner ones in Stace, adds to the
inconsistency. A useful feature found in Dandy but omitted here is the provision of genus numbers
in the index.

This is the first B.S.B.I. publication to have been produced by computer and yet it omits much
useful information that one imagines could have been included almost at the proverbial ‘touch of a
button’! Why, for instance, were the genus and species numbers and even the genus names not
repeated at the top of each left-hand page when a genus was split over two pages? It would make
consulting the List so much easier for the user and would not have taken much extra effort. Less
important but still annoying (to me at least) are the many instances of a single line of a two- or multi-
‘lined entry found at the top or bottom of a page. Similarly I prefer not to see authors’ initials and
surname, and genus initial and species name split over two lines or.even over two pages. With a little
extra effort in adjusting spaces (there is an awful lot of blank space on each page) these visual
imperfections could have been avoided. Another small point concerns the printing of hybrid names
and formulae — a multiplication sign (<) looks so much better than an x, and could have been
substituted easily.

Proof-reading appears to have been well done; spelling and punctuation mistakes are relatively
few, although on page 45 the specific epithet decalvans is twice misspelt decalvens and is once,
incorrectly, preceded by a hybrid sign. It also appears that the computer has occasionally slipped a
synonym into the wrong species, the most amusing instance is finding Cotoneaster humifusa listed as

302 BOOK REVIEWS

a synonym of Taraxacum hamatiforme not only on page 236, but also (by implication) in the index.
In a book such as this, which is relied upon to give the correct name, authority and synonym, it is
slightly worrying that so obvious an error was not spotted.

Despite the above points, I already find Mr Kent’s List an invaluable source of information which
I am referring to constantly. British and Irish botanists will remain in his debt for many years to
come.

R. G. EL.is

Atlas écologique des fougeres et plantes alliées — illustration et répartition des Ptéridophytes de France.
R. Prelli & M. Boudrie. Pp. 277 with 124 black and white plates. Editions Lechavalier, Paris. 1992.
Price F.fr. 260 (ISBN 2—225-82527-0).

If ever a book arrived at a good time for reviewing, this was it. Two days before I left for France was
more than enough time to convince me that this was the one book I must take. It now meant that I
had something more manageable than Atlas Florae Europaeae or the Atlas of North European
Vascular Plants when setting off to look for pteridophytes in a part of France that was new for me.

The book is not an identification manual as such as it contains no keys, although good descriptions
accompanied by beautiful photographs and distribution maps make this an ideal field guide for the
pteridologist in France. Prelli’s excellent Guide des Fougéres et Plantes Alliées (reviewed in
Watsonia 16: 98-99, 1986, lst ed.; Watsonia 18: 442, 1991, 2nd ed.) was written instead as an
illustrated key for identifications. Most of the commoner species are also found in Britain so English
keys will be sufficient for identification in most cases. The authors have not arranged the species in
any taxonomic or alphabetical order. Instead they have grouped them by habitat; firstly into
lowlands, mountains or Mediterranean and then subdivided either into open, woodland or wetland
species or Dv the type of rock on which they live. The absolute beginner will probably find this
approach difficult to follow and a little confusing because several species grow in more than one of
the delimited habitats; the authors have placed the species in the habitats in which they are most
commonly found. Nevertheless the photographs and the brief descriptions are good and the reader
is made aware of the other most similar species. The authors have thoughtfully chosen their
photographs to illustrate whole plants or enlargements of parts of them as appropriate for each
species. Perseverance should therefore direct the patient observer to the correct result. Reference
to the accompanying maps and range descriptions will give additional support to the identifications.

The maps themselves give information about the relative abundance of the plants in each of the
French Départements and the coloured map on the back cover shows at a glance which areas are the
richest for pteridophytes. |

The authors are to be coz agratulated on producing such a fine book. Anyone heading for France
with even a passing interest in pteridophytes would do well to acquire a copy.

B. A. THOMAS:

British plant communities. Vol. 3: Grasslands and montane communities. Edited by J. S. Rodwell.
Pp. x + 540, with 36 figures. Cambridge University Press, Cambridge. 1992. Price £95 (ISBN 0-521-—
39166-0).

This is the third of the five volumes describing plant communities in Britain, and follows the same
format as the previous two volumes (reviewed in Watsonia 19: 49, 1992, and 19: 198, 1993). A
general introduction is followed by three main sections describing 13 mesotrophic, 14 calcicolous
(calcicolous meaning ‘with calcicoles’) and 21 calcifugous grassland and montane communities.
Finally, there are indices to vegetation synonymy and the occurrence of species in the communities,
and an extensive bibliography.

Rodwell’s prose is so seductive that it is easy to get side-tracked and learn-much in the process.
There are mouth-watering accounts of the mesotrophic MG3 Anthoxanthum — Geranium hay
meadows of Teesdale, the calcicolous CG1 Festuca — Carlina oceanic rock garden grasslands, and

BOOK REVIEWS 303

in the calcifugous grasslands and montane communities, the U17 Luzula — Geum tall-herb
community of mountain ledges. A particular strength of the work is the synthesis of an enormous
amount of research, relating the vegetation extensively to soil, climate and treatment.

In general, the book is well-presented, though Fig. 20 is poorly integrated into the text; it takes
time to find the start of the sub-community descriptions as the synonymy runs into them, and the
sub-communities are not numbered in the text. The tables are simpler than the confusing ones of
Volume 1, but would still benefit from labelling (e.g. ‘Constants’, ‘associates’, etc.). The maps are
ordered across rather than down the page (cf. Volumes | and 2 respectively). Most of the records on
the map for Ul6c are missing from the map for U16. ‘Subspecies’ is excluded from the names (thus
adopting zoological convention) but ‘variety’ is included.

The influences of the sampling and analysis should have been dealt with in more detail. For
instance, data for the CG1f Carlina — Scilla sub-community were abstracted from the Biological
Flora for Draba aizoides whose quadrats were deliberately centred on that species, resulting in an
Over-estimation of its frequency in the tables. Over half of the samples analysed for the calcicolous
grassland chapter were collected by T. C. E. Wells et al. in a small area of Dorset, Hampshire and
Wiltshire using 1 x 1 m quadrats and including few bryophytes, resulting in under-estimations of the
frequencies of some species and the numbers of species per quadrat, and localized clusters of
records on the maps.

The practical problems of consistently identifying vegetative plants suggests that more judicious
lumping might have been adopted (e.g. Poa pratensis and P. humilis (P. subcaerulea) cannot have
been consistently separated). Oddly for the calcicolous grasslands, the recorders ‘“‘always attempted
to separate the subspecies of Cerastium diffusum . . .”’ though only one occurs in Britain.

An introduction to the three grasslands would help to place a grassland in its appropriate section,
and give the volume more coherence in its own right. There is considerably more variation in
mesotrophic grasslands than is described. For example, the grazing meadows on the levels in Kent
and Sussex characterized by Cynosurus cristatus, Hordeum secalinum and Lolium perenne are
distinctive in the field and clearly related to the MG6 Cynosurus — Lolium grasslands; in light of the
under-sampling of this community in South East England (cf. the map of samples of the
community), perhaps they should be re-examined. Grasslands heavily modified by broad-leaved
herbicides are both distinctive and common, and deserve more mention as does Elymus repens
grassland on neglected land and recent road verges (e.g. M25 embankments). Grasslands
dominated by Festuca arundinacea similar to the MG12a Festuca grassland Lolium — Holcus sub-
community occur widely inland. Any odd mesotrophic vegetation always keys out unsatisfactorily to
MG5 Centaurea — Cynosurus, MG6 Cynosurus — Lolium or MG7 Lolium grasslands at the end of
the key. Such criticisms are inevitable for the first airing of such a major work, and indicate possible
areas for revision, and it will be interesting to see whether the dissolution of metalliferous and chalk
heath vegetation types is adopted elsewhere.

For B.S.B.I. members, this book will be an invaluable reference work as a source of information
on vegetation and plant ecology. Accounts of vegetation for County Floras could be summarised
from the volumes, though widespread mapping of communities is unlikely to catch on due to the
need for bryophyte identification (especially in the north and west) and the high cost of the volumes.

After three years’ intensive application of British plant communities in the field, I remain firmly
convinced of its value and usefulness. It is a convenient, practical tool for describing and assessing
vegetation, and provides an excellent framework for understanding the relationships between
communities and the environment.

T. C. G. RIcH

counties

INSTRUCTIONS TO CONTRIBUTORS

Scope. Authors are invited to submit Papers and Short Notes concerning the taxonomy,
biosystematics and distribution of British and Inish vascular plants, as well as topics of a more
general or historical nature.

Manuscripts must be submitted in duplicate, typewritten on one side of the paper, with wide margins
and double-spaced throughout.

Tables, figure legends & appendices should be typed on separate sheets and attached at the end of
the manuscript.

Submission of manuscripts

Papers and Short Notes: Dr B. S. Rushton, Department of Biological and Biomedical Sciences,
University of Ulster, Coleraine, Co. Londonderry, N. Ireland, BTS52 1SA.

Books for Review: Dr J. R. Edmondson, Botany Department, Liverpool Museum, William
Brown St, Liverpool, L3 8EN.

Plant Records: the appropriate vice-county recorder, who should then send them to C. D. Preston,
Biological Records Centre, Institute of Terrestrial Ecology, Monks Wood, Abbots Ripton,
Huntingdon, PE17 2LS.

Back issues of Watsonia are handled by Messrs Dawson UK Limited,

Cannon House, Folkestone, Kent, CT19 5EE, 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.I., 68 Outwoods Road, Loughborough, Leicestershire, LE11 3LY.

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