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January-March 2010 Number 1072

A QUARTERLY JOURNAL OL NATURAL HISTORY LOR THE NORTH OL ENGLAND

Whiteflies (Hemiptera: Aleyrodidae) of Watsonian Yorkshire -

Chris Malumphy

Wall Ferns in East Hull, 2004-2008 - R. Middleton

Botanical Report for 2009 - Flowering Plants and Ferns -P.P. Abbott

The Ostracod Scottia pseudobrowniana Kempf, an Addition to the
Freshwater Fauna of Yorkshire, and a Note on the Evolution of
Terrestrial Habits in Ostracods and other Crustaceans -

Andy Godfrey, Geoffrey Fryer and David J. Horne

The Status of Moorland Breeding Birds in The Peak District National
Park 2004 - Peter Middleton and Geoff Carr

Published by the Yorkshire Naturalists’ Union

Editor M. R. D. Seaward MSc, PhD, DSc, FLS, The University, Bradford BD7 1DP

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A QUARTERLY JOURNAL OF NATURAL HISTORY FOR THE NORTH OF ENGLAND

Editor M. R. D. Seaward, msc, PhD, dsc,fls,
The University, Bradford BD7 1DP

Volume 135
2010

Published by the Yorkshire Naturalists’ Union

WHITEFLIES (HEMIPTERA: ALEYRODIDAE)
OF WATSONIAN YORKSHIRE

CHRIS MALUMPHY

The Food and Environment Research Agency, Sand Hutton, York Y041 1LZ
e-mail : chris .malumphy @ fera .gsi .gov.uk

Abstract

Collection details are provided for 1 1 species of Aleyrodidae (eight native and naturalised
introductions, one introduced species established on indoor plantings, and two non-
established introductions on growing plants) recorded in Watsonian Yorkshire. Two native
species ( Pealius quercus (Signoret)) and Siphoninus immaculatus (Heeger)) and one
introduced naturalized species (Aleurotuba jelinekii (Frauenfeld)) are recorded from the
region for the first time. Siphoninus immaculatus is reported damaging an ornamental ivy
plant for the first time in Britain and cultivated rose is recorded as a new host for Aleyrodes
lonicerae Walker. A list of six non-native species that have been intercepted in the region
by the plant health authorities of England and Wales on imported plant produce is also
presented.

Introduction

Whiteflies comprise a single family, Aleyrodidae, which currently contains 1556 species in
161 genera (Martin & Mound, 2007). Fifty-six species occur outdoors in Europe and the
Mediterranean basin (Martin et al., 2000). All whiteflies are phytophagous and several
species are economically important. Feeding by the immature stages reduces plant vigour
by depletion of the plant sap, foliage is contaminated with eliminated honey dew on which
black sooty mould grows, and the adults of a small number of species are important vectors
of plant viruses.

Harrison (1915, 1916, 1917, 1917a, 1920, 1920a, 1931) studied the whiteflies of the
North East of England and listed four native or naturalised species (i.e., an introduced
species that overwinters outdoors) in Yorkshire ( Aleurochiton aceris (Modeer), Aleyrodes
lonicerae Walker, Aleyrodes proletella (Linnaeus) and Tetralicia ericae Harrison). Walsh
(1918, 1920) recorded A. lonicerae, T. ericae and ‘ Aleurochiton avellanae ’ from Yorkshire
(the specific identity of the latter is uncertain and is discussed below). The purpose of this
communication is to review the whiteflies of Watsonian Yorkshire (VCs 61 to 65), based on
samples collected since 1997, published records and unpublished data held by The Food
and Environment Research Agency (Fera) .

Methods

Whitefly samples were collected on woody and herbaceous plants growing outdoors and
indoors between 1997 and 2009, mainly by the author. Puparia were slide-mounted
following standard published methods (Martin, 1987) and deposited at Fera. They were
identified using the diagnostic keys provided by Martin et al. (2000). The identity of some
of the non-native species was confirmed by Jon Martin of the Natural History Museum,
London. The nomenclature used follows Martin and Mound (2007).

Morphological descriptions and illustrations of the puparia of all native/naturalised
species listed below are provided by Martin et al. (2000) and the adults of the majority of
the species are described by Hulden (1986). The host and distribution data is obtained from
Martin et al. (2000), and Mound and Halsey (1978).

Results

Eleven species (8 native and naturalised introductions, 1 introduced species established on
indoor plantings and 2 non-established introductions on growing plants) of Aleyrodidae are
recorded here for Watsonian Yorkshire. Two native species ( Pealius quercus (Signoret) and

Naturalist 135 (2010)

Whiteflies (Hemiptera: Aleyrodidae) ofWatsonian Yorkshire

Siphoninus immaculatus (Heeger)) and one introduced naturalized species ( Aleurotuba
jelinekii (Frauenfeld)) are recorded from the region for the first time.

The white fly species have been divided into three categories below: the first includes
the native and naturalized species; the second, non-native species established under
artificial conditions (and occasionally found breeding outdoors in the summer but not
overwintering outdoors); the third, includes non-native species that have been found
breeding on imported plants but which are not known to have become naturalised or
established anywhere in Britain. The author collected all of the samples recorded under
‘Collection data’, unless stated otherwise.

For the sake of completeness, 6 non-native species that have been intercepted in the
region by the plant health authorities of England and Wales on imported plant material are
listed in Appendix 1 .

In addition to the whitefly species listed below, Walsh (1920) recorded the larvae of
‘ Aleurochiton avellanae’ in Raincliffe and Staintondale on Corylus avellana, in August and
September 1920. Aleurochiton avellanae (Signoret) was synonymised with Asterobemisia
carpini Koch by Mound and Halsey (1978), but this was not accepted by Zahradnik (1989,
1991) who considered them two distinct species. The species avellanae Signoret has also
been confused with P. quercus in Britain, for example by Trehan (1940) (Mound, 1966;
Mound & Halsey, 1978; Zahradnik, 1956). As both A. carpini and P. quercus occur on
Corylus in England and there are no preserved specimens of the whitefly collected by
Walsh available for study it is not possible to be certain which species he observed.

Native species and naturalised introductions

Aleurochiton sp. (Figures 1-2)

Host plants: Acer spp.

Biology: bivoltine; overwinters as puparia in leaf litter.

Distribution: Europe.

Comments: there are three species of whitefly that feed exclusively on Acer spp. in Europe
and they are all assigned to the genus Aleurochiton (Martin et al., 2000). Harrison (1916)
recorded A. aceris (Modeer) in great numbers on maple in Gunnergate, Middlesbrough. He
subsequently republished this record, adding that the host was field maple (Acer
campestris) (Harrison, 1920a).

Mound (1966) dismissed this
record, as he assumed it was based
only on adults. However, although
adult whitefly may be found on
non-host species, they are unlikely
to occur in ‘great numbers’,
particularly on a tree where the
foliage is relatively high. Mound
stated that the Targe size and
striking appearance of the
immature instars with their white
dorsal wax’ were not recorded by
Harrison, proving that he did not
see puparia. Aleurochiton species
in Europe, however, are bivoltine
and their puparia exhibit distinct
dimorphism: only the mature

winter puparia are conspicuous
(dark with a dense coating of white wax on the dorsal surface) (Figure 1), whereas the
summer puparia have a pale, translucent cuticle and are difficult to detect (Figure 2) (the
present author observed numerous adult A. aceris on Acer platanoides trees in July 2008 in

FIGURE 1. Aleurochiton aceris dark, waxy winter
puparium on Acer platanoides .

Whiteflies (Hemiptera: Aleyrodidae) ofWatsonian Yorkshire 5

Sawbridge worth, Hertfordshire, but found it difficult to find a single summer pupal case).
Adult Aleurochiton are only active for a brief period in the spring (when the dark, waxy,
overwintered puparia are in the leaf litter) and in the summer (when the puparia on the

foliage are inconspicuous). The
presence of A. aceris in Britain was
not confirmed until 1976 (Martin,
1978) and it is now locally common
in southern England. It feeds on
A. platanoides and is widely
distributed across Europe (Martin et
al., 2000). Aleurochiton acerinus
Haupt has also been recorded in
southern England by Dolling &
Martin (1985) on A. campestris. It
occurs widely in southern and
central Europe (Martin et al., 2000).
Unfortunately the specific identity
of the Aleurochiton recorded in
Yorkshire by Harrison cannot be
verified, as Harrison did not appear
to keep any voucher specimens. It
would not be surprising if A. aceris
was found to be present in the north of England as it occurs widely in Scandinavia and the
Baltic region (Malumphy et al., 2009), whereas A. acerinus is more common in southern
parts of Europe.

Aleurotuba jelinekii (Frauenfeld, 1867) - Viburnum Whitefly (Figures 3-4)

Host plants: appears to be mainly restricted to Viburnum tinus in Britain but on the
continent it is also recorded on Arbutus unedo, Arctostaphylos sp., Myrtus communis and
Viburnum spp..

Biology: univoltine; flight period early May to early August; overwinters as puparia.
Distribution: Europe and the USA.

Collection data: all records are on V. tinus. VC61, Elvington, 24.ii.2008, 10.iii.2009,
5.vi.2009 (pupal cases and eggs)

(leg. S. Reid) (large infestation).

VC62, Haxby, 7.iii.2009,

20.iv.2009 (abundant, wide-
spread); Howsham, ll.v.2009
(abundant); Sand Hutton, various
dates iii-viii.2009 (abundant,
widespread; adults present from
beginning of May; by June the
new plant growth was covered in
thousands of adults and eggs;
first and second instars present at
end of July); Stockton-on-the-
Forest, 24.iv.2009 (abundant,
larvae), 27.vii.2009 (adults);

Strensall, 22.iv.2009 (abundant,
widespread); York, city centre,
ll.v.1997 (leg. L. MacLeod),

23 .xi .2003, 3.iv.2006, 15.xi.2008, 18.vii.2009 (large infestations, widespread), Clifton,
7.iv.2009 (abundant), Clifton Moor, 9.iv.2009 (abundant, widespread), l.viii.2009 (first and
second instars), Homestead park, 12.vii.2009 (abundant, adults and pupal cases);

FIGURE 3 . Viburnum tinus leaf with large infestation
of Aleurotuba jelinekii

FIGURE 2. Aleurochiton aceris pale summer
puparium on Acer platanoides

Whiteflies (Hemiptera: Aleyrodidae) ofWatsonian Yorkshire

Wigginton, 9.iv.2009 (abundant, wide-
spread); Wigginton Moor, 10.iv.2009
(abundant). VC 64, York, city centre,
14.iii.2009 (small infestation), Acomb,
1 .iii.2009 (small infestation), Acomb
wood, 9.V.2009 (large infestation,
puparia and adults) .

Comments: introduced into Britain
in about 1936 (Mound, 1962, as
Aleurotrachelus jelinekii). It is now
very common and widespread in
southern England (Andrew Halstead,
pers. comm. 2009). It is recorded here
for the first time from Yorkshire and is
Figure 4. Aleurotuba jelinekii puparia with abundant in the centre of York,
dorsal wax tufts frequently occurring in very large

populations smothering the undersides
of foliage (Figure 3). It is one of the most common whiteflies found across Europe (Martin
etal., 2000).

Aleyrodes lonicerae Walker, 1852 - Honeysuckle or Strawberry Whitefly (Figure 5)

Host plants: broadly polyphagous.

Biology: multivoltine with overlapping generations; adults occur throughout the year;
overwinters in the adult stage.

Distribution: Europe and Iran.

Collection data: VC61, Allerthorpe Common, on Lonicera periclymenum, 25.vii.1998;
Beverley, plant nursery, Euphorbia pulcherrima (grown indoors), 5.xi.l998, from the
Netherlands, various dates xii.2001 (Fera), Lonicera sp., 10.ix.1998 (Fera); Dunnington
Common, Rabbit Warren, L. periclymenum, 30. vi. 2009 (adults, eggs, first instars);
Elstonwick, Rubus sp.. 19.xi.2003 (leg.

W. Dolling) (adults and wax patches);

Halsham, Fragaria x ananassa grown in
a poly tunnel, 13.x. 2000 (leg. W.

Dolling); Hull, Lonicera sp., 25 .ix .2007
(leg. T. Prior) (large infestation, all
developmental stages); Skipwith
Common, L. periclymenum, 2.viii.2009
(sparse, adults, eggs, first instars). VC62,

Bossall, west-belt wood, Rubus fruticosa,

21.iv.2009 (eggs); Buttercrambe Moor
wood, L. periclymenum and R. fruticosa,

22.x .2008 (several vacated pupal cases
and parasitised puparia), 21.iv.2009
(adults, eggs), 10.vi.2009 (abundant, all
stages); Haxby, Lonicera nitida, various
dates viii.2007 (abundant, all stages),

L. periclymenum, various dates iii.-
viii.2009 (abundant), R. fruticosa, 19.iv.2009 (eggs, larvae); Haxby allotments, F. x
ananassa, 31.vii.2009 (abundant, adults, eggs); Haxby, Westwoods, Aegopodium
podagraria, 31.vii.2009 (abundant, all developmental stages); Nunnington, Meconopsis
cambrica, 5.ix.2008 (leg. R. Hammon) (adults, eggs, first instars), Rosa sp., 28. vi. 2009
(pupal cases and adults); Sand Hutton, R. fruticosa, 1 3 .iii.2009 (sparse), Black Dike
Plantation, R. fruticosa, 19.xi.2008 (adults, vacated pupal cases, parasitised puparia),
5 .iii.2009 (adults, eggs, first instars), L. periclymenum, R. fruticosa, (adults, eggs, early

FIGURE 5 . Aleyrodes lonicerae adult with one
greyish spot on each forewing

Whiteflies (Hemiptera: Aleyrodidae) ofWatsonian Yorkshire 7

instars; abundant), Arctium lappa and Centaurea scabiosa (adults only, A. lappa and C.
scabiosa are not confirmed hosts), 20.viii.2009; Strensall, R. fruticosa, 29.ix.2008 (adults,
vacated pupal cases; adults were active, making short flights despite an ambient
temperature of -1°C); Strensall Common, L. periclymenum, 16.vii.2009 (adults, eggs).
VC64, Fountains Abbey, L. periclymenum , 25.vii.2009 (eggs, larvae); York city centre, R.
fruticosa, 8.vii.2003 (sparse, adults, eggs).

Comments: the puparia of A. lonicerae are morphologically very similar to A. proletella
(Linnaeus) and one of the main characters used to distinguish between them (the degree of
development of the caudal setae) should be used with caution, as this character exhibits
more variation in both species than reported in the literature. Several specimens should be
examined and an average character state used for identification purposes. The adults,
however, are easily separated, as each forewing of A. lonicerae bears a single grey spot
(Figure 5), whereas each fore wing of A. proletella bears a pair of spots (Figure 6).
The shape of the aedeagus also differs significantly between the two species (Hulden, 1986).

Harrison (1915, 1920) recorded this species in Gunnergate, Nunthorpe, and near
Stainton, Middlesbrough, on Lonicera sp. He subsequently reported that it was extremely
common everywhere in North Yorkshire (Harrison, 1931). Walsh (1920) recorded it in
Haybum Wyke, Raincliffe Woods and Staintondale, on Lonicera sp. during August and
September 1920. It is an occasional pest of blackberry and strawberry in Britain (Alford,
2007). Cultivated rose is recorded here as a new host for this species.

Aleyrodes proletella (Linnaeus, 1758) - Cabbage Whitefly (Figure 6)

Host plants: broadly polyphagous.

Biology: four or five overlapping generations each year; adults occur throughout the year;
overwinters in the adult stage.

Distribution: Europe, Iran, Africa, Hong Kong, New Zealand, Brazil, Mexico, Bermuda,
Puerto Rico and the Virgin Islands.

Collection data: VC61, Beverley, plant nursery, on Euphorbia pulcherrima (grown
indoors), l.ix.2002, Hebe sp. (grown
indoors), 6.viii.2003 (Fera); Elstronwick,

Brassica oleracea kale, l.xi.1998 (leg.

W. Dolling) (adults); Stamford Bridge,
allotments, B. oleracea, 23.vii.2009
(adults, eggs, larvae). VC62, Burythorpe,

B. oleracea, 20.vii.2009 (leg. J. Ostoja-
Starzewski); Haxby allotments, B.
oleracea, 31.vii.2009 (adults, eggs,
larvae); Nunnington, B. oleracea,

20.viii.2003, 2 1 .viii.2008 (leg. R.

Hammon) (abundant); Stockton-on-the-
Forest, herbaceous plant, 24.iv.2009
(abundant, adults), Chelidonium majus,

Sonchus arvensis and other herbaceous
plants, 25.vii.2009, (abundant, hundreds
of adults, eggs); York, city centre,

Tropaeolum sp., 5.vii.2009 (leg. A.

Korycinska) (adults only, Tropaeolum is
not confirmed as a host), Clifton allotments, B. oleracea, l.viii.2009 (spares, adults, eggs).
VC64, Knaresborough, plant nursery, E. pulcherrima (grown indoors), from the
Netherlands, 21.xi.2004 (Fera); Leeds, B. oleracea kale, 27.viii.1997 (leg. T. Prior)
(abundant); York, Acomb, B. oleraceae, 8.vii.2001 (abundant) (leg. R. Natt).

Comments: Harrison (1915, 1916) recorded this species in Great Ayton, on C. majus. It is a
frequent pest of Brassica spp. in Britain.

FIGURE 6. Aleyrodes proletella adult with two
greyish spots on each forewing

8 Whiteflies (Hemiptera: Aleyrodidae) ofWatsonian Yorkshire

Bemisia afer (Priesner & Hosny, 1934)

Host plants: broadly polyphagous on woody plants.

Biology: not recorded in Britain but suspected to be multivoltine if climatic conditions are
suitable.

Distribution: occurs throughout the warmer regions of the World and is widespread in the
Mediterranean.

Collection data: VC64, Huddersfield, plant nursery, on L. nobilis from Italy, ll.x.2000
(Fera); York city centre, L. nobilis (in sheltered walled private garden on a well established
tree with no recent import connection) 23 .vi .2003, 8.vii.2003 (sparse); Wetherby, plant
nursery, L. nobilis from Belgium (probably originated in Sicily), various dates between
ix.1999 to i.2001 (Malumphy, 2003).

Comments: B. afer has been introduced and become naturalised in Gloucestershire and
Greater London (Halstead, 1981, as B. hancocki (Corbett); Malumphy, 2003) but in
England has only been found breeding outdoors on L. nobilis.

Pealius quercus (Signoret, 1868)

Host plants: deciduous Betulaceae and Fagaceae.

Biology: univoltine; flight period May and June; overwinters as puparia in the leaf litter.
Distribution: north and central Europe.

Collection data: VC64, Gormire Lake, on Quercus robur, 27 .v. 1997 (adult and early larval
instar). Not found in the same location in May 2009.

Comments: W. Dolling observed adult whitefly without any wing markings on oak in Haw
Park, Wakefield, 17.V.2003, that were highly likely to be P. quercus. This species is
widespread and locally abundant in England and parts of Scotland (Mound, 1966). This is
the first time that it has been recorded in Yorkshire.

Siphoninus immaculatus (Heeger, 1 856) - Ivy Whitefly (Figures 7-8)

Host plants: Hedera helix in Britain; also recorded on H. canariensis on the continent.
Biology: univoltine; flight period from June to August; overwinters as first instars.
Distribution: central and northern Europe, and Iran.

Collection data: in all cases it was found on H. helix growing against a solid surface, for
example brick or stonewalls or a wooden fence, usually with a southerly aspect. VC61,
Elstronwick, 26.vi.2003 (leg.

W. Dolling) (adults, puparia,
eggs). VC62, Haxby, 31.vii.2009
(mature puparia, many parasitized,
foliage covered in sooty mould);

Howsham, ll.v.2009 (few first
instars, many second and third
instars, old vacated pupal cases
and parasitized puparia),

21.V.2009 (many third instars and
a few teneral puparia), 15.vi.2009
(mature puparia), 25 .vi .2009
(adult females and eggs); Castle
Howard, 2.iv.2009 (first and
second instars). VC64, Fountains
Abbey, on a variegated plant,

29.iii.2009 (first instars, vacated
pupal cases; foliage covered in
honey dew and black sooty
moulds and suffering from necrosis and premature leaf drop; vacated Syrphidae puparia
present), 27 .vi. 2009 (thousands of adults and vacated pupal cases, eggs, teneral and mature
puparia; puparia dorsal pigmentation varied from being absent, to a cepahalic and posterior

FIGURE 7. Hedera helix leaf covered in honeydew
and mould due to a large infestation of Siphoninus
immaculatus

Whiteflies (Hemiptera: Aleyrodidae) ofWatsonian Yorkshire 9

dark spot, to a broad longitudinal band; sex ratio 1 male to 4.4 females (n = 113); many
adults were caught in spiders webs but were not seen being eaten by spiders), 25.vii.2009
(abundant, hundreds of adults,
thousands of eggs, most laid on
leaves that were already heavily
infested with whitefly; three
predatory cecidomyid larvae present
among the eggs; several puparia
contained a single hymenopterous
parasitoid larva or pupa) .

Comments: occurs widely in

continental Europe and as far north
as Sweden (Gertsson, 1987; Martin
et al., 2000). It is infrequent in
southern England (Mound, 1966).

This is the first time that it has been
recorded in Yorkshire and appears
to be the most northerly published
record in Britain. This is also the
first occasion in Britain that it is reported causing damage to its host plant.

Tetralicia ericae Harrison, 1917 - Cross-leaved Heather Whitefly
Host plants: Erica tetralix in England; also E. carnea on the continent.

Biology: univoltine; flight period May; overwinters as third instars.

Distribution: Europe and Iran.

Collection data: VC62, Strensall Common, on Erica tetralix, 29 .vi. 1997 (single puparium).
Not found in 2007-9 despite extensive searching although sheep had heavily grazed the
heather plants.

Comments: Harrison (1917, 1917a, 1920a) originally described this species from
specimens collected on E. tetralix at Waldridge Fell, Durham and subsequently recorded it
from Skip with Common. G.B. Walsh and W.J. Fordham found it on every tuft of Erica at
Skipwith Common on 1 August 1918 (Walsh, 1918). It has also been recorded at
Middlesbrough (1917) (Bink-Moenen, 1989), and Hatfield Moor (1992) and Thome Moor
(1995) (Skidmore, 2006). It occurs widely in England from the south coast as far north as
Northumberland (Mound, 1966) but is rarely recorded. It also occurs widely in continental
Europe as far north as Sweden (Gertsson, 1987; Martin et al., 2000).

Introduced species established on indoor plantings

Trialeurodes vaporariorum (Westwood, 1856) - Glasshouse Whitefly
Host plants: broadly polyphagous.

Biology: multivoltine with overlapping generations.

Distribution: cosmopolitan.

Collection data: the data has been summarised due to the large number of records, mostly
from Fera; all are on indoor plantings either in plant nurseries, private premises or
laboratory glasshouses. VC61: Beverley, Halsham and Hull, on Ajuga reptans from the
Netherlands, 2008 ,Asclepias sp. from the Netherlands, 2000, Brachychiton sp. from Israel,
2000, Datura sp. from Israel, 2005, Euphorbia pulcherrima, 1997-2001, from Germany,
2003, 2007, Portugal, 1998, the Netherlands, 2001-2006, and Sweden, 2008, Fuchsia sp.,
1997, Hardenbergia violacea from the Netherlands, 2004, Hedera sp. from the
Netherlands, 2002, 2008 , Hypericum androsaemum from the Netherlands, 2008, Hypoestes
sp. from the Netherlands, 2001, Lycopersicon esculentum, 13.x .2000 (leg. W. Dolling),
Mentha sp. from Israel, 2001, Pelargonium sp. from Israel, 1997, Pyracantha sp., 2003,
Verbascum sp., 2002, and Zelkova sp. from China, 1999. VC62, Sand Hutton, on Borago

FIGURE 8. Electron micrograph of a Siphoninus
immaculatus pupal case showing the dorsal
siphons

Whiteflies (Hemiptera: Aleyrodidae) ofWatsonian Yorkshire

officinalis, 2003, Calendula sp., 2003, Camelina sp., 2003, Cannabis sativa, 2008,
Chrysanthemum sp., 2003, 2007, Fragaria sp., 2008, Lunaria sp., 2003 and Urtica dioica,
2002. VC64, Leeds, Richmond, Wakefield and Wetherby, on Aster sp. from Colombia,
1996, Lantana sp. from Italy, 1996, Lysimachia sp. from Israel 1996, Mentha sp. from
Israel, 2001, Mrytus communis from unknown origin, 2008, Origanum vulgare from Israel,
2001, Pelagonium sp., 2009, Phyla sp. from Israel, 2007, Rosmarinus officinalis from
Israel, 2003, Salvia officinalis from Israel, 2002, and Urtica sp., 2001 .

Comments: growing plants imported into England and Wales are routinely inspected
shortly after arrival by the Plant Health and Seeds Inspectorate (PHSI) . The countries listed
above refer to the origin of the plants that in most cases are the likely source of the
whiteflies. However, it is possible that some of the plants became infested after arrival in
Britain, as T. vaporariorum is very common throughout the country in commercial and
private ornamental indoor plantings. It also breeds outdoors during the summer, most
frequently in southern England and in large urban areas. Trialeurodes vaporariorum is
probably much more common and widespread in Yorkshire than the records here indicate.
It is under-recorded because it is so ubiquitous.

Non-established incursions

Bemisia tabaci (Gennadius, 1889) - Tobacco Whitefly
Host plants: broadly polyphagous.

Distribution: cosmopolitan.

Collection data: the tobacco whitefly has been found on imported plants and plant material
at a botanical garden and at plant nurseries distributed throughout the region (specific
locations are not given for reasons of commercial confidentiality as this is a regulated pest).
On Abutilon sp., Ajuga reptans, Argyranthemum sp., Asclepias sp., Begonia sp.,
Brugmansia sp.. Cassia sp., Crossandra sp., Diascia sp.. Euphorbia amygdaloides, E.
pulcherrima, Hardenbergia violacea, Hibiscus rosa-sinensis, Hibiscus sp., Gerber a sp.,
Jatropha sp., Lantana camara. Origanum vulgare, Pelargonium sp.. Salvia officinalis,
Solidaster sp., Solanum sp., Rosmarinus officinalis. Thymus sp., Trachelium sp., Verbena
sp. and Zelkova sp. imported from Belgium, Brazil, Canary Islands, China, Denmark,
Germany, Israel, Netherlands, Portugal, Spain and Sweden.

Comments: this species is listed in the plant health legislation of the European Union. All
interceptions and incursions of B. tabaci have been, or are being, eradicated by Fera. Any
suspected cases of B. tabaci must be reported to the PHSI.

Dialeurodes citri (Ashmead, 1885) - Citrus Whitefly

Host plants: broadly polyphagous but exhibits a strong preference for citrus.

Distribution: Central and North America, Caribbean, southern Europe, Asia, and Pacific.
Collection data: VC61, plant nurseries, Ligustrum chinensis penjing (grown indoors) from
China, various dates xi.2000 (Fera).

Comments: this species has been found on several occasions at commercial nurseries on
citrus plants imported from the Mediterranean and on penjing from China (Fera).

Discussion

Fifteen native and naturalised species of whiteflies occur in Britain (Burckhardt, 2007;
Malumphy, 2003, 2005; Martin et al., 2000) of which half have been recorded in
Watsonian Yorkshire. Two native species, Pealius quercus and Siphoninus immaculatus,
and one naturalized species, Aleurotuba jelinekii, are recorded from the region for the first
time. The whiteflies ofWatsonian Yorkshire, however, remain inadequately studied and it
is probable that other species will be found to occur in the region. It would be interesting to
confirm the presence and identity of the Aleurochiton species recorded by Harrison (1916,
1920, 1920a) and the identity of the whitefly observed by Walsh (1920) on hazel.

Siphoninus immaculatus was observed damaging a large variegated ivy plant growing

Whiteflies (Hemiptera: Aleyrodidae) ofWatsonian Yorkshire

against a wall at Fountains Abbey (Figure 7). This species has not been reported causing
plant damage in Britain before (Andrew Halstead, pers. com., 2009). The symptoms
exhibited by the plant were typical of those caused by sap-feeding bugs. The upper
surfaces of the leaves growing near the base of the plant were covered in sooty moulds
growing on honey dew eliminated by the whitefly larvae. The undersurfaces of the leaves
were covered in whitefly exuviae, pupal cases and waxy deposits. The most heavily
infested leaves were crinkled, showed signs of necrosis and were dropped prematurely.
Only a single plant was infested while other ivy plants growing nearby were completely
free from the pest.

There appears to have been an increase in the abundance and distribution of A. jelinekii
in and around York during the last decade; for example, only a single population of A.
jelinekii was recorded in York in 1997, yet by 2008-9 it was very common throughout the
centre of York, frequently occurring in very large populations smothering the undersides of
Viburnum tinus foliage (Figure 3). It was also found in Elvington, Haxby, Howsham, Sand
Hutton, Strensall and Wigginton for the first time. The puparia of A. jelinekii are
conspicuous, being black with white wax tufts (Figure 4), and it is unlikely that it was
simply over-looked previously. This is a thermophilic species and its increase in abundance
and distribution may be in response to climate change.

Bemisia afer appears to have an exceptionally wide climatic tolerance, ranging from
tropical Africa, through the Mediterranean and even over- wintering outdoors in the centre
of York. It is a broadly polyhagous species but has only been found on L. nobilis when
breeding outdoors in Britain.

The most common whiteflies in Yorkshire are Aleyrodes lonicerae on Lonicera and
Rubus\ Aleyrodes proletella on Brassica\ and A. jelinekii on Viburnum. All three of these
species frequently occur in enormous populations that can be damaging to their hosts
(covering the foliage with honey dew on which sooty moulds grow). Aleyrodes lonicerae
was found to be breeding on cultivated rose for the first time although it was not observed
causing any damage to the host plant. Trialeurodes vapor ariorum is very common on
indoor plantings and frequently found in large numbers.

Any finding of a suspected non-native whitefly in England and Wales should be
reported to the local PHSI office or to PHSI headquarters, Sand Hutton, York (Tel. 01904-
465625, Fax. 01904-465628).

Acknowledgements

I would like to express my sincere gratitude to Bill Dolling for allowing me to include his
collection data and providing me with several key references. I would also like to thank
Roger Hammon, Anastasia Korycinska, Louise MacLeod, Richard Natt, Tom Prior, Sharon
Reid and Joe Ostoja-Starzewski for collecting samples of whiteflies.

References

Alford, D.V. (2007) Pests of Fruit Crops. A Colour Handbook. Mansion Publishing,
London.

Bink-Moenen, R.M. (1989) A new species and new records of European whiteflies
(Homoptera: Aleyrodidae) from heathers ( Erica spp.). Entomol. Gaz. 40: 173-181.
Burckhardt, D. (2007) Hemiptera, Aleyrodidae. In: Burkhardt, D. (ed.) Fauna Europaea.

Version 1.1. [Available online at http://www.fauneur.org ]

Dolling, W.R. and Martin, J.H. (1985) Aleurochiton acerinus Haupt, a maple-feeding
whitefly (Horn, Aleyrodidae) new to Britain. Entomol. Mon. Mag. 121: 143-144.
Gertsson, C.A. (1987) Den svenska mjollusfaunan. Entomol. Tidskr. 108: 85-91.

Halstead, AJ. (1981) A whitefly pest of sweet bay. Plant Pathol . 30: 123.

Harrison, J.W.H. (1915) Notes and records. Aleyrodidae. Snowy winged flies. Vasculum 2:
57.

Harrison, J.W.H. (1916) Coccidae and Aleyrodidae in Northumberland, Durham and north-
east Yorkshire. Entomologist 49: 172-174.

1 2 Whiteflies ( Hemiptera : Aleyrodidae ) of Watsonian Yorkshire

Harrison, J.W.H. (1917) A new species and genus of Aleyrodidae from Durham. Vasculum
3: 60-62.

Harrison, J.W.H. (1917a) New and rare Homoptera in the Northern Counties. Entomologist
50; 169-171.

Harrison, J.W.H. (1920) Notes and records. Aleyrodidae. Vasculum 6: 59.

Harrison, J.W.H. (1920a) New and rare British Aleurodidae. Entomologist 53: 255-257.

Harrison, J.W.H. (1931) Some observations on Aleurodidae. Entomologist’s Rec. J. Var. 43:
84-86.

Hulden, L. (1986) The whiteflies (Homoptera, Aleyrodidae) and their parasites in Finland.
Notul. ent. 66: 1-40.

Malumphy, C. (2003) The status of Bemisia afer (Priesner & Hosny) in Britain
(Homoptera: Aleyrodidae). Entomol. Gaz. 54: 191-196.

Malumphy, C. (2005) The Neotropical solanum whitefly, Aleurotrachelus trachoides
(Back) (Hem., Aleyrodidae), intercepted in the UK on sweet potato leaves imported
from Gambia. Entomol. Mon. Mag. 141: 94.

Malumphy, C., Ostrauskas, H. and Pye, D. 2009. A faunistic review of whiteflies
(Hemiptera, Aleyrodidae) of Lithuania. Acta zool. litu. 19: 49-57.

Martin, J.H. (1978) Aleurochiton complanatus (Baerensprung) (Aleyrodidae) -
confirmation of occurrence in Britain. Entomol. Mon. Mag. 113: 7.

Martin, J.H. (1987) An identification guide to common whitefly pest species of the world
(Homoptera, Aleyrodidae). Trop. Pest Managmt 33: 298-322.

Martin, J.H., Misfud, D. and Rapisarda, C. (2000) The whiteflies (Hemiptera: Aleyrodidae)
of Europe and the Mediterranean. Bull. Entomol. Res. 90: 407-448.

Martin, J.H. and Mound, L.A. (2007) An annotated check list of the world’s Aleyrodidae
(Insecta: Hemiptera: Stemorrhyncha). Zootaxa 1492: 1-84.

Mound, L.A. (1962) Aleurotrachelus jelinekii (Frauen.) (Homoptera, Aleyrodidae), in
Southern England. Entomol. Mon. Mag. 97: 196-197,

Mound, L.A. (1966) A revision of the British Aleyrodidae (Hemiptera: Homoptera). Bull.
Br. Mus. (Nat. Hist.) ent. 17: 399-428.

Mound, L.A. and Halsey, S.H. (1978) Whitefly of the World. A Systematic Catalogue of the
Aleyrodidae ( Homoptera ) with Host Plant and Natural Enemy Data. British Museum
(Natural History), London / John Wiley, Chichester.

Skidmore, P. (2006) An inventory of the invertebrates of Thome and Hatfield Moors.
Thorne & Hatfield Moors Monographs 2: 1-162.

Trehan, K.N. (1940) Studies on the British Whiteflies (Homoptera - Aleyrodidae). Trans.
R. Entomol. Soc. Lond. 90: 575-616.

Walsh, G.B. (1918) Field Notes, Entomology, Hemiptera-Homoptera in East Yorkshire.
Naturalist 43: 302.

Walsh, G.B. 1920. Field Notes, New Scarborough Hemiptera. Naturalist 45: 328.

Zahradnik, J. (1956) Trois nouvelles especes des aleyrodides pour la faune tchecoslovaque.
Sb.faun. Praci Entomol. Odd. Nar. Mus. Praze 1: 43-45.

Zahradnik, J. (1989) La revision des aleurodes des pays Tcheques (Stemorrhyncha:
Aleyrodinea) III. Acta Univ. Carol. 31: 407-443.

Zahradnik, J. (1991) Taxonomisches und Faunistisches iiber europaische Mottenlaiise
(Aleyrodinea). Acta Univ. Carol. 35: 111-118.

Appendix 1.

Non-native whiteflies intercepted on imported plant produce in Yorkshire.

The six non-native whitefly species listed below do not occur anywhere in Britain and are
unlikely to survive for long on imported produce. Most of them were actually found on
foliage used for packing, rather than directly on the plant produce itself.

Aleuroclava bifurcata (Corbett, 1933)

Interception data: VC64, Bradford, Dimocarpus longan foliage used for packing longan
fruit, from Thailand, 2.x .2000 (Fera)

Whiteflies (Hemiptera: Aleyrodidae) of Watsonian Yorkshire

Aleuroclava psidii (Singh, 1931)

Interception data: VC64, Bradford, Psidium guajava foliage used for packing guava fruit,
from India, 9.ix.l999 (Fera).

Aleurodicus dispersus Russell, 1965 - Spiralling Whitefly

Interception data: VC64, Bradford, Mangifera indica foliage attached to mango fruit, from
Ghana, 6.xi.l996 (Fera).

Crenidorsum sp.

Interception data: VC64, Bradford, Dimocarpus longan foliage used for packing longan
fruit, from Thailand, 2.x .2000 (Fera).

Pealius misrae Singh, 1931

Interception data: VC64, Bradford, Psidium guajava foliage used for packing guava fruit,
from India, 9.ix.l999 (Fera).

Trialeurodes lauri (Signoret, 1882)

Interception data: VC61 , Beverley, Arbutus unedo foliage from Turkey, 14.xii.2002 (Fera).

BOOK REVIEWS

Darwin in Ilkley by Mike Dixon and Gregory Radick. Pp. 127, with numerous b/w
illustrations. The History Press, Stroud. 2009. £12.99, softback.

Darwin’s stay in Ilkley occurred at a momentous period in his life. The pressure of work
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published? Was Darwin genuinely ill or did he suffer from hypochondria? Recent research
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“psychological background”.

Debilitated yet again, Darwin could not face such pressure. He needed a bolt-hole and
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Darwin left Wells House to return home on 7 December, having stayed at Ilkley for nine
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These and many other fascinating topics are covered in this delightfully illustrated
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MRDS

Book reviews

Fumitories of Britain and Ireland by Rose J. Murphy. Pp. vi + 121, with descriptions,
line drawings, photographs and maps showing the distribution in Britain and Ireland of all
10 Fumaria species, plus four hybrids and two casuals; also contains a detailed section on
plant characters important for identification, glossary and summary of vice-counties where
Fumaria spp. have been recorded. Botanical Society of the British Isles Handbook No 12.
2009. £12.50 (paperback) plus p. & p., available from Summerfield Books, 3 Phoenix Park,
Skelton, Penrith, Cumbria CA11 9SD.

We are all aware of the critical and difficult botanical genera including Rubus, Hieracium,
Taraxacum, Euphrasia and Alchemilla , and various other publications have dealt in whole
or in part with these groups. The present volume has been accepted by the BSBI
Publications Committee as the first in a series of mini-handbooks describing small genera
that are difficult to identify.

In a small genus of only 10 British species, most botanists are readily familiar only
with common fumitory ( Fumaria officinalis ), and several of the others are indeed rare and
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synonymy where applicable. The whole topic has been thoroughly researched and there is
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tackle a fumitory which is unknown to him. It is a valuable addition to one’s bookshelf and
I recommend it to all who are interested in the genus.

GTDW

The Orchids of Ireland by Tom Curtis and Robert Thompson. Pp. 160, inch coloured
illustrations & maps. National Museums Northern Ireland, Belfast. 2009. £20.00 hardback,
plus postage & packing. [Available from: National Museums Northern Ireland, Cultra,
Holywood, Co. Down, Northern Ireland BT18 0EU, or via info@nmni.com]

This useful guide contains excellent coloured plates of each orchid in situ and close-up, as
well as distribution maps of all species to be found in Ireland. Keys to genera and to
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glossary and index.

A Dictionary of Environment and Conservation by Chris Park. Pp.vi + 522, incl. 21
line drawings. Oxford University Press. 2008. £10.99 paperback.

This valuable addition to the naturalist’s reference bookshelf is packed with up-to-date
information and well as definitions of more than 8,500 environmental terms and issues.
Highlighted key words have been treated in more detail, and appendices dealing with on-
line data sources, international treaties, the Beaufort (wind), Saffir-Simpson (hurricane),
Fujita (tornado), Torino (asteroid and comet impact hazard) and Richter and Mercalli
(earthquake) scales, geological time-scale, chemical elements, and SI units and conversions
are also provided.

WALL FERNS IN EAST HULL, 2004-2008

R. MIDDLETON
12 Meaux Road, Wawne HU7 5XD
e-mail: r.middleton@hull.ac.uk

Introduction

Casual observations over the last ten years have suggested that the fern flora of Hull has
become both more diverse and more abundant. This seems to match observations made in
London by Edgington (2000, 2003). In the Flora of East Yorkshire (Robinson 1902) no
ferns are recorded in the (admittedly smaller at that time) urban area of Hull. In the East
Riding as a whole, Phyllitis scolopendrium was described as ‘not common and apparently
vanishing’, Asplenium ruta-muraria as ‘frequent, but sparing in quantity’, A. trichomanes
as ‘frequent on old walls’, and A. adiantum-nigrum was only known on Easington Church.
The Millennium survey of Hull (Middleton 2000) revealed the presence of all of these
species growing on suburban walls in Hull (Figure 1 ) .

A subsequent study (Middleton 2005b), mapped the distribution of ferns in more detail
and established that low (c. 1 m) suburban garden walls, built of impervious bricks bonded
with lime mortar, provided an ideal environment for the growth of many calcicole fern
species. In the Hull area such walls are typical of suburban housing built during the 1920s
and 1930s. It was also proposed here that the observed increase in species diversity within
the city was a direct result of lower levels of atmospheric sulphur dioxide, a conclusion
also reached by Edgington (2003) for London ferns. Recent studies of the lichen flora of
Hull by Seaward (2004, 2007) revealed a similar increase in species diversity and
attributed this to the same cause.

Figure 1 . Phyllitis scolopendrium and Asplenium trichomanes growing on a garden wall in

suburban Hull.

Naturalist 135 (2010)

Wall Ferns in East Hull, 2004-2008

Method

The increase in fern diversity is relatively easy to monitor and since the Millennium Flora
Survey two new fern taxa have been added to the City’s flora, namely Asplenium marinum
and Cyrtomium falcatum. Although it is likely that improved air quality has made it
possible for these species to survive, long-term climate change and spore availability are
probably just as important.

In order to establish whether there is any change in the abundance of ferns a long-term
sampling study is required. Garden walls provide ideal study sites for repeat surveys as
there is always easy access and their gates are conveniently labelled with a number! Two
areas a little over 1 km apart in eastern suburbs of Hull, each comprising three adjacent,
sub-parallel, streets of inter- war housing and known “ferny” garden walls, were selected for
monitoring (Table 1 ) . The three species most commonly encountered on garden walls in this
part of the city are Asplenium trichomanes , A. ruta-muraria and Phyllitis scolopendrium.
Dryopteris spp. and Asplenium adiantum-nigrum are also to be found, but not with a high
enough frequency for meaningful comparisons to be made. In June 2004, July 2006 and
August 2008, the street-facing sides of all garden walls in the study areas were examined
and a count made of the number of walls bearing ferns of each of the target species. In 2006
and 2008 an attempt was also made to count the individual plants of each fern taxon but in
some clustered colonies it was not always possible to make a consistent count.

TABLE 1 Sampling areas

Area

Orientation

SSE - NNW

ESE - WNW

Ellesmere Avenue

Lancaster Drive

Waldegrave Avenue

Burlington Road

Shaftsbury Avenue

Skirbeck Road

Total length (m)

1050

560

Centroid

TA 130 316

TA 117 320

Results and Discussion

As can be seen from Table 2 and Figure 2a, the number of properties with “ferny” garden
walls has shown a steady increase, doubling over the 4-year period. The effect was smaller
in area 1 (55%), but very marked in area 2 (138%). The difference is due, at least in part, to
the observed loss of suitable walls for colonisation in the first area. Here the slightly wider
streets and bigger front gardens have encouraged some residents to demolish their garden
walls and create off-street parking areas, resulting in a loss of suitable habitat and offsetting
some of the potential for gain.

The increase in properties with any type of fern on the garden wall is mirrored by each
of the individual fern taxa monitored. Figures 2b-2d demonstrate that the colonisation of
walls has been proportionally similar indicating that the factor influencing the change is
affecting all species equally. If the change was solely a function of climate amelioration it
might be expected that it would influence the taxa differentially, which does not appear to
be the case. If it were warmer or moister conditions driving the change then it would be
expected that Hart’s Tongue, which is a species with a more dominantly south-western
distribution, would be favoured. The data presented here do not support this supposition.

An alternative explanation is that the expansion is a direct result of the improvement in
air quality in suburban Hull. Sulphur dioxide, and to a lesser extent oxides of nitrogen, in
the atmosphere dissolve in precipitated moisture to decrease the pH of rain water. The
reproductive biology of ferns is fundamentally different from that of vascular plants and

Wall Ferns in East Hull, 2004-2008

TABLE 2

a: Number of ferny garden walls in each street

2004

2006

2008

Ellesmere Avenue

Waldegrave Avenue

Shaftsbury Avenue

Lancaster Drive

Burlington Road

Skirbeck Road

Total

b: Walls with Asplenium trichomanes (total number of plants bracketed)

2004

2006

2008

Ellesmere Avenue

8(20)

10(80)

Waldegrave Avenue

4(13)

8(55)

Shaftsbury Avenue

0(0)

Lancaster Drive

1(1)

2(9)

Burlington Road

8(34)

11(37)

Skirbeck Road

5(47)

15(135)

Total

26(115)

46(316)

c: Walls with Asplenium ruta

muraria (total number of plants bracketed)

2004

2006

2008

Ellesmere Avenue

2(4)

1(1)

Waldegrave Avenue

3(6)

6(25)

Shaftsbury Avenue

1(1)

Lancaster Drive

13(137)

18(207)

Burlington Road

4(6)

6(23)

Skirbeck Road

6(107)

17(258)

Total

28(260)

49(515)

d: Walls with Phyllitis scolopendrium (total number of plants bracketed)

2004

2006

2008

Ellesmere Avenue

9(32)

11(78)

Waldegrave Avenue

7(13+)

9(25)

Shaftsbury Avenue

2(3)

Lancaster Drive

1(3)

5(7)

Burlington Road

2(2)

Skirbeck Road

12(64)

12(106)

Total

29(112+)

41(221)

Wall Ferns in East Hull, 2004-2008

a: any fern taxon b: Asplenium trichomanes

c: Asplenium ruta-muraria d: Phyllitis scolopendrium

2004 2006 2008 2004 2006 2008

Figure 2. Number of houses with a ferny garden wall; area 1 - hatched, area 2 - solid.

30
25
20
15
10
5
0

Figure 3. Mean hourly sulphur dioxide levels (pg.m3) in central Hull, 1994-2008.

Note that in 2002 the monitoring site was moved a short distance. The 2008 figure is for
the first eight months of the year, comparison with 2007 for the same period suggests that
the yearly average would be expected to rise by less than 0.2

involves the transmission of genetic material as gametes from one plant to another through
a film of water rather than through the air as pollen. This means that the composition of the
water film is critical to the reproductive success of ferns and can be enough to inhibit it
completely. In a recent, and temporally overlapping, study of the lichen flora of Hull
(Seaward, 2004, 2007), an increase in diversity was similarly attributed mainly to a
reduction in atmospheric sulphur dioxide.

In 1970 the total UK atmospheric release of S02 was around 6.4 Mt, slightly over half
of which was resulting from public power generation (Dore et al. 2008). By 1990 this had
fallen to a total of 3.7 Mt, with little of the decline being attributable to power stations.

Wall Ferns in East Hull, 2004-2008

After 1990 there was a marked reduction in emissions from power stations which, coupled
with a continued decline in other sources, reduced the UK total output to 1.0 Mt S02 by
2000 (Dore et al. 2008).

Although Hull’s own coal-fired power station was closed in the latter half of the 1970s
there are three of Britain’s largest power stations, Drax, Ferrybridge and Eggborough, sited
downwind of the city at the head of the Aire Valley. With this in mind, it is not to be
expected that background sulphur dioxide levels would have fallen significantly between
1970 and 1990. Road vehicle fuels were, until relatively recently, also a significant local
source of atmospheric sulphur with total UK output peaking at 64 kt in 1990, but falling
rapidly to 4 kt by 2001. Reduction in duty on low-sulphur fuels probably contributing, at
least in part, to this decline. Figure 3 illustrates the steady decline in atmospheric S02 as
measured in central Hull, the closest long-term monitoring station, over the last 15 years.
The concentration of the gas can be seen to have fallen by almost 78% over the decade
which preceded the study period and by almost 90% at the end.

Locally the situation is a little more complex as the Holliday Pigments factory, a
leading producer of ultramarine blue, was situated on the banks of the River Hull just
2.5 km WSW of area 1 and 1.5 km SW of area 2. Until flue-gas desulphurisation
equipment was installed in the late- 1990s, the factory was emitting around 6.6 kt of S02
per year. This would have contributed significantly to the atmospheric S02 in both survey
areas. Problems with the equipment meant that after a promising start the discharge consent
had to be increased from the initially stipulated 200 to 500 tonnes each year in 2004, but
still representing more than a 90% reduction (Hull City Council 2004). The site closed
completely in 2007.

Conclusions

The data show that over the monitoring period there has been a marked and consistent
increase in both the number of walls bearing ferns and the number of individual fern plants
within the study areas. This increase amounts to a little more than a doubling of the number
of walls colonised and is consistent for each of the three species monitored. Although it
was only measured over the last two years, the total number of plants of each taxon also
increased by a factor of 2 or more.

Measurements taken in central Hull over the same period show a consistent fall in the
background concentration of atmospheric S02. Local factors have also reduced the amount
of sulphur discharged into the air around the study areas and it would be tempting to
suggest a simple inverse relationship between the number of individual ferns counted and
the level of S02 What seems more likely is that regeneration is effectively inhibited when
the atmospheric sulphur is above a certain level. The increase observed in both the number
of sites and individual plants is evidence of a natural colonisation process which only
started when concentrations fell below a critical level, possibly over a decade ago. The
critical level was probably dependent on several factors including the amount of rain at
crucial periods in the fern’s reproductive cycle.

References

Crackles, F. E. (1990) Flora of the East Riding of Yorkshire. University of Hull, Hull.

Dore, C.J. et al. (2007) UK Emissions of Air Pollutants 1970 to 2005. Download available
at: http://www.airquality.co.uk/archive/reports/cat07/080 1 140937_2005_Report_FINAL.pdf
Edgington, J. (2000) Aspleniaceae and Polypodiaceae in London. The London Naturalist
79:51-54.

Edgington, J. (2003) Ferns of the metropolis - a status report. The London Naturalist 82:
59-73.

Hull City Council (2004) Air quality annual report. Download available at:
http://www.hullcc.gov.uk/pls/portal/docs/PAGE/HOME/ENVIRONMENT/
POLLUTION/ AIR% 20QU ALIT Y/AIR%20M AN AGEMENT/AIR% 20
MONITORING/MAY2004DETAILED ASSESSMENT.PDF

20 Wall Ferns in East Hull, 2004-2008

Middleton, R. (2000) The plants of Hull: a millennium atlas. Available online at:
http://www.hull.ac.uk/hullflora/ viewed Jan. 2009.

Middleton, R. (2005a) Suburban ferns in Kingston-upon-Hull - exploitation of a deco-
mural environment. BSBI News 98: 28-30.

Middleton, R. (2005b) Ferns in Kingston-upon-Hull. Unpublished report for the Hull
Biodiversity Partnership. 25pp.

Robinson, J. F. (2003) The Flora of the East Riding of Yorkshire. Brown, Hull.

Seaward, M. R. D. (2004) The lichen flora of Hull, with particular reference to zonal
distribution and environmental monitoring. Naturalist 129: 61-66.

Seaward, M. R. D. (2007) The lichen flora of Hull: biodiversity update, 2002-2006.
Naturalist 132: 41-49.

BOTANICAL REPORT FOR 2009
FLOWERING PLANTS AND FERNS

Compiled by
P.P. ABBOTT

It is amazing and gratifying that every year rare and uncommon plants are still being found
in new sites, some of them even new to Yorkshire although these are usually non-native
plants which have escaped from gardens. However, the star find this year must be Cotula
alpina, an Australian species native to New South Wales, Victoria and Tasmania, which is
believed to be new to Britain and to Europe. It was found and identified by Linda
Robinson, first on Kirkby Malzeard Moor and then in several other sites in VCs. 64 and 65
on the moorland between Nidderdale and Wensleydale. The identification was confirmed
by Eric Clement. In 1995 it had been found on Dallow Moor and misidentified as
C. squalida since C. alpina is not mentioned in any of the British Floras. Similarly, it was
found in September this year, by members of the Ryedale Natural History Society, on
Rudland Rigg in VC 62 and initially identified as C. squalida. The plants at this site have
now also been redetermined as C. alpina.

Other notable finds include Oenanthe pimpinelloides in fields near Priory Meadows in
Hull, its only site in northern England, and Salvia verbenaca, found this year in Staveley
churchyard, which had not been recorded in VC 64 since 1876. It was pleasing to learn that
the iconic Cypripedium calceolus has produced flowers in all its reintroduction sites. Other
interesting records are listed below.

Nomenclature is according to Vice-county Census Catalogue of Vascular Plants of
Great Britain (2003) edited by C.A.Stace et al. * denotes a new vice-county record.

SOUTH-EAST YORKSHIRE (VC 61)

Anacamptis pyramidalis Hull, Priory Meadows, TA0531 , D.R.Grant

Carex divisa North Ferriby, field edge, SE988252, P.Cook

Carex viridula ssp. viridula Allerthorpe Common, SE7647, D.R.Grant

Filipendula vulgaris Cleaving Combe, SE8646, D.R.Grant

Gentiana pneumonanthe Skipwith Common, SE653372, D.R.Grant

Hieracium umbellatum Allerthorpe Common, SE7547, D.R.Grant

Hordeum secalinum North Ferriby, field edge, SE998252, P. Cook

Oenanthe aquatica Hull, Priory Meadows, TA0531 , R.Middleton

Oenanthe fistulosa Hull, Priory Meadows, TA0531 , R.Middleton

Oenanthe pimpinelloides Hull, fields near Priory Meadows, TA0531, R.Middleton

Ophioglossum vulgatum Hull, Priory Meadows, TA05 31, R.Middleton

Silaum silaus Hull, Priory Meadows, TA0531 , R.Middleton

Thalictrum flavum Hull, Priory Meadows, TA053 1 , R.Middleton

Naturalist 135 (2010)

Botanical Report for 2009 - Flowering Plants and Ferns

NORTH-EAST YORKSHIRE (VC 62)

Actaea spicata Gilling, Scar Wood, SE619768, G.Smith
Allium scorodoprasum Broughton, north side of track, SE767720, G.Smith
Antennaria dioica Sand Dale, SE858848, G.Smith
Aphanes australis Lodge Hag, SE670741 , W.Thompson
Atropa belladonna Hutton Common, on hillside, SE7087, G.Smith
Berberis vulgaris Gilling, SE607771, G.Smith

Campanula glomerata Broughton, Flowery Lane road verge, SE766716, G.Smith
Carex divulsa ssp. leersii Hovingham, Socarrs Lane, SE675761, W.Thompson
Cirsium eriophorum Hutton Common, on hillside, SE7087, G.Smith
Cirsium eriophorum Wath, old quarry, good colony, SE676749, W.Thompson
Clinopodium acinos Gundale, base of old quarry, SE800881, G.Smith
Colchicum autumnale Castle Howard Arboretum in rough grassland, SE703701 ,
W.Thompson

Convallaria majalis Wath Wood, a large patch, SE673741 , W.Thompson
Cuscuta epithymum Gundale, on Rockrose, SE800880, G.Smith
Dactylorhiza traunsteineri Sand Dale, SE858848, G.Smith, det. R.Bateman
Epipactis palustris Sievedale Fen, at base of slope, SE853878, G.Smith
Equisetum hyemale Goathland, footpath behind hotel, NZ826010, G.Smith
Eriophorum latifolium Sand Dale, SE858848, G.Smith
Eilipendula vulgaris Yatts, SE807881 , G.Smith
Frankenia laevis Howkeld, roadside, SE685856, G.Smith
Gagea lutea Appleton, near top of bank, SE742880, G.Smith
Gagea lutea Kirkdale, SE677857, G.Smith, W. A .Thompson
Gymnadenia conopsea ssp. densiflora Sievedale Fen, SE853878, G.Smith
Helleborus viridis Ashberry, many plants, SE5784, G.Smith
Hordelymus europaeus Kitscrew Wood, SE67 1748, W.Thompson
Lathraea squamaria Gilling, on Hazel above church, SE616768, G.Smith
Lathraea squamaria Robson’s Spring, by bridleway east of Low Parks farm, SE6381 ,
G.Smith

Lister a cordata Damholme, 2 clusters, NZ838023, G.Smith
Lithospermum officinale Wath SINC, woodland edge, SE670746, W.Thompson
Monotropa hypopitys Sutherbruff, under Pine tree, SE870875, G.Smith
Narcissus pseudonarcissus Appleton, woods to east, SE742880, G.Smith
Neottia nidus-avis Yedmandale, old quarry; many flower spikes, SE978860, M.Walshaw
Ophrys insectifera Yatts, many plants, SE806879, G.Smith
Orchis morio Hole of Horcum, SE849936, G.Smith
Orchis ustulata Yatts, on bank, SE806879, G.Smith
Orobanche elatior Broughton, Flowery Lane, SE767721 , G.Smith
Orobanche elatior Swinton Lane, SE760720, W.Thompson
Paris quadrifolia Gilling, SE6 19769, G.Smith
Phegopteris connectilis Brandsby Dale, 4 colonies beside stream, SE595735, W.Thompson
Picris hieracioides Hovingham quarry, on quarry faces and spoil, SE669752, W.Thompson
Platanthera bifolia Yatts, SE806879, G.Smith
Platanthera chlorantha Hutton Common, SE7087, G.Smith

Populus nigra Stonegrave, regeneration from base of tree blown down in gale. Last known
Black Poplar in the area, SE756775, W.Thompson
Potentilla anglica Bulmer Hagg, SE7167, G.Smith
Ranunculus lingua Whitwell, SE7 15668, G.Smith

Rhamnus cathartica Wath SINC, woodland edge, SE670745, W.Thompson

Rorippa microphylla Hovingham, in and beside beck, SE666758, W.Thompson

Rubus amplificatus Carlton, NZ506041 , V. Jones

Rubus cinerosus near Great Ay ton, NZ574 108, A. Newton

Rubus conjugens Whitby, Upgang ravine, NZ850118, A.Newton

Botanical Report for 2009 - Flowering Plants and Ferns

Rubus incurvatiformis near Kildale, NZ6 13098, VJones

Rubus infestus Yearsley Moor, SE579750, A. Newton

Rubus pallidus Goathland, NZ824001 , D.R.Grant

Silene noctiflora Whitwell, set aside field, SE7266, G.Smith

Spergularia rubra Lastingham, SE729908, M.Walshaw

Stellaria neglecta Horse Coppice, grass verge, SE654751, W.Thompson

Stellaria neglecta Brandsby, laneside, SE595723, W.Thompson

Trientalis europaea Hovingham High Wood, SE64875 5, W.Thompson

Verbascum nigrum Hartoft, SE749931, G.Smith

Vicia sylvatica Cawthom, Peat Rigg Bank, SE770895, G.Smith

SOUTH-WEST YORKSHIRE (VC 63)

Asplenium adiantum-nigrum Horbury Junction, brick wall, SE302177, D.R.Grant

Clematis vitalba Shafton, Rabbit Ings, SE378114, J.Greaves

Dactylorhiza praetermissa Bullcliffe Wood, old mine site, SE285149, D.Proctor

Dactylorhiza maculata Wessenden Head, SE070070, M.J. Lucas

Frangula alnus Howell Wood Country Park, SE435096, J. Lambert

Hieracium umbellatum Bolton-on-Deame, old mine site, SE432028, T.Schofield

Narthecium ossifragum Wessenden Head, SE070070, M.J .Lucas

Picris echioides Kilnhurst, railway wall-top, SK459971 , T.Schofield

Rubus echinatus Bullcliffe Wood, old mine site, SE285149, D.R.Grant

Rubus rufescens Howell Wood Country Park, SE4309, YNU, det. D.R.Grant

MID- WEST YORKSHIRE (VC 64)

Allium oleraceum Leeds, Gipton Wood, SE325367, P.P. Abbott
Allium scorodoprasum Newton Kyme, roadside, SE451447, P.P.Abbott
Allium vineale Leathley, SE229482, N. Vernon
Andromeda polifolia Red Syke Head, SD679605, PC .G .Green
Aquilegia vulgaris Oxenber Wood, SD7868, Bradford Botany Group
Asperula cynanchica Ledsham Banks, SE4630, S.Joul
Asplenium adiantum-nigrum Ilkley, SE1046, Wharfedale Naturalists’ Society
Campanula glomerata Famham, road verge, SE354669, P.P.Abbott
Campanula glomerata Roach Lime Hills, many plants, SE419315, P.P.Abbott
Campanula trachelium Follifoot, railway cutting, SE340576, 1.Wallace
Carex disticha Ribblehead Quarry, SD767787, J.Newbould, T.Whitaker
Carex elongata Bishop Wood, second.record in V.C. 64, SE551326, M.Hammond
Ceterach ojficinarum Foxup, by Foxup Beck, SD860766, B. Burrow
* Dactylorhiza xformosa Ribblehead, SE776784, N.Barrett, M.Wilcox
Dactylorhiza x insignis Otley Wetlands, SE1844, Bradford Botany Group
Dipsacus pilosus Bolton Abbey Woods, SE075555, B. Brown
Dipsacus pilosus Fountains Abbey, SE280688, 1.Wallace

Dryopteris submontana Giggles wick Scar, SD809654, R. Wilding, conf. M.Canaway
*Dryopteris x ambrosiae Ingleborough, SD743747, B. Brown, conf. K.Trewren
Equisetum x litorale Ribblehead Quarry, SD766787, M.Wilcox, conf. B. Brown
Filago vulgaris near Hellifield station, SD8557, E.Shorrock
Filipendula vulgaris Ledsham Banks, SE462302, Leeds Naturalists’ Club
Groenlandia densa Harden Bridge, in Austwick Beck, SD762678, C.R. Abbott, P. Abbott
Gymnocarpium robertianum Feizor, in scree below scar, SD789681, Bradford Botany
Group

Hottonia palustris Spofforth, SE364514, 1.Wallace
Huperzia selago Antley Gill, SD670596, P.C.G.Green
Hypericum humifusum Fewston, SE 160564, M. Atkinson
Lathraea squamaria Hubberholme, SD918783, N.Moore

Botanical Report for 2009 - Flowering Plants and Ferns

Neottia nidus-avis Fountains Abbey, 29 spikes, SE281688, 1.Wallace
Onobrychis viciifolia Follifoot, SE340576, 1. Wallace

Orobanche minor Newton Kyme, road verge; on Vicia sepium, SE455447, A. Hanson,
R.Masheder det. P.P. Abbott

Polystichum setiferum Dunsop Fell, SD6854, EE.Greenwood
Polystichum setiferum Otley, riverbank, SE206461 , B. Brown
Pulicaria dysenterica Otley Chevin, SE20 1445, B. Brown
Ranunculus penicillatus Clitheroe, SD7242, EE.Greenwood
Rhamnus cathartic a Kettle well, SD7242, T. Whitaker
Ribes spicatum Barden, SE054584, B. Brown, conf. M. Wilcox
Rubus polyanthemus Adel, SE281401 , D.R. Grant
Salix x ambigua Ribblehead, SD776784, M.Wilcox
Salvia verbenaca Staveley churchyard, SE362626, B.Lobo, conf. K.Walker
Saxifraga x polita Slaidbum, SE7052, EE.Greenwood
Stellaria neglecta Edisford, SD725414, M.Wilcox
Thlaspi caerulescens Yambury, old lead mine, SE017657, Wharfedale Naturalists’ Society
Tilia platyphyllos Ledsham, Claypit Lane, small tree in hedge, SE450293, Leeds
Naturalists’ Club

Veronica polita Edisford, SD724414, M.Wilcox

NORTH-WEST YORKSHIRE (V.C. 65)

Alopecurus borealis Great Shunner Fell, SD842970, R.Comer, L.Robinson
Allium scorodoprasum West Burton, SE0186, YNU Botany Section
*Asperula cynanchica Widdale Bridge, hay meadow, SD825876, F. Graham
*Dactylorhiza x grandis Marfield Wetlands, SE2082, L.Robinson, A. Gendle
*Epilobium xfacchinii Shunner Fell, flush, SD844971, J. Somerville, M.Wilcox
Equisetum x litorale Ballowfield NatureReserve, SD9889, BSBI
*Equisetum x rothmaleri Ballowfield Nature Reserve, SD9889, BSBI
Euphrasia arctica ssp. borealis West Burton, SE0086, YNU Botany Section
Galium x pomeranicum Kilmond Scar, NZ0212, J.Clarke, L.Robinson
*FIieracium leyi Widdale, gill, SD799886, B. Burrow, conf. D.McCosh
Vaccinium uliginosum Great Shunner Fell, peat hags on summit plateau, SD848972,
R.Dimon, M.Owen

Vaccinium uliginosum Great Shunner Fell, acid mire, SD843969, S. Hewitt

ALIEN PLANTS

Aconitum lycoctonum Amotherby Church grounds, SE749733, W.Thompson
Allium subhirsutum Chellow Dene, SE131339, B.A.Tregale, M.Wilcox
Alopecurus myosuroides Dentdale, head of dale, SD764843, J.Clarke
Alyssum saxatile Thackley, on garden wall. Old Windhill Road, SE168384, B.K.Byme,
BATregale, M.Wilcox

Amaranthus retroflexus Middlesborough, waste ground, Douglas St, NZ503193 V.Jones,
W.Thompson

*Anemanthela lessoniana York, waste ground, SE599519, V.Jones, W.Thompson
*Astilbe x arendsii Stonehouse Bridge, riverside, SD7684, M.Canaway, L.Robinson
Bassia scoparia Middlesborough, NZ503 193, V.Jones, W.Thompson
* Bergenia x schmidtii Eldwick, woodland, SE121386, B.A.Tregale, M.Wilcox
*Bergenia x schmidtii Bradford Horton Bank, SE126310, BATregale, MWilcox
Camelina sativa Thornaby, Tees-side, NZ47 1184, V.Jones, W.Thompson
Chiastophyllum oppositifolium Dentdale, by road bridge, SD7686, B. Burrow
Chrysanthemum segetum Whitwell, 1 plant, SE7 1 966 1, G. Smith
Clematis tangutica Scarborough Crossing, TA030459, J.Killingbeck
Cordyline australis Bradford, Queens Avenue, SE 164349, B.A.Tregale
*Cotula alpina Kirkby Malzeard Moor, SE176760, L.Robinson

Botanical Report for 2009 - Flowering Plants and Ferns

*Cotula alpina Potts Moor, SE1 19754, L.Robinson, conf. E.Clement
*Cotula alpina Rudland Rigg, SE649956, Ryedale Natural History Society
* Cyclamen coum Hirst Mill, on dumped soil, SE131384, B.A.Tregale, M.Wilcox
Cyclamen coum Thackley, field, Old Windhill Road, SE 163 3 81, B.A.Tregale, M.Wilcox
Cymbalaria pallida var. beguinotii Thackley, outside of garden wall, Old Windhill Road,
SE163381, B.A.Tregale, M.Wilcox

Cyrtomium falcatum Hull, Wilberforce House, garden wall of museum, TA 10328 8
R. Middleton

Dipsacus laciniatus Otley, Crow Lane, SE205455, J. Hartley, N.Vemon
Doronicum pardalianches Harden Bridge, SD762678, C.R. Abbott, P.P. Abbott
Duchesnia indica Yarm, West Street, NZ417129, VJones, W.Thompson
*Escallonia x langleyensis Bradford, bank by Boar’s Well, SE164357, B.A.Tregale
Geranium macrorrhizum west of Cowgill, in lay-by, SD742864, J.Somerville et al.

Geum macrophyllum Shipley Glen, SE128389, Bradford Botany Group
*Helianthus x laetiflorus Champion, SD7452, E.F.Greenwood
*Iris x robusta Stonegrave, by pond, SE654777, W.Thompson
*Jasminum officinale Yarm, waste ground, Bridge Street, NZ418 131, VJones,

W.Thompson

J uncus tenuis Skip with Common, old runway, SE647370, R. Middleton
*Knautia macedonica Bradford, waste ground, Fairweather Green Mills, SE133334,
B.A.Tregale, M Wilcox, det. E.J. Clement
Kniphofia uvaria Dalton-on-Tees, woodland edge, NZ2808, L.Robinson
Lemna minuta South Wood, in pond, SE666742, W.Thompson
Leucojum vernum Shackleton Lane End, SE653721, W.Thompson
Lilium martagon Austwick, wide verge by track, SD773768, Bradford Botany Group
Luzula luzuloides Cowgill, edge of rail track, SD764875, J. Clarke
Lychnis coronaria Masham, edge of lagoon, SE2 19821, L.Robinson
Malva alcea West Marton, roadside, SD889503, J.Clarke
Mimulus moschatus Northdale, marsh, Castleton Rd, SE7 18978, M.Walshaw
Nectaroscordum siculum Famhill, below trees, SE009460, M.Canaway
*Osteospermum jucundum Saltwick, near caravans, NZ915107, VJones, A.Ritson
*Oxalis tetraphylla Nunthorpe, lane near Grange Farm, NZ542140, VJones
Petrorhagia prolifera Redcar, South Gare, amongst slag, NZ557273, VJones
*Phlox paniculata Dent, in hedge, SD705866, J.Clarke
Phytolacca acinosa Skelton-on-Ure, by windmill, SE375695, 1.Wallace
Pratia pedunculata Pickering, lawn in Lintondale, SE792 847, VJones, W.Thompson
*Primula denticulata west of Cowgill, in lay-by, SD742865, J.Somerville et al.

*Primula x pruhonicensis Thackley, field by Old Windhill Road, SE 1 6338 1 , B.A.Tregale,
M.Wilcox

Rhus typhina Chellow Dene, SE 129340, B.A.Tregale, M.Wilcox
Saxifraga x geum Dentdale, along river, SD7686, B. Burrow
Sedum hispanicum Greenhow, stone wall, SE1 13641 , BSBI det. G.M.Kay
Senecio inaequidens Hull, Priory Road, TA049320, J.Dews
Solanum physalifolium York, Water End, Clifton, grass verge, SE590528, VJones,
W.Thompson

Sorbus commixta Shipley, riverside in scrub, SE131384, B.A.Tregale det. H.. McAllister
*Trollius x cultorum Thackley, disused railway, SE168384, B.A.Tregale
*Tulipa gesneriana Leybum Shawl, SE094903, B. Armstrong, L.Robinson
*Viola x wittrockiana Masham, edge of lagoon, SE2 19821, L.Robinson

THE OSTRACOD SCOTTIAPSEUDOBROWNIANA KEMPF, AN
ADDITION TO THE FRESHWATER FAUNA OF YORKSHIRE,
AND A NOTE ON THE EVOLUTION OF TERRESTRIAL HABITS
IN OSTRACODS AND OTHER CRUSTACEANS

ANDY GODFREY,

90 Bence Lane, Dart on, Barnsley, South Yorkshire S75 5DA
GEOFFREY FRYER

Elleray Cottage, Windermere, Cumbria LA23 1AW
and

DAVID J. HORNE

Dept, of Geography, Queen Mary University of London, Mile End Road, London El 4NS
and Dept, of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD
email: d.j.home@qmul.ac.uk

Introduction

In 1850 T. Rupert Jones described from Pleistocene deposits at Clacton, England, an
ostracod which he named Cypris browniana, of which only the carapace valves (the so-
called shell) were preserved (Jones, 1850). The genus Cypris was at that time used to
embrace forms that were later assigned to several genera. In 1889 Brady and Norman
reported that they had received from Thomas Scott material of an ostracod that he had
found living “in pools near Loch Fadd” on the Island of Bute, Scotland, which they
regarded as being conspecific with Jones’ fossil C. browniana. Although the fossils were of
Pleistocene age, and therefore not very old, geologically speaking, this was nevertheless an
interesting situation. From their living Scottish material they described and illustrated some
of the appendages (Brady & Norman, 1889). They also assigned the animal, adult females
of which achieve a length of up to c. 0.83 mm, to a new genus, Scottia, named for the
finder of the living specimens.

Its describers were sufficiently intrigued by the discovery of living Scottia for Norman
to visit the site where it was collected, which was later described by Brady and Norman
(1896). Although the animal was originally said to be from “pools near Loch Fadd”, they
described its habitat more precisely as a spring which “rises on the bank close to the loch,
into which the water finds its way among the grass; the water is nowhere trickling more
than two or three inches deep among the herbage”. Here Scottia was living with a rich
assemblage of other ostracods, Psychrodromus robertsoni, Herpetocypris reptans , Eucypris
pigra, Candona Candida , Candonopsis kingsleii and Plesiocypridopsis newtoni, whose
generic assignments given here are the modem ones, not necessarily those used by Brady
and Norman. Of these, all save P. newtoni are non-swimming, creeping species.

Examples of living Scottia, assigned to S. browniana, were subsequently recorded from
various sites in Europe under this name. However, Kempf (1971) showed that living
representatives of what had long been regarded as conspecific with the Pleistocene fossils
referred to as S. browniana are in fact distinct from it, and he therefore renamed the living
species S. pseudobrowniana. The situation is, however, more complex than this simple
statement implies. In fact three distinct species are involved: S. browniana,

S. pseudobrowniana and S. tumida. The last-mentioned had been described (as Cypris
tumida) as a Pleistocene fossil from Essex by Jones (1850) and subsequently, also as a
Pleistocene fossil, as Cyclocypris huckei by Triebel (1941), but Kempf (1971) recognised
not only that these were conspecific but also that the species they represent belongs in the
genus Scottia. Fossil S. browniana is sometimes a major component of ostracod
assemblages in certain palaeoenvironmental settings in British Pleistocene deposits
(Robinson, 1980). In Europe, both S. browniana and S. tumida are extinct, Early to Middle
Pleistocene species, the youngest British occurrence of S. browniana being in the Hoxnian
interglacial (Marine Isotope Stage 11), while S. tumida ranges higher, as far as the Purfleet

Naturalist 135 (2010)

The Ostracod Scottia pseudobrownian Kempf

interglacial (MIS 9) (Whittaker & Horne, 2009). The extant S. pseudobrowniana, which is
now also known as a fossil, is apparently confined as such to the Late Pleistocene and
Holocene (De Deckker, 1979; Robinson, 1980). Thus, although Brady and Norman (1889)
were in fact in error when they assigned the living Scottia from Bute to S. browniana, it
does indeed belong to a species that also occurs in the fossil state !

As no appendages of S. browniana and S. tumida are known, discrimination of the
three species has to be based entirely on carapace shape, but Kempf (1971) demonstrated
convincingly that this is possible. Compared to S. pseudobrowniana , both S. browniana
and S. tumida have relatively shorter, approximately semi-circular carapaces (in lateral
view); that of S. browniana has a short, straight dorsal marginal section, while that of
S. tumida is relatively higher with a more arched dorsal margin. It is interesting to note
that, in common with several other British Pleistocene species such as Ilyocypris salebrosa
(see Bates et al., 2002), S. tumida may still be living in North America although it is
evidently extinct in Europe; however, records from the USA of both S. tumida and
S. pseudobrowniana (e.g. in the NANODe database: Forester et al., 2005) have yet to be
verified by careful comparison of specimens with European material; for example,
S. pseudobrowniana has been reported in the USA from Tennessee (Cole, 1966) and
Nevada (Kiilkoyliioglu & Vinyard, 2000), but Smith et al. (2002) point out differences in
the appendages of the latter record which suggest that it may be a different species.

It should be noted that the illustrations of the whole animal (i.e. the carapace and
appendages) given by Brady and Norman (1889) and reproduced by Klie (1938) were
based on material of the living S. pseudobrowniana from near Loch Fadd and not on fossil
S. browniana. Meisch (2000) provided a description and excellent illustrations, including
the appendages, of S . pseudobrowniana.

The known distribution of all three species is shown on a map provided by Kempf
(1971) and their ranges have only been extended slightly by more recent finds. Although
Kempf differentiated between sites at which living and fossil representatives of
S. pseudobrowniana were found, curiously he indicated the Bute site, from which the first
living examples were described, as that of a fossil find. Its distribution (Fig. 1) is
predominantly north-central and eastern European but it is also known from northern Italy

Figure 1 . European distribution of extant Scottia pseudobrowniana, from the NODE
database (Home et al., 1998).

The Ostracod Scottia pseudobrownian Kempf

and has been reported in Turkey (Kiilkoyliioglu, 2003, 2005). As noted above, it may also
live in North America. In Britain it is known from relatively few sites. Apart from the
original site on Bute, it has been found near Birmingham (a few males only) by Lowndes
(1930) and at the margin of Skelsmergh Tarn in Cumbria by Danielopol et al. (1996).
Henderson (1990) indicated, on a map, a site apparently in Essex, which may in fact be the
fossil type locality of S. browniana. It is known from one site in Ireland (Douglas &
McCall. 1992).

First record of Scottia pseudobrowniana in Yorkshire

Here we report the discovery of S. pseudobrowniana in Yorkshire by Andy Godfrey who
collected it in moss from tufaceous springs and seepages at Seive Dale Fen, Low Dalby,
North Yorkshire (SE854881) on 1 July 2005. The seepages and springs are on an open and
gently sloping flushed hillside and the streams issuing from these flow eastwards where
they join Dalby Beck at the foot of the slope. Vegetation around the open seepages and
springs included Schoenus nigricans , Carex spp., Filipendula ulmaria , Dactylorhiza sp.,
Cardamine pratensis, Lychnis flos-cuculi, Succisa pratensis, Pedicularis palustris and
Hydrocotyle vulgaris, with fringing Salix caprea and Alnus glutinosa woodland.
Stone worts were also frequent at the springheads and in the channels. The streams issuing
from the springs and seepages were generally no more than 10cm wide and usually less
than 5cm deep. Their water was clear brown with flow only perceptible in the narrower
parts of the channels. The following single measurements were made in the springs: pH
7.81, water temperature 11.6°C, conductivity 612pS cm1 and total dissolved solids
308ppm. The substrate was a combination of mud and organic detritus and included
obvious tufa fragments. The area around the seepages and springs was poached but no
cattle were present on the day of the visit. Other invertebrates associated with the springs
and seepages included the aquatic larvae of the Red Data Book 1 and UK Biodiversity
Action Plan soldier-fly Odontomyia hydroleon and the larvae of other scarce or local
soldier-flies ( Oxrycera morrisii, O. pygmaea and Stratiomys singularior) , the Nationally
Scarce water-penny beetle Eubria palustris and the local Sphagnum-bug Hebrus ruficeps.

Habitat requirements

Kempf (1971) surveyed the published information on the ecology of S. pseudobrowniana
that was available at the time. A preference for springs or spring-fed situations is evident;
these are of various kinds, often swampy and sometimes grassy or mossy and, not
surprisingly, predominantly cool, though there are records of it at 22°C in a spring-fed area
of horsetails ( Equisetum ) and sedges and at 17°C in a stand of Char a fed by a highly
calcareous spring. Springs with lime-encrusted moss, or in which ferruginous deposits are
present, are among those frequented.

A recent find by Kiilkoyliioglu (2003, 2005) in Turkey confirms earlier suggestions
that it is a crenobiont, preferring cool springs or waters connected to such springs. Its
associates in spring-fed areas include several ecologically-tolerant species of ostracods but
also Cryptocandona vavrai and Potamocypris zschokkei as well as (most frequently)
Psychrodromus olivaceus and Eucypris pigra, all of which favour such sites and seepages.
It has also been found in springs and swamps in woodland among wet leaf-litter (Meisch
2000) where, as it does in moss, it leads a semi-terrestrial (as opposed to fully aquatic)
existence. In a Romanian lake it has been found inhabiting floating “swamp islands”
(Danielopol & Vespremeanu, 1964) comprising masses of Phragmites , Typha, Carex and
plant detritus, quite similar to the “floating fen” habitat in which it was found at the
margins of Skelsmergh Tarn in Cumbria, UK (Danielopol et al., 1996). In such habitats it is
commonly associated with the ostracods Candonopsis kingsleii and Metacypris cordata.

The ecology of the Seive Dale Fen animals is in accord with previous knowledge of the
species. The Seive Dale springs are tufaceous, cool and characterised by vegetation
including mosses and charophytes, which closely matches the observations recorded by
Kempf (1971), Meisch (2000), Kulkoyliioglu (2003) and others. They are also

The Ostracod Scottia pseudobrownian Kempf

characterised by the local black bog rush Schoenus nigricans and it is possible that
searches in other springs where this distinctive calcicole plant occurs may reveal further
occurrences of this scarce and probably under-recorded ostracod. It is interesting that as a
result of extensive work on Pleistocene fossil faunas, the genus Scottia has come to be
regarded as an indicator of small, shallow, permanent water bodies, with a marked
tendency to weed growth and the ultimate development of a fen (e.g. Robinson, 1980),
which is not at variance with the preferences shown by its living representative in Europe.

FUNCTIONAL MORPHOLOGY

Scottia pseudobrowniana is a crawling species, as is apparent from the shortness of the so-
called natatory setae on its antennae and from the robust nature of the furcal (or uropodal)
rami that are armed distally with a pair of strong claws (among the stoutest of any
podocopid species) which are used for levering the animal forwards. The antennae, which
have stout distal claws, serve mainly to pull the animal along and cannot be used for
swimming (in many freshwater ostracods the antenna has a dual role in both crawling and
swimming). Probably related to its crawling habits is the presence, at the distal extremity of
the sixth limb (the so-called walking leg), of not only the robust, curved, terminal claw or
spine usual in cypridids, but also a second spine, more slender but of similar length. Both
spines are denticulate. In cypridids other than Scottia , the second terminal spine is usually
represented only by a small, slender seta. Exactly how the twin terminal spines are used is
not clear, but they may aid in climbing among the vegetation through which the animal
crawls. The carapace valves (Fig. 2) are thick and robust, which may be an advantage when
the animal leaves the cushioned aquatic environment, as is probably the ability to close
them tightly when necessary.

Figure 2. Scottia pseudobrowniana from Seive Dale Fen, North Yorkshire.

Figured specimens deposited in the Zoology (Crustacea) collection at the Natural History
Museum, London; catalogue numbers given in brackets. A: female carapace, right lateral
view (2010.245); B: male left valve, internal view (2010.246); C: female carapace, ventral
view (2010.247); D: detail of framed area of (C) showing rows of sensilla near outer margin
of right valve. Arrows indicate anterior direction. Scale bars A-C: 100 pm; D: 50 pm.

The Ostracod Scottia pseudobrownian Kempf

Scottia and the evolution of terrestrial habits in ostracods
For more than 70 years S. pseudobrowniana remained the only known extant species of
Scottia. Subsequently three living congeners have been discovered which throw interesting
light of the history of the genus from geographical and ecological standpoints. The
description of S. audax from New Zealand by Chapman (1961) (who originally placed it in
the genus Mesocypris ) gave Scottia an unexpectedly wide and disjunct distribution (De
Deckker, 1980). A second species, S. insularis (Chapman, 1963) is known from Australasia
but is inadequately described. More recently S. birigida has been described from Japan by
Smith et al. (2002). Such a wide distribution hints at an ancient origin for the genus
although pre-Pleistocene fossil records appear to be sparse and uncertain; Swain (1999)
lists several species from the Tertiary of the USA that are assigned to Scottia with varying
degrees of confidence, the oldest being of “late Cretaceous or early Tertiary” age, but these
records are in need of verification. Ecologically, the extant species share with
S. pseudobrowniana a propensity for a semi-terrestrial or even entirely terrestrial way of
life. S. birigida, for example, was found in wet leaf litter, and exploration of its habitat
revealed it to be living among wet fallen leaves and on the soil surface at all times of the
year, in a climate where the temperature reaches 30-35°C in summer and may fall as low as
-5°C in winter, when the area is often covered with snow. S. audax was described as a truly
terrestrial species.

This preference of species of Scottia for habitats that lead to a semi-terrestrial or even
terrestrial way of life is reflected in the affinities of the genus which lie with ostracods,
most of which have been described relatively recently, that have similar habits. Scottia is
assigned to the Subfamily Scottiinae (of the Family Cyprididae, Superfamily Cypridoidea),
the only other members of which are Mesocypris and the recently described
Austromesocypris , neither of which has British representatives, and all of which show a
marked preference for terrestrial or semi-terrestrial habitats. The vast majority of members
of other subfamilies of the Cyprididae, which are common and diverse in non-marine
waters, are fully aquatic. That ostracods include truly terrestrial taxa was not known until
1953, when Mesocypris terrestris was described from forest humus in the Knysna Forest,
South Africa. Of this species Harding (1953) gives a well-illustrated account. It is
interesting that there is nothing in its morphology that immediately indicates that its habits
are terrestrial and not aquatic. An earlier known species of this genus, M. pubescens, was
already known as a frequenter of wet moss in the splash zone of a waterfall and similar
situations in Kenya (Klie, 1939), but M. terrestris has taken the tendency to emancipate
itself from water even further. Since its discovery, two more species of Mesocypris have
been found to be terrestrial, as have three species of Austromesocypris from Australia
(Martens et al., 2004). Mesocypris and Austromesocypris are believed to be more closely
related to each other than to Scottia and have been recognised as constituting a tribe, the
Mesocypridini, while Scottia has been allocated to a second tribe, the Scottiini.

The discovery that some ostracods have adopted terrestrial habits led to searches and
discoveries elsewhere (e.g. Danielopol & Betsch, 1980; De Deckker, 1983); indeed we now
know that the terrestrial mode of life has arisen more than once among ostracods.
Terrestrial species not closely related to the Scottiinae have been found in widely scattered
parts of the world, and most are related to freshwater taxa. For example there are terrestrial
representatives of the subfamilies Callistocypridinae and Candoninae (family Candonidae)
and Terrestricypridinae (family Cyprididae), all belonging to the superfamily Cypridoidea,
as well as the family Darwinulidae (superfamily Darwinuloidea) (Pinto et al., 2003, 2004,
2005; Martens & Home, 2009). The route from freshwater, via damp mosses and leaf litter,
seems an obvious one. More surprising, perhaps, has been the discovery of a completely
separate superfamily, the Terrestricytheroidea, making the transition to terrestrial habitats
from marine and brackish water environments. Schomikov (1969) found in Iturup, one of
the Kuril Islands north-east of Japan, an ostracod that he described as Terre stricy there
pratensis, which he found living in a coastal locality among small pebbles which were
always kept damp by mist, rain and sea spray. The same author described a second species

The Ostracod Scottia pseudobrownian Kempf

of the genus from a monsoonal salt marsh with brackish groundwater at a coastal site near
Vladivostok, and also recorded T. pratensis living nearby among supralittoral salt-tolerant
plants (Schomikov, 1969). Two more species of Terre stricy there were discovered in
England, one (which remains un-named) represented by a single male specimen found in
an intertidal rock pool on the Bristol Channel coast, the other, described as T. elisabethae,
by a large population inhabiting a shaded locality on a tidal creek in Hampshire, where the
animals were found crawling among damp leaves (Home et al., 2004). More recently,
further species have come to light in Japan (Hiruta et al., 2007) and the Black Sea
(Schomikov & Syrtlanova, 2008). It is clear that adaptation to terrestrial habitats was an
entirely independent evolutionary venture for Terre str icy ther e , not only because it
originated in marine/brackish-water rather than freshwater habitats, but because it
constitutes a separate superfamily only distantly related to other terrestrial ostracods. While
the Terrestricytheroidea, Darwinuloidea and Cypridoidea all belong to the same order, the
Podocopida, each is very distinct. Although no fossil representative has yet been
recognised, the Terrestricytheroidea may be a very ancient superfamily, possibly hundreds
of millions of years old (Home, 2003; Home et al., 2004).

Convergent evolution of terrestrial and semi-terrestrial habits in

OSTRACODS AND OTHER SMALL CRUSTACEANS

The acquisition of terrestrial and semi-terrestrial habits by ostracods involved the solution
of ecological and physiological problems. Although small, the animals concerned are often
too large and bulky to live completely within the thin film of water that covers surfaces in
damp situations (which is otherwise sufficiently “deep” for protozoans, rotifers, tardigrades
and even harpacticoid copepods). Many, if not all, ostracods (aquatic as well as terrestrial)
are capable of surviving adverse conditions by sealing themselves tight shut inside their
carapaces, and this undoubtedly serves them well if terrestrial environments become too
dry. However, at least some terrestrial ostracods are capable of continuing to move in
relatively dry conditions, dragging with them a small bubble of water as they crawl, as was
observed and illustrated for Terre str icy there elisabethae by Home et al. (2004). In this they
are aided by having “hairy” carapaces, furnished with abundant sensilla that are
particularly densely spaced, and sometimes brush-like, around the ventral margins of the
valves, which help to retain water. However, while some taxa (e.g. Terre stricy there , some
species of Mesocypris and Scottia) are abundantly adorned with such sensilla, others,
including S. birigida and S. pseudobrowniana are not particularly well-endowed in this
respect although the latter, at least, displays rows of closely-spaced sensilla near the free
outer margins of the valves (Fig. 2D) . It may be that the degree of “hirsuteness” of these
ostracods is a reflection of how far along the adaptive path to terrestrial habits they have
progressed, but there is clearly much to learn about how these animals cope with their
environment.

Semi-terrestrial habits have evolved also in branchiopod crustaceans of the Order
Anomopoda that are very distantly related to ostracods and provide a remarkable example
of convergent evolution. The branchiopod exponents of this way of life are members of the
family Chydoridae which, like many ostracods, are predominantly crawlers. They live in
wet moss on forest floors and among mosses growing on tree trunks in cloud forests where
the air is saturated with moisture for most of the time. All are very small, which enables
them to live in small interstices among moss leaves where a film of water is easily retained.
Thus the first two terrestrial species to be described, by Frey (1980), which he assigned to a
new genus Bryospilus, are no more than 0.35mm in length and are among the smallest
known chydorids. Fike ostracods they have a functionally bivalved carapace, but in their
case this is a single, folded structure with no dorsal hinge, although its two valves can be
brought together by means of adductor muscles, just as can those of the hinged bivalved
carapace of ostracods. They crawl by the use of the trunk limbs, assisted at times by
levering with a median post-abdomen that is the functional equivalent of the posteriorly-
located furca (or uropods) of many cypridoidean ostracods, including terrestrial species.

The Ostracod Scottia pseudobrownian Kempf

The antennae, used for propulsion when anomopods swim (which most of them can), or,
less frequently, for levering them through detritus, are reduced in size and play no part in
locomotion. In this respect semi-terrestrial chydorids are unique among anomopods. The
small size of these animals presumably enables them to retain water within the carapace
chamber when living in a thin film of it. Bryospilus proves to be a very distinctive genus,
not closely related to previously known genera. Among its peculiarities is that it lacks the
single compound eye (derived from two fused eyes) possessed by most anomopods, but it
retains the small ocellus. The first semi-terrestrial chydorids, of which Frey (1980) gave
excellent illustrations, were found in damp forests in New Zealand, Venezuela and Puerto
Rico. Additional species, one of them allocated to a new genus, with similar habits, have
since been described from wet forests in West Africa (Chiambeng & Dumont, 1998).

The other group of small crustaceans that is well represented in freshwater, the
Copepoda, also includes a few species that have emancipated themselves from truly aquatic
conditions. Unlike ostracods and anomopods, copepods lack an enveloping carapace that
can retain water. One member of the Cyclopoida, Ectocy clops phaleratus, which has
creeping habits and can move over wet surfaces, pushing a film of water with it (as perhaps
can its congeners), may exhibit a potentially early stage in the evolution of terrestrial
habits. Already better- adapted to a semi-terrestrial life, however, are various smaller,
creeping species, often no more than 0.5 mm in length, that belong to several genera whose
relationships are uncertain. These, whose four pairs of locomotory thoracic limbs have
short rami, often reduced to two segments from the basic three, often live among mosses,
in springs, seepages and groundwater. British representatives are few but include
Speocyclops demetiensis, two of whose three recorded sites in Britain are in Yorkshire, and
Graeteriella unisetiger.

Perhaps even better exponents of such habits are found among the Harpacticoida,
freshwater representatives of which are predominantly crawlers rather than swimmers, and
of which some frequent wet moss or the leaf carpet of woodlands where they use their
thoracic legs as oars when they “row” through a thin film of water, or as levers where the
film is thinner. Species of Maraenobiotus and Epactophanes, both of which have British
representatives, live in wet (or sometimes even only damp) moss and occasionally among
decaying leaves or on moist peat. In Yorkshire two species of Moraria , one very common,
the other much less so, usually frequent Sphagnum, another has been found in a seepage,
and two other species of the genus have been found in the leaf carpet of woods (Fryer,
1993).

While of less relevance in the present context, for completeness it can be mentioned
very briefly that various larger, malacostracan crustaceans have also to a large degree
adopted terrestrial habits. Large size is beneficial in that, for example, surface area to
volume ratios are more advantageous in combating losses by evaporation and a thick, more
water-retaining cuticle can be employed than in minute animals - though terrestrial
crustaceans lack the waxy layer of epicuticle typical of other terrestrial arthropods. Gills
can also be housed in an enclosed humid chamber, as they are in crabs that adopt such
habits.

Most successful on land have been the isopods which, as well as being very diverse in
the oceans where they have, for example, given rise to a spectacular array of deep sea
forms, are well represented on land as the wood lice. Amphipods have been less successful
in this respect, though “beach-hoppers” are common on sea coasts, but a few species thrive
in the damp tropics and one semi-terrestrial invader, Orchestia cavimana , has been found
in Yorkshire. Various crabs are highly successful in terrestrial habitats - the Coconut Crab
Birgus latro even climbs trees - but many retain aquatic larval stages, which means that
adults have to return to the sea to reproduce and are therefore restricted in their ability to
venture far inland. Some freshwater crabs, which have eliminated larval stages from the
life cycle, roam freely on land, especially when conditions are moist. Perhaps surprisingly,
hermit crabs of the genus Coenobita have successfully colonised terrestrial habitats, where
they use the shells of terrestrial snails as “homes”, but have to return to the sea to breed.

The Ostracod Scottia pseudobrownian Kempf

Nevertheless the New World C. clypeatus is a highly successful species in the Caribbean
and adjacent mainland areas.

Tailpiece

When, just over 120 years ago, Thomas Scott discovered the ostracod whose name
perpetuates his memory, and George Stewardson Brady and Alfred Merle Norman
described it, they had no idea that it would eventually prove to be related to terrestrial
species whose existence was then not even suspected, and were not to begin to be known
until more than 60 years later. Nor did they know that their findings, with those of later
naturalists, would enable its ecological preferences, habits and morphological attributes to
be compared with those of other ostracods, both related and unrelated, and throw light on
the route or routes whereby these seemingly unlikely animals have come to occupy semi-
and fully terrestrial niches. In recalling this chain of events the role of three pioneers in the
study of small crustaceans is inevitably and happily remembered.

Acknowledgments

Andy Godfrey would like to thank the Environment Agency (York office) who funded the
project which was aimed at studying the ecology of the Red Data Book 1 and UK
Biodiversity Action Plan soldier-fly Odontomyia hydroleon. Werner Hollwedel kindly
informed us that the term “Schachtelhalm”, translated as “shavegrass” (which we were
unable to equate with any common English name), in fact refers to “horsetails”, the
common name for Equisetum. We thank Gilly Walsh-Kemmis (nee McCall) for supplying
coordinates for the Irish record of Scottia pseudobrowniana.

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(Cypridoidea: Ostracoda) from western Honshu, Japan and a key to the subfamily
Scottiinae Bronstein, 1947. Zootaxa 126: 1-20.

Swain, F.M. (1999) Fossil nonmarine Ostracoda of the United States. Developments in
Palaeontology & Stratigraphy 16: 1-401.

Triebel, E. (1941) Die ersten Ostrakoden aus der Paludinenbank. Z. Geschiebeforsch. 17:
61-75.

Whittaker, J.E. and Home, D.J. (2009) Pleistocene. In: Whittaker, J.E. & Hart, M.B. (eds),
Ostracods in British Stratigraphy, 447-467. The Micropalaeontological Society Special
Publications, The Geological Society, London.

BOOK REVIEW

Palms of Southern Asia by Andrew Henderson. Pp. 199 (inch line illustrations & 336
maps), plus 64 pages of coloured plates. Princeton University Press, Princeton & Oxford.
2009. £53.00 hardvack.

Although rather an exotic title for most readers of The Naturalist, this is an exemplary field
guide in presenting a wealth of taxonomic, ecological and biogeographical information
within the compass of a relatively small volume. The pages are packed with succinct but
informative data (including detailed keys and maps) on 352 naturally occurring palms in
southern Asia.

The author, Andrew Henderson (Curator at the New York Botanical Garden), will be
known to some of us as one of the speakers at the Linnean Society Conference on the
Yorkshire botanist Richard Spruce, held at York University in September 1993. His
contribution on ‘Richard Spruce and the palms of the Amazon and Andes’ is to be found in
Richard Spruce (1817-1893) Botanist and Explorer edited by M.R.D. Seaward and S.M.D.
Fitzgerald (1997) - see The Naturalist 122: 24-26 (1997).

MRDS

THE STATUS OF MOORLAND BREEDING BIRDS IN
THE PEAK DISTRICT NATIONAL PARK 2004

PETER MIDDLETON

33 Wilthorpe Road, Redbrook, Barnsley S75 1JA
and

GEOFF CARR

300 Higham Common Road, Higham, Barnsley S751PF

The decline of lowland farmland breeding birds in the UK is now well documented and
reasonably understood, but does this national trend also extend to upland breeding birds? A
survey of the upland breeding birds of the Peak District National Park was conducted in
1990 by AJF.Brown and K.Shepherd and various other sample surveys have been
conducted by the RSPB, the National Trust, the Peak District National Park Authority and
English Nature, but it is 14 years since the whole area was surveyed for upland breeding
birds. This article presents a summary of the results from a repeat Moorland Breeding Bird
Survey of the Peak District National Park in 2004 conducted by the Moors for the Future
Partnership and funded by the Heritage Lottery Fund.

The moors of the Peak District National Park present a diversity of breeding bird
habitats of national importance. Nevertheless, the Peak District moorland has suffered
significant losses and degradation of moorland habitats through fringe agricultural
improvement, past poor management of burning and grazing regimes and plantation
afforestation. Surrounded on all sides by the heavy industry of northern England, the
vegetation of the Peak District has also been severely degraded by atmospheric pollution.
Furthermore, the area is within close proximity to huge urban areas and this has resulted in
enormous visitor pressure. The designation as Britain’s first national park, rights of access
to large areas of wilderness through open access agreements and the increase in car
ownership has resulted in huge numbers of visitors engaging in an array of activities
throughout the year. In spite of this, the Peak District retains the most southerly viable
populations of typical moorland breeding birds in England. For a number of species,
particularly European Golden Plover Pluvialis apricaria, the South Pennines Peak District
may hold the most southerly breeding populations in the world.

The 2004 survey covered the unenclosed upland area of the Peak District National
Park. It was therefore mainly concentrated within the SSSIs of the Dark Peak (between
Edale in the South and Marsden in the north), the Eastern Moors and the South-West Peak
(Staffordshire moors between Buxton and Leek). The 1990 census covered the Dark Peak
and South Pennines as far north as Ilkley. Therefore to analyse the population trends and
spatial distribution of breeding birds it was necessary to deduct the data from the northern
section north of Marsden from the 1990 survey.

The Peak District vegetation

An important correlation exists between the breeding birds of the Peak District moorlands
and the vegetation of the area, which in turn influences both densities and the distribution
of typical moorland birds. The vegetation of the Peak District moors can broadly be
described in terms of five vegetation types, each associated with particular facets of
topography or management. The vegetation generally is severely degraded and much of the
SSSIs have been designated by English Nature as being in unfavourable condition.

On the plateau of the Dark Peak, where the depth of peat can exceed four metres, the
vegetation is dominated by Eriophorum vaginatum. Relatively huge expanses of this
vegetation exist on the moorlands and it is particularly characteristic of the South Pennines
blanket bog (Ratcliffe, 1997; Brown & Shepherd, 1991). E. angustifolium can be abundant
along the edges of pools and where the peat is eroded; typical associates are Calluna
vulgaris, Empetrum nigrum, Vaccinium myrtillus and Rubus chamaemorus. Although these

Naturalist 135 (2010)

36 The Status of Moorland Breeding Birds in the Peak District National Park 2004

blanket bogs are amongst the most species poor floras in Britain they are unique and
support high densities of European Golden Plover and nationally important numbers of
Dunlin Calidris alpina. The highest densities of both these species are found on
Eriophorum bog that contains abundant Empetrum nigrum and Rubus chamaemorus . The
latter is certainly characteristic of moorlands that support high densities of these waders. In
contrast there are extensive areas devoid of vegetation that are seriously eroding and in
these areas birds are at very low densities or absent. The present extent of peat occurred
some 4000 years ago and severe erosion is perhaps part of a natural cycle due to climate
change. However, this process has been accelerated by un-natural events such as moorland
fires, overgrazing, increased public access, atmospheric pollution and particularly acid
precipitation (Anderson & Yalden, 1998). The latter is thought to be responsible for the loss
of peat-forming sphagnum mosses in the mid- 18th century. Once peat is exposed it is
particularly vulnerable to erosion and oxidization that prevents colonisation of plants, thus
it remains bare and the process accelerates.

Away from the plateau the vegetation of the moorland slopes is mostly dominated by
Heather, and these areas are mostly managed for Red Grouse Lagopus lagopus scoticus. As
a consequence, the heather moors are burned at regular intervals to provide nutritious
young shoots for the grouse to forage. Empetrum nigrum and Vaccinium myrtillus are
common associates and the latter is often present in relative abundance. According to
Anderson and Yalden (1981) areas dominated by pure heather in the northern part of the
Peak District appear to have declined by some 35% since 1913. These losses are thought to
have resulted from past overgrazing and poor burning management. Peak District bird
surveys in recent years have revealed that these pure heather moorland or monocultures
attract only small assemblages of moorland birds and often only Red Grouse and Meadow
Pipit Anthus pratensis are present in such areas. However, recent heather bums,
particularly on wet moors with regenerating Eriophorum angustifolium, often attract
breeding European Golden Plovers. Emphasising those differences in land management
may affect breeding bird densities (Sim et al., 2005).

Much of the Heather moorland has been replaced by some form of acid grassland.
These areas of grassland are particularly favoured by Sky Lark Alauda arvensis, Eurasian
Curlew Numenius arquata and Meadow Pipit, the latter often found in local abundance.
Acid grassland is an underestimated resource, indeed, there appears to be a misconception
that extensive heather moorland is best for biodiversity and this is often not the case.
Molinia caerulea is often dominant on gentle, wetter slopes where it usually forms a
species poor, tussocky sward. Nardus stricta also forms smaller but similar tussocks and
this type has probably spread most in recent times as a consequence of heavy grazing, this
grass is unpalatable to sheep. The most interesting and species rich acid grassland type
found in the Peak District is Deschampsia flexuosa. It often occurs on the steeper ground
and it is particularly palatable to sheep, this is perhaps the reason why this grassland type is
not more abundant in the area.

J uncus spp. are also common throughout all areas of the Peak District where water is
abundant at the surface. J. efjusus is the most abundant and occurs in the damp places
particularly where there is a marked change in slope, and variable sized areas of Juncus
also occur alongside streams. Two species of birds are associated with this habitat, notably,
Common Snipe Gallinago gallinago and Reed Bunting Emberiza schoeniclus.

Cloughs, tors and gritsone edges, with their associated rock litter are often dominated
by Pteridium aquilinum which may invade heather or acid grassland. This vegetation type
is a very important habitat for typical moorland birds such as Northern Wheatear Oenanthe
oenanthe , Whinchat Saxicola rubetra, Common Stonechat S. torquata and Ring Ouzel
Turdus torquatus. Whinchats are never found away from this vegetation type and Twite
Carduelis flavirostris also have a strong association with it.

Clough woodlands containing Quercus sp., Betula pendula and Sorbus aucuparia
occur in the Peak District but generally they are considered by many conservationists as a
threat to the important moorland habitats. Nevertheless, natural regeneration of managed

The Status of Moorland Breeding Birds in the Peak District National Park 2004

and fenced areas has occurred in some cloughs and as a consequence, an assemblage of
birds has colonised these areas. Lone trees high in cloughs have been found to attract
breeding Willow Warblers Phylloscopus trochilus. Chaffinches Fringilla coelebs and Tit
Parus species are also attracted to such locations. The survey also revealed that Woodcocks
Scolopax rusticola and Redstarts Phoenicurus phoenicurus breed in the wooded cloughs
and, as these trees become mature, they may attract Wood Warblers Phylloscopus sibilatrix
and Pied Flycatcher Ficedula hypoleuca.

Results

During the 2004 survey, data were gathered concerning the distribution and numbers of 39
species of breeding birds, unlike the 1990 survey when data were presented for 27 species.
However, not all the species are considered typical upland breeding birds therefore
discussion on the numbers and distribution change will be largely centered upon what is
considered to be typical of moorland. Notable exceptions include some wildfowl species,
Common Buzzard Buteo buteo and Reed Bunting. Three species of wildfowl were found
breeding on the Peak District moors and increases in their numbers have occurred since
1990. Three pairs of Teal Anas crecca were found in 2004 compared to none in 1990 and
likewise, the breeding population of Mallard Anas platyrhynchos has increased from two
pairs in 1990 to 23 pairs in 2004. The largest increase is not surprisingly of Greater Canada
Goose Branta canadensis. Three pairs were present in 1990 compared to 56 breeding pairs
in 2004, also many more birds were present on the reservoirs. These birds were found
grazing at all altitudes and therefore any increase in population could result in additional
pressure on the moorland vegetation.

Raptors have also seen similar increases in population since 1990. However, the
dramatic increase in their population and distribution has followed the cessation of much of
the pesticide usage originally responsible for their population crash. Peregrine Falcon
Falco peregrinus and Merlin F. colombarius were particularly affected and it was not until
the 1980s that a slow recovery began. Both species have increased their populations and
distribution in the Peak District since 1990. Peregrines have increased from seven pairs in
1990 to 25 pairs in 2004. Likewise the Merlin has increased from 15 pairs to 31 pairs.
Common Buzzards have also increased considerably since 1990 when only a single bird
was recorded during the entire survey. During the 2004 survey Common Buzzards were
recorded in 20 1 km2 and two pairs were confirmed as breeding. This species nevertheless
remains a scarce breeder in the Peak District in spite of being a common resident in the
lowlands of Derbyshire. Likewise, there are also limiting factors inhibiting the successful
colonization by Hen Harrier Circus cyaneus. This species successfully bred in the area in
1997 after an absence of approximately 200 years (Frost, 1978), however, the
establishment of a small breeding population has not followed.

Red Grouse is a species synonymous with heather moorland, and where this habitat
exists, their densities are highest. Management of the moors appears to be the key to the
species range and abundance over time and therefore the heavily managed moors have
densities in excess of 50 pairs/km2. Interestingly, on the heather monocultures where
densities of Red Grouse are greatest the overall biodiversity is far less and often the only
other bird species present is Meadow Pipit. A total of 2337 individuals was recorded in the
study area during the survey in 1990 compared to 5416 individuals in 2004. The increase in
the population is dramatic and therefore may reflect the intensive management that occurs
on keepered estates; burning is much more evident than a number of years ago. Some
caution must be applied when considering the present increase in numbers because these
can vary significantly from year to year resulting from a variety of factors, particularly
intestinal parasites. The distribution of Red Grouse has also changed considerably and
birds were found in 422 km2 in 2004 compared to 324 in 1990.

Waders overall, within the Peak District National Park have remained stable or
increased in numbers; the only exception is Dunlin where the losses are directly correlated
with the loss of suitable habitat in the Bleaklow area. This loss is nonetheless significant

38 The Status of Moorland Breeding Birds in the Peak District National Park 2004

because Dunlins were found in only 33 1 km2 in 2004 compared to 54 1 km2 in 1990. This
species is found exclusively on blanket bog, preferably with pool complexes. A total of 91
pairs of Dunlins were recorded during the survey in 1990 compared to 67 in the same area
in 2004. The data suggest that there has been a reduction of 25% in the breeding population
of the Dunlin in the study area over the period. The average density throughout the study
area was 2.27 pairs/km2 and the highest density was 4.2 pairs/km2 in prime habitat on
Howden Edge.

European Golden Plovers were common throughout the study area in areas above
400m and where the vegetation is dominated by Eriophorum vaginatum. Birds were also
found nesting on areas of recently burnt heather moorland. Their distribution has changed
little: a total of 419 pairs was recorded in 2004 compared to 436 in 1990. The data suggest
that the population has remained stable; however there is clearly a paucity of records in the
east of the Dark Peak where Heather dominates; therefore, the species has a distinct
westerly bias. The average density of this species in the study area was 1.90 pairs/km2.

The Brown and Shepherd (1991) methodology used for this survey is clearly
inadequate for censusing Common Snipe Gallinago gallinago. Nevertheless, the same
methodology was used in 1990 and 2004 therefore one can still determine the trend for this
species in the Peak District National Park; 135 territorial males were recorded in the survey
area in 2004 compared to 56 in 1990. The birds recorded may well be a fraction of the true
population and this species is perhaps the most common breeding wader in the Peak
District National Park.

The Curlews range and population in the Peak District appears to have enjoyed a
significant increase since 1990; 259 pairs were recorded in the study area in 1990
compared to 453 pairs in 2004. Furthermore, birds were found in 288 km2 in 2004
compared to 189 km2 in 1990. There appears to be no obvious preference of nesting habitat
and the species is evenly distributed throughout the study area, the only requirement of
habitat is its proximity to inbye land where this species does much of its foraging.
Consequently, Eurasian Curlews are largely absent from the central massifs of the Kinder
and Bleaklow areas. The average density of Curlews in 2004 was 1 .54 pairs/km2.

An increase in population and distribution is also evident for Short-eared Owl Asio
flammeus and Raven Corvus corax , Indeed, Ravens have undergone a successful
colonisation of the district since 1990 when Brown and Shepherd (1991) stated that ‘there
seems little immediate prospect of the return of this species’. This emphasises how the
fortunes of some species can change dramatically in only a very short time and how little
we know about the changing numbers and distribution of birds. There has been a gradual
increase in the population of Ravens since the first confirmed breeding in 1992 and the
estimated breeding population in 2004 is in the region of 65 pairs, however, only 18 pairs
were confirmed as breeding. Short-eared Owls have also increased significantly from 18
pairs in 1990 to 27 pairs in 2004. This species is well known for considerable fluctuations
of breeding pairs that can occur from one year to the next depending on the vole ( Microtus
agrestis ) population.

The smaller passerines do not all share the upward trend in numbers of the species
already described. Indeed, some have suffered serious declines in numbers and distribution.
However, this cannot be said for both Whinchat and Common Stonechat. The former has
increased by more than 60% from 59 pairs in 1990 to 97 pairs in 2004. This species
favours the bracken covered lower slopes of the moorland fringe where they can be easily
detected by there loud characteristic song that is delivered from a prominent perch. One
reason for this rise may be an increase in suitable habitat resulting from the spread of
bracken since 1990. Even more pronounced is the increase of the Common Stonechat.
Interestingly, no breeding Stonechats were found in the Peak District in 1990, but since that
time colonisation has been dramatic and 91 pairs were recorded in 2004. Stonechats were
often found at higher altitudes than Whinchats but unlike the latter, there appears to be no
obvious preference of vegetation type for nesting. Northern Wheatear is also a species that
inhabit all of the upland area, but in the Peak District is often associated with rocky areas

The Status of Moorland Breeding Birds in the Peak District National Park 2004

of moorland. The numbers of territorial Wheatears recorded in 2004 was more than half the
number recorded in 1990, only 45 pairs compared to 109 in 1990. This species is not
considered to be of conservation concern, nevertheless, Sim et al. (2005) also found a
similar significant decrease in the South Pennines between 1990 and 2000.

In contrast, Ring Ouzel is a Red Listed species of high conservation concern due to its
serious breeding population and range decline over the last 25 years. Interestingly however,
no significant decline was witnessed in the Peak District between 1990 and 2004 (98 and
83 pairs respectively). Although a slight decline of 15% was proven from the data
collected, it is assumed that this species also occurred in reasonable numbers in the doughs
of the private grouse moors where access was denied. The distribution of this species has
however changed a little and in 1990 it was notably absent from the eastern moors, but this
was not the case in 2004. This is perhaps not significant as the birds will take up territories
wherever the opportunity arises as they do not appear to be site faithful. This was probably
proven during the foot and mouth outbreak in 2002 when many pairs bred on Stanage Edge
when access was denied to climbers. This species is especially vulnerable to disturbance;
therefore they do not usually nest in areas frequented by recreational users.

Of all the species recorded during the survey the biggest and most alarming change
was of Twite. This is a Red Data Book species (Batten et al., 1990) and a Red Listed
Species of conservation concern due to an historical decline in the UK: a total of 136 pairs
of Twites were found in the study area during the survey in 1990, compared to only ten
pairs in the same area in 2004. The species was found at three localities within the study
area: Wessenden and Melthem moors west of Huddersfield, Edale in the extreme south of
the Dark Peak and the Combs Moss area in Staffordshire. Twites are thought to breed in
loose colonies therefore it is unlikely that these small numbers reflect the true breeding
population, nevertheless, the decline in the breeding population has been catastrophic.
Anecdotal evidence does however exist of a slightly larger population in Staffordshire than
the data suggest. Furthermore, small populations were known at two other localities in the
northeast of the study area in 2004, Winscar west of Barnsley and Black Hill near
Holmfirth (pers. obs.). The decline of the British breeding population has been attributed to
the deterioration in quality and quantity of the inbye land, in particular the seed rich hay
meadows in which the species feeds in summer (Brown et al., 1997). However, the
explanation is perhaps more complex: indeed, a small study by English Nature conducted
in 2003 revealed that the species preferred to feed in semi-improved pasture with a higher
nutrient level resulting in an abundance of Common Sorrel Rumex acetosa (Middleton,
2003). The fact remains that this species has vanished from areas that appear to provide
prime habitat {pers. obs.). This species has a strong preference for nesting in bracken;
indeed, there are many bracken-covered areas of moorland adjacent to seed rich hay
meadows in the study area that no longer support breeding Twites which were found in
only seven 1 km2 during the 2004 survey, compared to 88 in 1990. The species is, however,
known to breed in a few other localities within the study area. Nevertheless, Twites have
vanished from former haunts and their range contraction is dramatic. In 1990 the species
bred in 14 tetrads in the Sheffield area and ten years previously they nested in 26 tetrads.
The decline has continued and they are literally ‘hanging on’ in a few localities in the south
of the study area.

Brown and Shepherd (1991) stated that the Twite was thinly distributed throughout the
study area following the survey in 1990; furthermore, although they had declined in the
Sheffield area there was no evidence to suggest any marked change in its distribution in the
study area. The birds were said to be more thinly distributed south of the line between
Stalybridge and Holmfirth and it is therefore interesting to note that south of this line is
where they have declined most in the intervening time. If the birds are indeed loose colony
nesters, could it be possible that sites are abandoned once the minimum critical level is
reached? There is perhaps little evidence to support this hypothesis at present, but recent
ringing studies in the South Pennines may provide more information. Following the survey
in 1990, Brown and Shepherd (1991) estimated the total South Pennines’ population at

40 The Status of Moorland Breeding Birds in the Peak District National Park 2004

around 415 pairs. This was perhaps however an underestimation of the true breeding
population at that time. Interestingly, the population at present is estimated between 200
and 500 pairs (RSPB 2001), so if Brown and Shepherd (1991) were accurate in their
estimation, perhaps the species has not suffered a population decline but perhaps a
pronounced range contraction. English Nature conducted a small study in 2003 that
revealed that it was indeed abundant in certain parts of the Pennines and that there was a
‘hot spot’ for this species in the circle of moorland between Marsden, Halifax, Todmorden
and Rochdale (Middleton, 2003). Radiating away from this area the birds become more
thinly distributed. Considering that Brown and Shepherd (1991) stated that the birds were
thinly distributed throughout the study area, it is worth considering that perhaps the Twites’
overall population in the South Pennines may have remained stable, and it is just their
distribution that has changed. Indeed, they have vanished from areas that provide prime
habitat; therefore their foraging requirements are perhaps not the main reason for the
decline. A study by Batty et al. (1999) provided evidence of considerable distribution
change since the 1990 Brown and Shepherd survey. Whilst Twites were found in only 20%
of the squares in which they were recorded in 1990, they were found in 18% of the sample
of squares where they were not recorded in 1990. This might suggest that the species has
undergone some redistribution. Most important, however, is the finding that the increase in
densities in occupied squares has increased from 2.91 to 5.75. This again, is evidence that
may suggest the species has undergone a range contraction rather than a decline in
population.

Summary

Providing evidence of increases in the distribution and numbers of many moorland
breeding birds in the Peak District National Park is a pleasing outcome from the survey that
is in direct contrast to the decline of many lowland farmland birds in the UK. Indeed, many
of the birds in the study area have remained stable or increased, and of the 27 species that
were recorded during both the surveys of 1990 and 2004, only eleven species have suffered
a decline and only nine of those have undergone a significant decrease (see Table 1).

There is perhaps little doubt that densities of Red Grouse of the race scoticus,
indigenous to the British Isles, are found nowhere in such abundance as on the eastern area
of the Dark Peak. However, although this species appears to have doubled over the last 14
years, the figures mean little when the species is managed and kept in such artificially high
numbers; for example, 2005 was a bad year for this species and the numbers were
subsequently much reduced.

What is however, of great consequence is that many species of national importance
have remained in high densities, such as European Golden Plover. This species is at the
southern extremity of its range and its population has remained stable, therefore there is
perhaps little evidence to suggest any affects from global warming. Other species have
increased in range and numbers such as Peregrine Falcon, Merlin and Short-eared Owl.
Furthermore, Raven has colonised the area since 1990 and Hen Harrier is perhaps
attempting to re-colonise. Waders have also fared reasonably well; Dunlins are slightly
down in both range and numbers resulting from loss of suitable habitat in the Bleaklow
area, but Eurasian Curlew, Northern Lapwing and Common Snipe have all increased in
numbers.

There is no real evidence to suggest that the status of Ring Ouzel has changed since
1990, whilst Whinchat, Common Stonechat, Reed Bunting and Dipper have all increased in
both range and numbers. There is perhaps little change for Sky Lark and Meadow Pipit,
especially as these species were not recorded in the same detail as other species.

The declines of some species are perhaps less evident but nonetheless significant.
Common Cuckoo Cuculus canorus has undergone a massive 69% reduction in numbers in
14 years that is greater than the national trend (Glue, 2006). Northern Wheatears were also
much reduced in 2004, but most notable was the catastrophic reduction in both range and
numbers of Twite .

The Status of Moorland Breeding Birds in the Peak District National Park 2004 41

Table 1 . 2004 Peak District Moorland Breeding Bird Survey

Species

Breeding Pairs

Occupied km2

Percentage

Change

1990a

2004a

1990a

2004a

Abundance

Occupancy

Common Buzzard*5

1 18(2)

1700%

Greater Canada Goose

1800%

825%

Carrion Crow

117

-91%

-84%

Common Sandpiperd

100%

Common Cuckoo

-69%

-71%

Eurasian Curlew

259

453

189

288

-75%

-52%

Dipper

75%

67%

Dunlin

-26%

-39%

European Golden Plover

436

419

225

218

-4%

-3%

Grey Wagtail

24%

41%

Common Kestrel*5

84(5)

24%

13%

Northern Lapwing

131

115%

48%

Meadow Pipitc

10410

8432

476

472

-19%

-1%

Merlin*5

21(9)

31(6)

48(50)%

43%

Peregrine Falcon*5

7(3)

25(5)

286(40)%

257%

Red Grousec

2337

5416

324

422

132%

30%

Common Redshank

-33%

-50%

Reed Bunting

143

794%

600%

Ring Ouzel

-15%

-6%

Short-eared Owl*5

18(5)

27(18)

50(260)%

50%

Sky Larkc

1153

1069

311

306

-7%

-2%

Common Snipe

135

141%

95%

Common Stonechat6

Twite

131

-92%

Northern Wheatear

109

-59%

-43%

Whinchat

64%

73%

a data for only those 489 km2 surveyed in both periods
b raptor sightings (in brackets confirmed breeding pairs)
c number of individuals, not pairs (1990 counts in broad classes, 2004 actual)
d indication of change not reliable, as species associated with reservoirs not included in
surveys

e no records of Common Stonechat during 1990 survey

42 The Status of Moorland Breeding Birds in the Peak District National Park 2004

Perhaps some of the declines were predictable, especially in the case of Dunlin,
because large areas of blanket bog in the Bleaklow area have deteriorated to such an extent
that they have become devoid of vegetation and are eroding. The purpose of the Moors for
the Future Project is to preserve and enhance the moors for wildlife but also for future
generations to enjoy, therefore work in the Bleaklow area to restore and revegetate the
badly eroding moors has begun and already there are signs of success. Furthermore, paved,
formally eroded footpaths have surprisingly had the positive effect of increasing numbers
of breeding waders in these areas and both Dunlins and European Golden Plovers were
found nesting extremely close to the Pennine Way. Indeed, some of the work to be
addressed by the MFF project will be to repair such badly eroded rights of way. Therefore,
if the long term aim of the ‘Moors for the Future’ project is achieved then perhaps we can
look forward to further increases in the range and numbers of some of our nationally
important species in the Peak District National Park.

Acknowledgements

The authors would like to thank the land owners who gave access to their land, the
surveyors, English Nature, RSPB and the ‘Moors for the Future’ team.

References

Anderson, P. and Yalden, D.W. (1981) Increased sheep numbers and the loss of heather
moorland in the Peak District, England. Biological Conservation 20: 195-213.

Batty, R., Langston, R.H.W. and Gregory, R.D. (1999) South Pennines Twite survey 1999.
RSPB, Sandy.

Brown, A.F. and Shepherd, K. (1991) Breeding birds of the South Pennine Moors. JNCC,
Peterborough.

Frost, R.A. (1978) Birds of Derby shire . Moorland, Buxton.

Glue, D. (2006) Cuckoos in crisis? BTO News 263: 22-23.

Middleton, P. (2003) Survey of moorland fringe wading birds and Twite around the South
Pennine moors SSSI/SPA/cSAC. Unpublished report to English Nature
Ratcliffe, D.A. (1997) A Nature Conservation Review. Cambridge University Press,
Cambridge.

RSPB (2001) Twite inventory, Peak District to M62. RSPB, Denby Dale.

Sim, I.M., Gregory, R.D., Hancock, M.H. and Brown, A.F. (2005) Recent changes in the
abundance of British upland breeding birds. Bird Study 52: 261-275.

BOOK REVIEWS

Grasses of the British Isles by T.A. Cope & AJ. Gray. Pp. 612, with numerous line
drawings. Botanical Society of the British Isles. 2009. £19.99 (plus p. & p), paperback with
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This is the latest (No 13) and largest (in terms of content) of any of the BSBI handbooks,
and has been eagerly awaited for some time. The previous definitive and authoritative text,
Hubbard’s Grasses, upon which many of us relied for the last fifty years or so, has been
somewhat overtaken by advances in grass taxonomy and the need to develop more
workable and user-friendly keys to identification. The present volume can be best summed
up in two words - detailed and comprehensive - since it contains descriptions of 220 native
and introduced grass species including subspecies, hybrids and varieties, together with line
drawings of each species. It also contains detailed sections on the morphology,
reproduction and fruiting characteristics of the grass plant; classification from the
subfamily level, with very explicit keys for tribes, genera and species; and a very useful
section on how to use the book and interpret the species descriptions. There is a
comprehensive glossary of terms, bibliography and index at the end of the book.

Book reviews

The major part of the book deals with 220 species, covering the newly coined native
and casual categories as identified in New Atlas of the British & Irish Flora (2002). Within
each species, the textual content follows a clear and well-defined sequence, usually in five
paragraphs. Paragraph one gives a formal description of the structural and physical features
necessary to identify the species; paragraph two deals with the plant’s distribution and
habitat; paragraph three is concerned with its biology and ecology, discussing annual,
biennial and perennial status, reproductive and dispersal strategies; paragraph four looks at
its status, i.e. native or introduced in the British Isles, and its wider distributional (i.e.
European) context, and the IUCN categories of conservation importance, as outlined in The
Vascular Plant Red Data List for Great Britain (2005), i.e. Critically Endangered,
Endangered, Vulnerable and Near Threatened, are given for those species, such as
Calamagrostis scotica , which are included on the Red Data list. The page reference is also
given where the species can be found in New Atlas of the British & Irish Flora (or in its
accompanying CD). The species accounts are accompanied by superbly executed pen and
ink line drawings of all parts of the plant requisite for accurate identification. These
illustrations, by Margaret Tebbs, who is internationally acclaimed as a botanical artist, are
among the best I have ever seen of any plant material.

The very comprehensive section on Keys (pp. 34-103), may appear, at first glance,
somewhat daunting to the average botanist. The introduction of keys on Tribes will be
unfamiliar to many of us in Great Britain, and has been developed from Tom Cope’s
familiarity with writing Tropical Grass Floras, where Tribes and, indeed, Genera, are just as
important as species. Do not be put off, however, by the amount of detail and the
sometimes difficult terminology employed in the keys. I did a trial run, employing the
keys, on one or two specimens of well known grasses in my herbarium, and I have to say
that they seem, with practice, to work quite well. As with anything else, one needs practice
to assimilate and become familiar with new techniques, and I think this new system of keys
will, in time, largely remove the risk of going off up ‘blind alleys’ that we have all done
when using various grass keys in the past. To supplement the section on Keys, there is also
a short discussion on Classification, in which the characteristics of the six sub-families
making up all grass genera are highlighted. The last part of the comprehensive introductory
part of the book deals with structure and reproduction, covering the following features:
root, stem and leaf; inflorescence; spikelet; reproduction and breeding behaviour; vivipary
and proliferation; fruit; and anatomy and metabolism. Some of the terminology within this
section is somewhat esoteric, but, by and large, the average reader can gloss over these
areas, with little detriment.

This is an eminently useable book, of a convenient size to be carried in the field and
with a plastic cover to minimise wear and tear. It is a very worthy successor to Hubbard’s
Grasses , and the authors are to be congratulated in producing what is now and will
continue to be the standard identification text for grasses in the British Isles for many years
to come. I recommend it to amateurs and professionals alike who have an interest in or
involvement with the study of grasses or grassland ecology.

GTDW

The Scramble for the Arctic by Richard Sale and Eugene Potapov. Pp.223, inch b/w
illustrations, plus 20 colour plates. Frances Lincoln, London. 2010. £16.99 hardback.

This is a readable, fascinating, yet sobering book which takes a global perspective. Each
chapter recounts how the different countries possessing arctic regions have dealt with
particular issues. The book begins with an historical account of how humans moved into
and settled in the Arctic. It ends with an account of the first contacts between the aboriginal
peoples and the explorers sent out by countries to the south. The following chapter
documents how these countries took ownership of arctic lands and how the indigenous
people suffered. If you wondered how the USA rather than Britain came to purchase Alaska
and why, you will find interesting insights here into politics and power struggles. The

Book reviews

attempts to reach the North Pole and navigate the Northwest Passage are described. The
gradual regaining, by indigenous people, of some autonomy is compared between the
various countries with arctic lands. The key treaties which led to current ownership of
particular arctic island and land masses are mentioned.

The central chapters in the book deal with exploitation. The first phase was for furs and
one is astounded by the enormous number of animals that were sacrificed. It was only after
Prince Albert popularized the wearing of silk top hats that the demand for beaver pelts to
make hats dropped. The development of central heating and the economic crash of the
1930s further reduced the demand for furs in the West. The Arctic was considered of
relatively little importance after the Second World War so that nuclear bombs were tested
there by both the Russians and the Americans. The former did value the extraction of coal
in the Arctic and there had been a gold-rush in Alaska earlier; both are mentioned. A second
wave of exploitation starting around the 1960s involved mineral extraction, which
expanded in all the arctic countries in spite of the problems of dealing with the climate,
waste and local inhabitants. Finally it was realized what a rich resources of oil and gas lay
in the Arctic. Countries rushed to exploit these huge reserves which are documented in the
book. Conservation issues and protests by aboriginal people are described. There is a
chapter on international law in the Arctic which describes how countries hammered out
agreements on environmental protection for the Arctic, the law of the sea, and the
Northwest and Northeast Passages.

The final chapter deals with the future and it is clear that the Arctic is one of the most
fragile ecosystems in our planet, vulnerable to pollution, a thinning ozone layer, and global
warming. The potential danger of the release of frozen gas hydrates and methane from
permafrost with their capacity to exacerbate global warming needs to be appreciated more
widely. The likely rush to exploit the most northerly reserves of oil and gas to fuel a world
that refuses to address energy conservation is likely to lead to another ‘scramble for the
Arctic which will result in a new and far more dangerous cold war’. This book is illustrated
by 20 colour photographs and a few black and white illustrations. It is packed with
fascinating information on wildlife and is highly recommended to naturalists, students and
teachers, as well as to a wider readership.

DHSR

The Buildings of England. Yorkshire West Riding by Peter Leach and Nikolaus
Pevsner. Pp. xx + 824, incl. numerous line drawings, plus 64 pages of full colour plates.
2009. Yale University Press, New Haven & London. £29.99 hardback.

Naturalists who spend a significant amount of time in the field usually appreciate their
cultural as well as their natural heritage, often working in a wide range of habitats
supporting historically and architecturally interesting buildings. For those, like me, whose
special interests are attracted to churchyards, a copy of the relevant ‘Pevsner’ has been an
essential item in their field kit; the useful detail contained therein, particularly geological
and chronological, is of great importance to environmental interpretation. A revision of the
West Yorkshire volume (essentially vice-county 64), the last edition appearing in 1967, is
long overdue. For a more balanced view of our heritage, the complementary use of such
texts with the usual armoury of field guides is strongly recommended. Remarkable value -
a mine of information contained in a reasonably handy volume.

MRDS

Irish

Naturalists’ Journal

The Irish Naturalists’ Journal, sucessor to the Irish
Naturalist, commenced publication in 1925. The quarterly
issues publish papers on all aspects of Irish natural history,
including botany, ecology, geography, geology and zoology.
The Journal also publishes distribution records, principally
for cetaceans, fish, insects and plants, together with short
notes and book reviews.

Current subscription rates for four issues (including
postage): €33.00 (£20.00 stg); Students €11.00 (£7.00 stg).
Further details from: Mr Brian Nelson, INJ, Department of
Zoology, Ulster Museum, Botanic Gardens, Belfast
BT9 5AB.

Titus Wilson

Kent Works • Bumeside Road • Kendal • Cumbria • LA9 4RL
Tel. 01539 720244

Specialist printers /binders of
Academic Journals, Catalogues and Private Publications

Our service includes attending to worldwide distribution

Binding

Why not have your copies of The Naturalist bound into
volumes? One year’s issues per volume, or alternatively two
years in one volume at less cost than binding as two separate
volumes. We are also experienced in binding theses and
experts in the re-binding and repairing of all books.

Spink & Thackray
Broomfield Binder

Back Broomfield Crescent Telephone: 0113 2780353

LEEDS LS6 3BP Fax 0 1 1 3 2304 1 55

www .spinkandthackray .co .uk

Printed in Great Britain by Titus Wilson & Son, Kendal

ISSN 0028-0771

SMITHSONIAN INSTITUTION LIBRARIES

3 9088 01535 1356

ALIEN PLANTS OF YORKSHIRE

GEOFFREY WILMORE

The first comprehensive modem publication dealing with the alien
plants of any major British county, detailing the flora (almost 1300
species) of Yorkshire’s wool waste dumps, railway sidings, sewage works,
shoddy fields, industrial wasteland, dockland and oil mill sidings.

Pp. 316, including 15 line drawings; published 2000
Normal price: £11.00 (plus £2.50 p.&p.)

Reduced price for members of the YNU: £9.00 (plus £2.50 p.&p.)

Please make cheques payable to Yorkshire Naturalists’ Union.

Available from: Mr John A. Newbold, Stonecroft, 3 Brookmead

Close, Sutton Poyntz, Weymouth, Dorset DT3 6RS

THE FRESHWATER CRUSTACEA OF YORKSHIRE

a faunistic & ecological survey

GEOFFREY FRYER

The cmstacean fauna of Yorkshire reflects the great physiographic diversity of
the region. Adopting an ecological approach, this book considers the Yorkshire
fauna in relation to climate, topography, geology, soils and water chemistry,
always keeping in mind that it is dealing with living organisms whose habits,
requirements and physiological limitations determine exactly where they live.
Matters covered include the ecological background; faunal assemblages and their
regional attributes; an analysis of the factors that determine distribution patterns,
many of which are mapped; wide geographical aspects; and conservation. Large
areas, such as the Pennines, Howgill Fells, North Eastern uplands and the lowland
plains are surveyed. So too are localised regions including Whemside, the Malham
area, lowland heaths, and the largest lakes, as well as habitats such as upland tarns,
seepages, cold springs, small lowland ponds, inland saline waters. Notes are given
on every species recorded, including parasitic forms.

Price £5.00 (plus £2.50 p.&p.) Special offer to members of the Yorkshire
Naturalists’ Union £3.50 (plus £2.50 p.&p.)

Please make cheques payable to Yorkshire Naturalists’ Union.

Available from: Professor M. R. D. Seaward, Department of Geography
and Environmental Science, University of Bradford, Bradford BD7 1DP.

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