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August 2015
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Number 10®

Union

Journal of Natural History for the North of England

The Naturalist

Vol. 140 No. 1089 August 2015

Contents Page

From Yorkshire to China via Borneo: a biological excursion through tropical moth 81
ecology (YNU Presidential Address) Terry M. Whitaker

Nine years of change in the flora of Ellerburn Bank, a limestone grassland in the North 96
York Moors* Peter J. Mayhew, Susan E. Firth and Paul R. Waites

Local effects of climate change - has the date of first emergence changed in several 112
species of Lepidoptera in Yorkshire during the period 1995 to 2014? *

David R. R. Smith and Heather A. R. Smith

A Question of Ecology - answers from biological recording Paula Lightfoot 119

An investigation of the caddisfly (Insecta: Trichoptera) fauna of the Malham Tarn 121
NNR; with special reference to the Malham Sedge Agrypnetes crassicornis

S. Flint and P.W.H. Flint

Notes on the dolichopodid flies of two contrasting Yorkshire bogs Roy Crossley 128

Geological and land use influences on Badger sett densities across South Yorkshire 132

Colin Howes

Increase in bog-mosses Sphagnum and other changes in the vegetation of 134

Ringinglow Bog (Southern Pennines) since the 1940s R. Goulder

Additions and corrections to the Yorkshire Diptera list (part 6) Andrew Grayson 145

YNU VC63 Field Excursion to Thorpe Marsh 14th June 2014 Bryology Report 152

Colin Wall

Botanical Report for 2014 Phyl Abbott, Richard Middleton, Gill Smith & Linda 153

Robinson

Book Review 159

YNU Calendar 2015 160

Notices:

Erratum pl31

YNU Annual General Meeting pl58

An asterisk* indicates a peer-reviewed paper

Front cover: Fly Orchid Ophrys insectifera, a rare plant of Ellerburn Bank YWT reserve in the North York
Moors (see p97). Photo: P. Mayhew.

Back cover: Potamogeton lucens bed in Malham Tarn, looking towards Tarn House and the East Boathouse
(see pl26). Photo: S. Flint.

ipsasi

^ Union

HEALTH AND SAFETY GUIDANCE

Potential hazards and procedures to consider before
starting any field work or wildlife survey

Please spend a few minutes reading this guidance. If you
have any questions, please ask the organiser/leader. Health
and safety is an individual’s responsibility and each person
is responsible for drawing the attention of the group to any
hazard observed during the course of the work.

A risk assessment should be carried out before field work is started. Consider
the following:

General points

Appropriate clothing. Consider the weather forecast and the terrain to
be covered, including scrubby and prickly vegetation.

Sun protection cream, even in cloudy weather.

Sufficient drinking water, even on cool wet days.

Ensure supplies of any required medicine e.g. hay fever tablets. Diabetics
should have adequate food and medication.

An up to date tetanus vaccination is advised.

Have a small first aid kit available.

Latex gloves are useful in handling biological material, e.g. dead animals or
dung or when working in water.

Working in pairs/lone working

It is advisable to work in communicative pairs whilst surveying, i.e. in earshot of
each other, but there are times when this is not possible. Consider the following
issues:

• Be aware of your surroundings and be conscious of the dangers
associated with the different natural habitats in which you are working,
e.g. fissures in the ground hidden by vegetation, wet flushes and fallen
wet timber. Many places are on steep slopes, be aware of cliffs or wet
rocks or flushes. Remain aware of the surroundings of the site.

• Isolation: Check your mobile telephone is packed, charged and is within
range of a mast. Carry a card for telephone boxes and have change
available in case you need to ask another person to telephone on your
behalf. Check the location of the nearest settlement.

• Advise a reliable contact of the exact site where you are working, its
location and likely route and parking place. Advise the person of the
estimated time you will leave the site. Most organisations have an
agreed contingency plan should you not return. Call your contact on
leaving the site.

Surveying along roads and railways

When surveying hedges or verges running along a road or a railway, consider
the following:

• Wear fluorescent clothing for visibility.

• Try to keep off roads and work from the verge.

• Place warning signs 100m either side of a survey area where there is
no verge. Consider the need for a look out.

• Remain aware of where you are.

• If the survey involves a railway, you need consent of Network Rail or
other track owner.

•

Fauna and flora

Most wildlife and livestock are not a threat to humans or will avoid humans, but
in some situations you may become bitten or be stung. Consider the following:

• Avoid startling animals and livestock where possible.

• Some plants and caterpillars release irritants and toxins.

• Are there deer or sheep in the area? If suspected, protect the skin from
ticks to reduce the risk of Lyme Disease. Check yourself thoroughly
after returning home. If you find ticks, remove them as soon as possible
with a proprietary tick-removal device - do not use chemicals such as
washing-up liquid, soap, or meths as these are more likely to cause the
tick to expel bacteria into your bloodstream. Keep an eye on the
affected area and if it develops into a 'bulls-eye' pattern, seek medical
treatment quickly. Antibiotics will be needed to prevent Lyme Disease,
which is a serious long-term condition if not dealt with.

• Bracken spores are carcinogenic. It is advisable to avoid bracken-
covered areas from mid- July onwards.

Water bodies

• Consider the risks associated with water bodies and wet flushes,
especially areas of soft mud with no vegetation in fens and bogs.

• Avoid steep banks which could allow you to slip into a water body. Be
aware of becoming trapped in muddy conditions.

• If working close to a water body, it may be advisable to carry
emergency flotation equipment or 'throw line'. Carry dry clothing and
equipment to prevent hypothermia.

• Leptospirosis (Weil’s disease): most commonly associated with rats,
through bacteria in their urine, which can survive up to four weeks in
water. Humans can become contaminated via infected urine, water or
mud. Bacteria enter humans via cuts or through mucous membranes
(e.g. eyes, mouth or nose). Simple precautions include cleansing and
covering cuts with waterproof plasters. Avoid rubbing eyes, nose and
mouth and wash hands thoroughly after the survey. If flu like symptoms
develop within three to 19 days of the survey, contact a doctor and ask
for an ELISA blood test. Other animals can carry the bacteria.

Surveying at night.

Such surveys should always be carried out in communicative pairs.

Equipment should include primary and back-up torches as well as a whistle for

each surveyor. Sites should be visited in daylight and a risk assessment carried

out before night-time surveys are undertaken. Consider the following:

• It may be advisable to wear high visibility clothing

• Mark hazards with hi-viz tape, lamps or light sticks.

• Consider marking the way to the survey site.

• If using a generator, follow the manufacturer’s safety instructions.

Insurance

Some landowners require the surveyor to have insurance or indemnification.

The organiser is responsible for ensuring such insurance is in place.

The guidance given above is based on appendix II of the Hedgerow Survey

Handbook by Catherine J. Bickmore. English Nature, Peterborough.

Risk assessment

Some definitions:

• An accident is an unforeseeable event. Most incidents that occur during
surveys are not accidents.

• A hazard is an object, which is capable of causing harm to people.

• A risk occurs when somebody is exposed to a hazard that causes
serious harm, is in close proximity to a hazard or is exposed to a
hazard for an extended period of time.

Five steps to risk assessments:

1 . Identify hazards

2. Identify who is at risk from hazards identified

3. Establish how hazards can be removed or risk of injury reduced.

4. Record steps 1-3 above

5. Review the risk assessment periodically in the light of working
experience.

Each surveyor should take part in the risk assessment process. Surveys
should only go ahead if measures could be taken to reduce risks to an
acceptable level.

All incidents must be reported to the survey leader, who should record the

incident in writing.

BIO-SECURITY GUIDANCE

In the past few years, there have been a number of outbreaks of disease
amongst farm animals. We ask that field naturalists working in areas where
farm animals are present should obey the following guidelines:

• If visiting farmland by car, try to avoid parking in a farmyard where
animals are kept. Try to park on hard standing, not in muddy gateways,
likely to be used by animals.

• Avoid touching farm animals or deer. Always wash your hands if an
animal is touched.

• Clean mud from boots and/ or your car after each farm visit.

• In areas of potential high risk, wash/ spray your boots with an approved
disinfectant (see the Defra website for a list of suitable disinfectants
obtainable from an agricultural merchant).

• Seek permission from the farmer to enter premises or fields.

• We also alert you to an increase in Lyme Disease from ticks. The
number of cases has doubled in the past two years. Should you have
difficulty removing a tick or suffer from a fever after a tick, seek medical
assistance and ask for a Lyme disease test.

JAN/RPS Aug 2015

Mcdjutf&JjLsL

August 2015 Volume 140 Number 1089

From Yorkshire to China via Borneo: a biological excursion
through tropical moth ecology

The Presidential Address delivered following the Annual General Meeting at Mai ham Tarn Field
Centre , 15th November 2014.

Terry M. Whitaker

4 Crowtrees, Low Bentham, Via Lancaster, LA2 7EE.

Email: t.whitakerl@btinternet.com

In this address I hope to introduce you to some of the questions about moth species
distributions and population diversities that I have had a minor part in helping study in South-
east Asia. Some Yorkshire-born or based researchers have played a major part in researching the
associated ecology, especially Dr. Stephen Sutton, a past YNU President; more recently by the
research group of Professor Roger Kitching of Brisbane, Australia. Many of the questions have
not been fully answered yet.

The Big Questions:

1) What is the moth diversity in various biotopes?

lb) Does it depend on the biotope involved?

lc) What is the taxonomic structure of the species composition?

ld) How many moth species are there in Borneo?

2) Which areas & biotopes have the highest diversities?

3) How does moth diversity change with geographic distances?

3a) At what geographic scale?

3b) Are moths stratified vertically(ground to canopy)?

3c) How does moth diversity change temporally?

3d) How does moth diversity change on altitudinal transects?

The Naturalist 140 (2015)

3e) How does moth diversity change on latitudinal transects within forest biotopes?

3f) How do patterns of moth diversity change between continents?

Biodiversity: the importance of robust data (and big numbers)

A variety of objective measures have been created in order to measure biodiversity empirically.
The basic idea of a diversity index is to obtain a quantitative estimate of biological variability
that can be used to compare biological entities, composed of direct components, in space or in
time. Indirectly this can be used to estimate the number of species.

The term alpha diversity (a-diversity = within habitat diversity) was introduced by Whittaker
(1960, 1972) together with the terms beta diversity ((3-diversity= between habitat diversity) and
gamma diversity (y-diversity = whole landscape diversity (a larger geographical unit)).
Whittaker's idea was that y-diversity is determined by two different things, the mean species
diversity in sites or habitats at a more local scale (a-diversity) and the rate of change within
those habitats ((3-diversity). Definitions of a-diversity can also differ in what they assume
diversity to be. Often researchers use the values given by one or more diversity indices, such as
species richness, by the Shannon index or the Simpson index. Species richness is the number of
different species represented in an ecological community, landscape or region. It is simply a
count and does not take into account abundances, relative abundance distributions or the rarity
of individuals, whereas biodiversity can. However, it has been argued that it would be better to
use the effective number of species as the universal measure of species diversity. This measure
allows weighting of rare and abundant species in different ways, just as the diversity indices
collectively do, but its meaning is intuitively easier to understand. The effective number of
species is the number of equally abundant species needed to obtain the same mean
proportional species abundance as that observed in the dataset of interest (where all species
are not equally abundant) (Hill, 1973; Jost, 2007; Tuomisto, 2010). It is important to distinguish
'richness' from 'diversity'. Diversity usually implies a measure of both species number and
'equitability' (or 'evenness'). Fisher's logarithmic series model (Fisher et ol. 1943), log Series a
distribution (like the more commonly used Poisson log series) describes the relationship
between the number of species and the number of individuals of those species, and is one
measure that has been commonly adopted (Taylor et ai, 1976) which I am going to use to
introduce some of the concepts of species diversity in moths. At its simplest for a sample of a
given number of individuals the diversity will be highest in the sample containing the greater
number of groups (species). In the tropics moth species rank abundance curves from the most
diverse sites and the least diverse sites show similar inverse exponential curves but they differ
markedly in detail. The former have few abundant species and a long tail of rare ones whereas
less diverse sites have several abundant species and many fewer scarce ones. In order to
compare biodiversity of samples from different locations it was necessary to elucidate the
statistics of sampling and a measure was needed to understand species-richness expressed in a
way that was independent of samples size (Robinson & Tuck 1993a, 1996). This led to the use of
rarefaction curves (Sanders 1968).

la. What is the moth diversity in various biotopes?

The first people to estimate moth species richness in SE Asia were Barlow and Woiwod (1989,
1990) and they were the first to consider the contribution of 'micro-moths' to diversity. Their
samples proved what had been suspected, that the values of a-diversity of moths in tropical

The Naturalist 140 (2015)

forests could be extremely high (Table 1). This was confirmed by Holloway et al. (1990), who
obtained an a-diversity value of 309 for macrolepidoptera in lowland dipterocarp forest in
Sarawak. Robinson & Tuck (1993a, 1996) found that micromoths from two primary forest sites in
Brunei were particularly diverse, showing a-diversity values of 414 in pooled samples. Diversity
will also vary within an order related to the infra-ordinal taxonomic status of the sample group.
Samples restricted to micromoths, macromoths or pyralids will show lower values of a than
those counting all moths (Table 1). Sample size is important but larger samples will have greater
diversity values, especially in highly diverse places. This effect is the result of casual migration
and an increasing opportunity for rare moths from a wider area to be captured over a longer
time period. In effect, collecting is an open-ended enterprise. In practice, samples greater than
1000 individuals tend to have stabilized variances and were thought to provide useful 'snapshot'
measures of diversity.

Holloway et al. (1992) and Chey et al. (1997) used pooled samples of fewer than 900
individuals from single locations, which seemed to show that primary natural forest in Danum
Valley had a low moth diversity leading to the assertion that a-diversity at Danum appeared to
be lower than that recorded in other Bornean primary forests but similar to disturbed forest
habitats in Brumas. Also, nine equally spaced sample sites within 1.5km in the Danum Valley
Conservation area sampled by Beck et al. (2006) showed a very low a-diversity of only 14.5 ±
2.9. It was based on only 1596 geometrids (127 species) and his conclusion, that one sample in
an area of 225ha for the investigated taxon is representative, is now considered totally
unjustified. This is refuted here as Table 2 (from Willott et al. unpubl.) demonstrates, a for the
pyraloids (186) is similar to that recorded from Brunei (203) (Robinson & Tuck, 1993a) and
higher than for Peninsular Malaysia (91) (Barlow & Woiwod, 1989), although another study
based on small samples and methodological differences between the latter and the collections
from Borneo makes comparisons difficult. Similar criticism of measurements based on very
small numbers can be applied to some of the a-diversity values from West Java reported by
Sutrisino (2008).

The monumental dataset collected by Henry Barlow and analysis based on the staggering
number of 70,529 individuals of Macros and Pyraloidea plus Thyridoidea should draw some of
the above arguments to a close (Ashton et al., 2014). The authors say that there is no substitute
for long-term 'old fashioned' accumulation of data. Snapshot surveys will only provide a relative
measure of species richness.

To put the a-diversity figures in Table 1 into context: in Yorkshire (with c.500 species of
macromoths) the richest sites have an a-diversity of value <41 (Sutton & Beaumont 1989, Anon
1996). An a-diversity of 40 would represent 221 species for every 10,000 individuals sampled. In
contrast the mean UK a-diversity of Pyraloidea is 2.9 whereas at Genting it is 91 (Barlow &
Woiwod 1989). In comparison considering the Brunei case an a-diversity of 414 suggests c. 1,333
species of microlepidoptera occur near the Batu Api Forest Reserve sites (Robinson & Tuck
1993a, b; 1996).

The Naturalist 140 (2015)

Table 1. Some estimates of moth diversity in South-east Asia.

Location

Biotope

■

Trapped Types

a-diversity ±
S.E.

Study

Reference

W. Malaysia, Genting

150

h 309

Holloway 1987

W. Malaysia, Genting

650

imp

386 + 11

Barlow & Woiwod 1989

W. Malaysia, Genting

650

91 1 5

Barlow &. Woiwod 1989

(in Schulze 2000)

Sarawak

150

309

Holloway et al. 1990

Sulawesi, Dumoga-Bone NP

225

303.1 +9.2

Barlow & Woiwod 1990

Sulawesi, Dumoga-Bone NP

PI*

225

|234.5 1 9.4

Barlow & Woiwod 1990

Brunei, Kuala Belalong 1

125

355 1 46

Robinson & Tuck 1993a

Brunei, Kuala Belalong 2

125

413 1 74

Robinson &Tuck 1993a

Brunei, Kuala Belalong 1+2

125

414 1 39

Robinson & Tuck 1993b

Brunei, Kuala Belalong 1

125

193 1 35

Robinson & Tuck 1993a

Brunei, Kuala Belalong 2

125

218 + 46

Robinson & Tuck 1993a

Brunei, Kuala Belalong 1+2

125

203 1 25

Robinson & Tuck 1993b

Brunei, Kuala Belalong 1+2

125

^216132

Robinson & Tuck 1993a

Brunei, Kuala Belalong 2

125

222 1 77

Robinson & Tuck 1993a

Brunei, Kg. Kapok 1

39 1 19

Robinson &. Tuck 1993a

Brunei, Kg. Kapok 2

56 1 9

Robinson & Tuck 1993a

Brunei, Kg. Kapok (1+2)

69 + 11

Robinson & Tuck 1993a

Brunei, Kg. Kapok (1+2)

120 + 13

Robinson &. Tuck 1993b

Brunei, Kg. Kapok,

47111

Robinson & Tuck 1993a

Brunei, Kg. Kapok

105 1 13

Robinson & Tuck 1993a

Brunei, Kg. Kapok

120 1 13

Robinson & Tuck 1993a

Sabah, Crocker Range

1500

226135

Robinson &. Tuck 1993a, 1996

Sabah, Crocker Range

1500

64 + 20

Robinson & Tuck 1993a, 1996

Temengor, W. Malaysia

275

120 1 24

Robinson et al. 1995

Sabah, Danum Valley

125-150

292 1 14

Willott 1999

Sabah, Danum Valley

125-150

234 1 15

Willott 1999

Sabah, Danum Valley

150

244 1 15

Willott 1999

Sabah, Poring (Pori))

580

129113

Schulze 2000

Sabah, Poring (Por2)

630

147+13

Schulze 2000

Sabah, Poring (Por3)

600

138+20

Schulze 2000

China, Xishuangbanna

PA*

600-800

25517.15

Kitching et al. (2015)

China, Xishuangbanna

600-800

29218.32

Kitching et al. (2015)

W. Malaysia Sg. Halong

311

821.7+8

Ashton et al. (2015a in press)

W. Malaysia, Genting 1980

650

469.7+7

Ashton et al. (2015a in press)

W. Malaysia, Genting 2000

650

337.6+9.5

Ashton et al. (2015a in press)

Sabah, BRL

742.7+9

Ashton et al. (2015a in press)

KEY PI = primary lowland forest; P2 primary hill forest; P3 primary submontane forest; PC primary

lowland forest canopy; PA = primary alluvial lowland forest; PL = primary lowland forest on limestone;
L = logged over forest; S = Secondary Forest (Mature); M = Mangrove Forest; * disturbance present;
h = macro-Lepidoptera; m = micro-Lepidoptera sensu stricto; p = Pyraloidea.

The Naturalist 140 (2015)

lb. Does it depend on the biotope involved?

Beck et al. (2002) presented geometrid data along two habitat gradients ranging from primary
rainforest to cultivated land in Sabah, North Borneo. The moth diversity in the forest plantations
such as Acocio mongium, Gmelino orboreo, Paraserianthes falcotorio, Pin us coriboeo and
Eucalyptus deglupta, is unexpectedly high. Moth diversity in the E. deglupta plantation was
particularly high and comparable to that in old-growth secondary forest, possibly because this
plantation has a very diverse understorey both in terms of plants (secondary regrowth species)
and architecture. Disturbed tropical forest usually shows lower a-diversity (but higher p-
diversity) and peat swamp forests also show lower a-diversity (Holloway 1992; Chey 1994, 2000,
Chey et al. 1997; Schulze 2000; Fiedler & Schulze 2004). In some other woodland biotopes such
as mangrove forests, oil palm and agricultural landscapes, a-diversity is inherently very low,
reflecting the smaller pool of specialist moths dependent on the lower vegetational diversity
and the simpler forest architecture (Table 1, Robinson & Tuck, 1993a, b, 1996; Willott, 1999;
Beck et al., 2002). Kitching et al. (2015) found that faunas in lowland alluvial forest and
limestone forest in China (Yunnan, Xishuangbanna) had similar a and P-diversities and
comparable moth species compositions. Submontane forest sites can show quite high a-
diversities until the vegetation changes and it becomes depauperate in the higher altitude
montane forests.

In trying to estimate how many moths were restricted to primary forest (a presence - absence
criterion), Willott (1999) showed that c.55% were not sampled in primary forest and 11% were
only encountered in logged forest. The proportion changed if moths with a minimum
abundance were excluded. If this was restricted to those represented by at least ten individuals
it suggested that at least 10% were confined to primary forest while <1% were restricted to
logged-over forest. Unpublished data from the Whitaker & Kitching (2008) study did not confirm
this over a larger geographic scale. 602 of 2283 morphospecies (26%) were unique to secondary
and 537 (24%) confined to primary forest. A morphospecies is an unidentified taxon close to
species level which can be reliably distinguished by external appearance. However, as primary
forest samples are typified by a large number of rare moths represented by only singletons this
may not be a realistic measurement. Species totals in primary and post-logging secondary
forests confirm that each forest type presents similar levels of species richness (Hamer et al.
2003, Kitching et al. 2012, Willott 1999) but these are not composed of the same species.
Deaton (1993 unpubl.) showed similar results (but with limited data) when comparing primary,
secondary and heath forest (kerangas) at Barito Ulu, Kalimantan.

lc. What is the taxonomic structure of the species composition?

As reported by many authors working worldwide in perhumid tropical forests, micromoths of
the superfamily Geometroidea, Crambidae plus Pyraloidea (sensu Regier et al. 2012) and
Noctuoidea were known to form a large proportion of all tropical forest moths and the fauna is
dominated by these and other tympanate moth families (e.g. Sutrisino 2008). After recent major
taxonomic changes following DNA sequencing studies (Zahiri et al. 2011), the quadrifid
Noctuoidea (traditionally divided into the Noctuidae, Lymantridae and Arctidae (LAQ clade)) is
now partitioned into the Erebidae. This forms a large family with 21 subfamilies and several as
yet unassigned groupings which contain the lymantrids, arctiids and other parts of the LAQ clade
(especially the catocalids, originally considered with the Noctuidae). To illustrate this, Whitaker

The Naturalist 140 (2015)

& Kitching working at 10 sites in primary (virgin) tropical forest and disturbed (logged-over)
forest in Sabah (North Borneo), constructed a morphoseries from 14,013 specimens captured in
Pennsylvania-pattern actinic light traps (Kitching et al. 2005). Those that were identifiable were
allocated to 39 'families'. Converting the dataset to reflect modern taxonomy, the tympanate
families Erebidae (36.8%), Geometridae (14.1%), and the Pyraloidea (20.7% (Crambidae 14.2%
and Pyralidae 6.5%)) were found to make up the majority (72%) of the taxa caught. The other
tympanate families contributed less than 2.1% individually (Notodontidae (1.9), Noctuidae
(1.3%), Nolidae (1.2%) and Uraniidae (0.9)) as did the non-tympanate families with the
exception of the Limacodidae (7.0%) and Drepanidae (4.1%). There was no significant difference
in familial composition between primary and disturbed forest samples.

Id. How many moth species are there in Borneo?

This question has been asked of many geographic areas and for the apocryphally speciose
tropical forest it was undoubtedly a very large number. Holloway (1987), using his experience of
the region, initially suggested a total of 4,500 larger moths in Borneo. Initial estimates were
made by intuition, a variety of indirect sampling methods (which have been refined in time with
repeated sampling and improved statistical treatments such as the Chao 1 estimator (Chao,
1984)) and cumulative catalogues of species and morphotypes. As an example, Robinson & Tuck
(1993a, b) initially estimated that a-diversities of macrolepidoptera, Pyraloidea and
microlepidoptera of 310, 170 and 210 respectively, suggested c.3,750 moths occur in lowland
Bornean forest and, extrapolating from a number of taxonomic and field observations,
suggested a Bornean total for all moths of 8,628 species (3,614 macrolepidoptera and 5,014
microlepidoptera) (Robinson & Tuck, 1993a, b, 1996). Holloway (1986-2011) produced a list of
4,563 macromoths which, combined with 936 butterflies and c.6,331 microlepidoptera, gave
a minimum total of 1 1,830 lepidoptera. The conclusion to his massive revision of the taxonomy
of Bornean macromoths amended this to an estimated total of 12,684, which was revised to
12,777 using data in Ashton et al. (2014). This may still be an underestimate as Pyraloid
micromoths alone probably total in excess of 3,000 species (Sutton, Barlow & Whitaker, 2015 in
press).

Species richness estimators estimate the total number of species present in a community. The
Chao 1 index is commonly used and is based upon the number of rare classes found in a sample
(Chao, 1984): Using the Chao 1 estimator (Colwell & Coddington, 1994), an increment to the
numbers observed is derived from the square of the singleton number divided by twice the
doubleton number in a series. Applying the Chao 1 estimator to the Whitaker & Kitching Sabah
dataset of 14,013 specimens of 2,283 morphospecies of macros and micros (1,681 (6,831
individuals) in primary forest and 1,746 (7,182 individuals) in logged-over forest with 1,144
morphospecies in common) suggests at least 4,285 morphospecies in primary and disturbed
forest across northeast Borneo (2,430 in primary forest and 1,962 in logged-over forest). Using
the same methodology, Ashton et al. (2014) estimate 5,422 macros and micros at Genting
(disturbed forest) and 5,097 macros and micros in primary forest at Borneo Rainforest Lodge
(Sabah).

The Naturalist 140 (2015)

3. How does diversity change with geographic distances?
a) Measuring Beta Diversity; Does 'McDonaldisation' Exist?

At a landscape scale there is often a continuum of habitat change and it is difficult to determine
the limits of a particular biotope. Beck & Chey (2003) attempted to address this and other
problems of temporal change and altitudinal change using published datasets of geometrid
assemblages across 700km in Borneo. They concluded that the variation between the moth
ensembles was mainly explained by factors such as elevation, habitat disturbance and temporal
factors such as weather and habitat changes over three decades. The validity of their conclusion
that residual variation still contains a pattern was tentatively explained by geographical distance,
particularly <20km, but methodological differences made their comparison unreliable.

Prof. Roger L. Kitching set out to start to answer the question of (3-diversity and its role in y-
diversity. This included the statistically robust study on two forest biotopes in Sabah introduced
above (Whitaker, 2007, 2008 unpubl.; Whitaker & Kitching, 2008; Kitching et oi, 2013). The
results showed that there were very similar patterns in moth species richness and levels of
diversity in both the primary and logged-over forest. However, a strong relationship was
observed between moth assemblages in primary forest, decreasing in similarity with increasing
geographic distances over 80km but no such pattern was found in post-logging forest (Figure 1).
We speculated that the strong spatial heterogeneity in moth assemblages in the primary forest
landscapes may be absent from logged-over forest, through the removal of canopy niche space
(Kitching et oi, 2013). The small scale spatial heterogeneity in primary forest may be the reason
why some a-diversity measurements in primary forest show relatively low values. However,
Novotny et oi (2007) found a low rate of change in species composition (p-diversity) of many
insect groups across lowland primary forest in Papua New Guinea.

Sharp (2014) applied Jost's (2007) 'true' diversity measures with a functionally diverse group
(Coleoptera) to estimate ecosystem-level changes in a-diversity, p-diversity and y-diversity
associated with disturbance. Calculating diversity, y-diversity and P-diversity decreased with
disturbance based on taxa richness but not on proportions of taxa counts. In all cases p-diversity
was influenced by interactions between distance and disturbance, and provided a more
descriptive insight into changing community structure than either other component. Declines in
P-diversity are likely to result from reduced habitat heterogeneity and a switch from niche
differentiation to habitat filtering (Kitching et oi, 2013)

3b) Are Moths Stratified (Ground to Canopy)?

The concept of the tropical forest canopy as a distinct biotope that determines arthropod
vertical distribution due to its abiotic factors, forest physiognomy and tree architecture,
resource availability and arthropod behaviour, has been proposed for many years. Many workers
(e.g. Stork, 1988) postulated that a significant proportion of the biodiversity of primary tropical
forests was held in the canopy. The canopies of tropical forests were once thought to contain
the most rich and diverse assemblages of insects, and moth abundance was shown to be higher
in the forest canopy in Borneo, Papua New Guinea and Panama (Sutton et oi, 1983; Kato et oi,
1995), although whether this was a real difference cannot be established as identification was
not to species so there can be no estimate of species richness or correction for sample size.
However, there was growing evidence that the canopy was no more diverse than the

The Naturalist 140 (2015)

understorey (Hammond, 1990; Wolda et al., 1998; Willott, 1999), though the distributions of
insects in woodland appear to be strongly vertically 'compartmentalised' with more or less
isolated assemblages of species occupying closely adjacent habitat components within a forest
(Basset etol., 1992a; Amedegnato, 2003; Wardhaugh et. al., 2012; Ashton et al., 2015b in prep).
Intachat & Holloway (2000) found that there was no consistently significant difference in species
richness, abundance or diversity (as measured by a) between the three levels 1, 15 & 30m at
Brumas, although values tended to be lower at the highest level. The diversity for the canopy
(high) level was significantly lower when all samples were pooled. Here plant diversity is lower
and the forest architecture is less complex. Individual flight height preferences varied according
to the taxa studied.

Figure 1. Relationships between geographical distance and (A) Chao-Sprensen & (B) Sprensen
similarity values using moth assemblages collected from primary (triangle) and post-logging
secondary (circle) forest. Trend lines are drawn for primary forest only.

From Kitching et al., 2012

These comparisons of canopy and understorey faunas in tropical forests are of more than purely
theoretical interest. Several papers have examined the extent to which lepidopteran diversity is
maintained in disturbed forests or plantations but without samples from the canopy of the
primary forest 'control' site (Holloway et al., 1992; Hill et al., 1995; Chey et al., 1997; Holloway,
1998; Hamer et al., 2003). If a large number of canopy specialists are missed in the process then
the species richness of the primary forest may be severely underestimated or the taxonomic
composition misjudged. Furthermore, there is some evidence that canopy insects may fly closer
to the ground in disturbed or plantation forest where the canopy is lower and more open (Davis
& Sutton, 1998; Willott, 1999). If these species are detected in ground-based sampling in the
disturbed habitat but not the primary, then the estimate of species richness in the former will be
inflated relative to that of the latter. Willott (1999), confirmed by Beck et al. (2002), showed that
there was a substantial turnover of larger moths between the canopy and ground faunas with a
combined a-diversity of 367 ± 15, larger than that of its independent components (Table 1).

The same is true of the Pyraloidea: Willott et al. (unpubl. ms.) collected a total of 5,322
individual moths comprising 1,207 species on just four nights from April to July 1995. Light traps
were sited at Danum Valley Field Station at ground level and at 40m in primary forest canopy.
Randomised species accumulation curves for both the macrolepidoptera and Pyraloidea were

The Naturalist 140 (2015)

generated by randomly sampling without replacement from each dataset to compensate for
more species and individuals being collected from the understorey, taking the mean of 1000
iterations for each sample size. Following the same procedure, the mean number of species
expected in a sample of 1000 individuals (Siooo) was generated to provide an estimate of species
richness independent of sample size. The data are presented in Table 2.

Table 2. Numbers of species (S), individuals (n), diversity (alpha of the log-series) and rarefied
species richness (Siooo) (both mean ± 95% Cl). From Willott et al. (unpubl.)

Alpha

Siooo

Macrolepidoptera

Canopy

414

1133

235 ±22
242 ± 19

382 ±7

Understorey

500

1673

375 ± 12

Total

709

2806

305 ±18

410 ±14

Pyraloidea

Canopy

307

1084

143 ± 14 295 ± 6

Understorey

383

1432

171 ±14

325 ±11

Total

498

2516

186 + 12

| 328 + 13

Combined

Canopy

721

2217

371 ±25

447 ± 18

Understorey

883

3105

412 ±23

471±17

Total

1.207

5322

487 ± 21

487 ±20

A randomised species accumulation curve for the macrolepidoptera was steeper than that for
the Pyraloidea, suggesting that a greater proportion of the species pool of Pyraloidea had been
sampled. The most abundant macromoth was Amato prepuncta Holloway (Erebidae: Arctiinae,
Syntomini) with a total of 151 individuals and the most abundant pyraloid was Pagyda salvalis
Swinhoe (Pyralidae: Spilomelinae) with 102 individuals. In this limited unreplicated study the
significant species turnover between the understorey and canopy confirmed that sampling in
the canopy is a pre-requisite for an accurate estimate of the diversity and faunal composition of
a site. While comprising fewer species, the number of individuals of Pyraloidea approached that
of the macromoths in total but the Pyraloidea in Brunei comprised approximately half the total
of species and individuals of the micromoths as a whole (Robinson & Tuck, 1993a). This suggests
that up to two thirds of the abundance of flying moths may be microlepidopterans. A large
proportion of lepidoptera are layer-specialists, with mixing between ground and canopy being
the exception rather than the rule (Schulze et al., 2001; Schulze & Fiedler, 2003; Basset et al.,
2003; Brehm, 2007). In addition, most are narrowly oligophagous, often being restricted to a
single genus or genus-group within a particular plant family while some feed on leaf litter and
detritus (Bassett, 1992; Novotny et al., 2002, 2003; Dyer et al., 2007). However, Willott et al.
(unpubl.) tested groups with >20 individuals in total against the null hypothesis of equal
abundance between canopy and understorey in a numerically limited study and showed that the
Herminiinae, thought of as leaf litter and detritus feeders, did not conform to that expectation,
being found mainly in canopy samples. Possibly this is related to the high biomass of canopy
detritus. The families found predominately in the understory were Nolidae: Chloephorinae
(Sarrothripini & Chloephorini), Euteliidae (Stictopterinae) and (Pyralidae) Epipaschiinae.

The Naturalist 140 (2015)

Beck et al. (2002) showed that canopy assemblages of geometrids were not very similar to each
other and more closely resembled their corresponding understorey samples, a feature noted by
Ashton (pers. com., Ashton et al., 2015b in prep.) in Yunnan, China, where different types of
forest all showed distinct canopy components to their biodiversity at all sites. Differentiation
between samples in the understorey and in the canopy is still a valid concept but may not be
universal. Stratification within the canopy itself (which can be >20m thick in Borneo) still
remains a mystery and may not exist.

3c) How does moth diversity change temporally?

Wolda (1983) considered that seasonal variation of tropical animals was the rule, even where
seasonal weather changes are minimal, but definitive studies involving large datasets were
lacking. Studies in Sulawesi and Peninsular Malaysia showed slight evidence of temporal
heterogeneity (Barlow & Woiwod 1989, 1990). Generally in undisturbed perhumid forests there
often seems little difference between samples in successive years (Kitching et al., 2013) but
there is usually a significant increase in diversity when the data sets are added (Barlow &
Woiwod, 1989; Kitching et al., 2013). A principal component analysis on monthly catches in
traps at various levels in primary lowland dipterocarp forest in Sarawak detected non-random
seasonal trends of insect abundance (Kato et al., 1995). These could have been due to
vegetational succession, phenology or slight seasonal changes during short dry spells (Robinson
& Tuck, 1993a, b). Fiedler and Schulze (2004) showed higher temporal variation at primary forest
and agricultural sites than in sites with intermediate disturbance. This could be traced down to
the species level. That diversity changes over longer periods has been demonstrated on Mt.
Kinabalu (Sabah; Borneo) by Chen (2011). This could probably be ascribed to climatological
changes (Chen et al., 2009, 2011). A similar long-term change was demonstrated at Genting,
West Malaysia, by Ashton et al. (2015a in press) using the data sets from 1980-2000 and 2000-
2013. This distinct temporal component was possibly associated with vegetational changes.

3d) How does moth diversity change on altitudinal transects?

Depending on the geographic location of the tropical forest transect, the position of highest
species richness can vary and this is not uniform at smaller taxonomic scales. In many cases this
is a result of vegetational changes along the transects - see Axmacher & Fiedler (2008) on Mt
Kilimanjaro, a-diversity of Geometridae, Pyraloidea and Arctiidae significantly declined in
Ecuador between 1,040m and 2,677m a.s.l., but was overall remarkably high: 250.1, 185.1, and
96.1 respectively (combined a-diversity of 531.6) and a was 120-185 even at the highest sites.
The Geometroidea maintained their diversity to the highest levels whereas most of the
Pyraloidea showed the expected decline of ectothermic herbivores from a 100 at 1,040m to <35
above 2,000m while the Arctiidae showed a much less steep decline, similar to the Crambidae as
compared with the Pyraustinae (Brehm et al., 2003; Fiedler et al., 2008).

Maximum diversity for macromoths in Borneo was noted around 1,000m a.s.l. (Holloway, 1987;
Holloway et al., 1990; Chey, 2000). These differences may relate to vegetational richness of food
plants (Holloway & Nielsen, 1999; Beck et al., 2002) or to moth clade tolerance of lower
temperature regimes (Fiedler et al., 2008). On Mt Kinabalu maximum vegetational richness is
around 1,000-1, 299m (Ashton, 2003).

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The Queensland-Chinese Academy of Sciences project (Q-CAS) was born to test these ecological
concepts on an ever greater scale across the length of Yunnan Province from the south-western
lowland tropical forests at Mengla through mid-altitude subtropical and deciduous forests of
Aila Shan to the alpine coniferous forests near Lijiang on the borders of Tibet. It involved
replicated sampling of canopy and ground. The enormous dataset of botanical and
entomological samples from replicated altitudinal transects between 800m and 3,800m is still
being worked on (Ashton et al., 2015b in prep.).

3e) How does moth diversity change on latitudinal transects within forest biotopes?

There are two current models of community assembly based on one hand on ideas of neutral
species replacement and on the other of resource-based niche partitioning. Niche dimensions
can be determined by two non-overlapping sets of physico-chemical parameters (or their
surrogates: altitude, latitude or substrate) and the physiologically defined envelope in which an
organism can exist. Kitching (2013) examined the geographic scale of when wholly stochastic
processes give way to deterministic processes (the stochastic-deterministic switch line (SDL))
and concluded that there is a point for each class of ecological community at which an assembly
cannot be distinguished from a random combination of species. This model explains the
distance-decay change observed in primary forest moth communities over 80km first elucidated
in Sabah and its absence in logged-over forest (Kitching et al., 2013). Kitching (2013) considered
that the associated changes in moth assemblages are driven by locally changing availability of
larval food plants. The underlying vegetation changes can usually be explained by neutral ideas
over a scale of several hundred kilometres as modified to include dispersal distances. The bio-
geography of lowland forest moths in Malaysia over several hundred kilometres (Ashton et al.,
2015a in press) can probably be explained by a combination of niche suitability and random
(stochastic) dispersal plus colonisation and speciation events.

The Q-CAS Project with its large collections of several invertebrate groups will undoubtedly add
to our knowledge of these processes. Combined moth collections (macromoths and pyralids,
determined as morphospecies) from each of the three Q-CAS altitudinal transects showed the
following approximate numbers; 2,500 from Mengla (800-1400m); 1,600 from Ailoa Shan (2000-
2700m) and 750 from Lijiang (Mt Satseto) (3200-3800m). At Mengla it is probable that
Lepidopteran diversity is extremely high, higher than any measured in the Malaysian Peninsular
and probably approaching those of Borneo (Ashton et al., 2015a in press).

3f) How do patterns of moth diversity change between continents?

Ashton et al. (2015a in press), working on 64 data sets from China (Yunnan), Panama, Vietnam,
Borneo and Papua New Guinea (containing 175,768 moths) showed consistent differences
between canopy and ground assemblages at almost all rainforest locations and across altitudinal
and latitudinal gradients. Vertical beta diversity increases with increasing elevation in each
of the northern hemisphere transects that have been undertaken. This is a reverse (and
unexpected) pattern compared to that in the southern hemisphere. This is possibly related to
structural differences in vegetation patterns.

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Acknowledgements

I would like to acknowledge the contribution of many fellow workers in the research field and in

the Queensland-Chinese Academy of Sciences (Q-CAS) Project. I am especially grateful to Prof

Roger Kitching, Dr Louise Ashton, Dr A. Nakamura and Dr J. Willott.

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Nine years of change in the flora of Ellerburn Bank, a limestone
grassland in the North York Moors

Peter J. Mayhew, Susan E. Firth, Paul R. Waites and the Second-Year Ecology Field Course.

Department of Biology, University of York, Heslington, York YO10 5DD.

Email: peter.mayhew@york.ac.uk

Introduction

Limestone grassland is one of the most biodiverse habitats on Earth at a small scale (Wilson et
a!., 2012). Floristic richness within Europe can reach 80 species per m2 (Butaye et al., 2005) and
many plants are found in no other habitat. Limestone grassland also supports important
invertebrate communities as well as specialist birds (UK Steering Group, 1998). However, it is
also one of Europe's most threatened habitats (WallisDeVries et al., 2002). In 1998 there was a
maximum of 41,000 hectares of lowland calcareous grassland remaining in the UK (UK Steering
Group, loc. cit.). Loss of habitat continues from forestry, conversion of pastureland to crops and
land abandonment (WallisDeVries et al., loc. cit.), leading to reductions in the area and increases

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in the isolation of habitat patches (Fisher & Stocklin, 1997). Quality reductions come from the
abandonment of traditional agricultural practices leading to land-use intensification such as
increased fertilization, herbicides, reseeding and frequent or early mowing (WallisDeVries et a!.,
loc. cit.). They may also result from overgrazing, undergrazing leading to the encroachment of
scrub, Bracken Pteridium oquilinum and coarse grasses (Bobbink & Willems, 1987), and
atmospheric nitrogen deposition leading to loss of richness (van den Berg et al. , 2011). The
latter threats mean that their quality may decline even if sites are enclosed in protected areas,
due to pervasive forces beyond the control of reserve managers (van den Berg et ol., loc. cit.) or
as a consequence of suboptimal management. Indeed, JNCC declared that only 29% of Sites of
Special Scientific Interest (SSSI) and 27% of Special Areas of Conservation (SAC) on lowland
calcareous grassland in the UK were in favourable condition in 2006, below the average for
habitats in general (Williams, 2006). Optimal management is difficult to achieve because the
effects of different practices can vary depending on local conditions (Klimek et ol., 2007) and the
taxonomic group under consideration (WallisDeVries et ol., loc. cit.).

Ellerburn Bank is a 2.91ha grassland site sloping south-east on oolitic limestone on the southern
edge of the North York Moors near Pickering (SE853849: VC62). Despite its modest size, it is one
of the most extensive areas of unimproved limestone grassland remaining in the North York
Moors (Sykes, 1993). It was notified as a SSSI in 1983 and has been managed as a nature reserve
by the Yorkshire Wildlife Trust (YWT) since 1966, having been informally managed for nature
conservation perhaps for the previous decade (Yorkshire Wildlife Trust, 2012). The flora and
fauna of the site is exceptional for the region, with over 150 species of plant recorded (Sykes,
loc. cit.) including large displays of Cowslip Primula veris in spring, orchids in early summer (Plate
1, centre pages) and Felwort Gentianella omorello in late summer (Leadley & Richards, 2012).
Flowering plants of regional note include Dropwort Filipendulo vulgaris, Woolly Thistle Cirsium
eriophorum, Saw-wort Serrotulo tinctorio, Fly Orchid Ophrys insectifero and Greater Butterfly
Orchid Plotonthero chlorontho. The site is noted for its extensive Lepidoptera fauna (Frost,
2005), including butterflies of regional note such as Dark Green Fritillary Argynnis oglojo and
Dingy Skipper Erynnis toges, and the site also supports a population of Glow-worm Lompyris
noctiluco. Management of the reserve currently consists of low-intensity winter grazing by
Hebridean sheep (Leadley & Richards, loc. cit.) and rotational scrub clearance (often burnt on
site). The site is listed as in 100% favourable condition by Natural England (2014) and light
winter grazing is its official management advice (English Nature, 2004). The upper, north-
western margin borders an agricultural field and consists of a Bronze Age earthwork (a double
ditch and bank) partially covered with Hawthorn Crataegus monogyna and Blackthorn Prunus
spinosa scrub, whilst the lower, south-eastern side bordered by forestry, consists of a patchwork
of taller grass and Gorse Ulex europaeus scrub (Fig. 1, Plate lc, centre pages).

From 1999 to 2011, with the exception of 2001 due to the Foot-and-Mouth disease epidemic,
the bank and surrounding areas of Dalby Forest were visited in the first week of July as part of
the second year Ecology Field Course run by the Department of Biology at the University of York.
In the early years of this field course it was noticed that one of the taller grasses, False Brome
Brachypodium sylvaticum, common around the scrubby fringes of the reserve, also occupied
large visible patches across the central and south-eastern parts of the pasture where
colonization of young woody scrub plants was noticeable (see Plates 1(b), 1(c) and 1(d), centre
pages). It was decided to attempt to monitor the spread of False Brome and woody scrub as well

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as their potential effects on the other flora from year to year. 24 permanent lm2 quadrats (Fig 1)
were sited throughout an area where small woody scrub plants and False Brome were
noticeable in 2003, from which time they were systematically surveyed for flowering plants. The
quadrats were re-surveyed in five of the nine years subsequent to that (2004, 2005, 2008, 2009,
2011). Changes to the structure of the degree programme at York in 2012 necessitated running
the field course earlier in the year, in May and then June. Although the quadrats were surveyed
in 2012 and 2013, the different survey dates make the data less comparable with those from
previous years because of the different apparency of above-ground parts of the plants used for
identification. It therefore seems timely to summarize here some of the findings from the initial
years of survey work.

During or just prior to the years of study reported here, the following management on the
reserve was carried out (Yorkshire Wildlife Trust, loc. cit.): in February 1999, a strip of Gorse
adjacent to the south-eastern boundary of the reserve, extending south-west from quadrats 5
and 6 (Fig. 1), was removed by British Trust for Conservation Volunteers. In November 1999, an
area of Gorse on the south-eastern boundary of the Reserve, close to quadrats 18 and 22, was
probably removed by contractors. Other small areas of scrub on the earthwork and Gorse
further south from the study quadrats were removed in 1999. In October 2001, because of
absence of grazing due to the Foot-and-Mouth epidemic, a strip of long grass along the north-
eastern and south-eastern boundaries of the reserve was mown. This may have encroached
over quadrats 5, 9, 10, 13, 18 and 22. In 2003 an area of Hawthorn saplings immediately to the
north of quadrats 15, 20, 23, and 24 was removed by strimming. In January 2007 a small area of
scrub on the earthwork was cleared by a National Park volunteer group. In October 2007 YWT
staff cut and treated an area of Gorse to the north-east of the study quadrat area but also in the
vicinity of quadrats 17, 18, 21-24. In January/February 2008 the Hobs volunteer group also cut
and burnt an extensive area of scrub along the earthwork and Gorse close to quadrats 5, 6, 9
and 10. Bracken was cleared along the north-eastern reserve boundary in 1998 and 2008.

From 1998 to 2004 a creep grazing regime was used, which saw 21 sheep being grazed
progressively on three enclosed paddocks on the centre of the pasture, from south-west to
north-east, the last of which covered the area of the study quadrats, for around 18 days on each
during the winter period. An extra paddock was grazed in 1999 extending grazing north-east. In
2003 grazing pressure was increased from 21 sheep to 40 sheep which grazed the four
successive pastures, still in a creep grazing routine. From 2005-2010 a continuous winter grazing
regime was used. Approximately 30 Hebridean sheep were located on the site between October
and March. Overall then, annual grazing was the main management activity immediately
affecting the quadrated area, though some scrub removal took place immediately adjacent to
the quadrated area, and over the north-eastern part of it in October 2007.

In this article we first summarize the community of flowering plants found in the permanent
quadrats. We then test four hypotheses of vegetation change related to the efficacy of
management of calcareous grassland sites which motivated the work: that False Brome is
increasing in frequency in the sampled area; that woody scrub plants are increasing in frequency
in the sampled area; that richness and alpha diversity are decreasing over time; and that the
Ellenberg indicator nitrogen scores (Hill et al., 1999) are increasing over time.

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Methods

Twenty-four permanent lm2 quadrats were sited across an area of the central and south-eastern
parts of the pasture measuring 120m x 40m using a stratified random design (Fig 1, Appendix 1).
The total area was divided into twelve 20m x 20m units and two lm2 quadrats were randomly
sited in each. The corners of each lm2 quadrat were marked by six inch steel nails hammered
into the ground. In subsequent years the quadrats were relocated by metal detector, aided in
later years by the use of a hand-held GPS receiver.

Figure 1: Map of the northern part of Ellerburn Bank showing the location of the survey

quadrats and local surroundings.

Plants were recorded using a semi-quantitative method. A gridded quadrat consisting of 25
divisions of 20cm x 20cm was placed over each lm2 location. Presence/absence of each plant
species was recorded in each of these sub-units of the quadrat and the number of these
subunits totalled to give an occupancy score between 0 and 25, representing how widely that
plant occurred in each quadrat. This method allowed year-to-year consistency and is relatively
rapid, given that surveying had to be completed in 2-3 days each year and by different students
in each year. Leaf shape was generally sufficient to allow accurate identification for the herbs
but presence of a grass was only recorded if a flowering stem was present, allowing accurate
identification. The exceptions were Cock's-foot Dactylis glomerota, whose fleshy leaves are
distinctive, and False Brome, whose wide tough and yellow leaves are also distinctive. One grass,
Creeping Bent Agrostis stolonifero, was also initially identified by leaf alone but subsequent
observations cast doubt on the efficacy of this identification and only flowering stems were used
in subsequent years. However, because of this inconsistency, this plant is eliminated from the
analyses below requiring occupancy estimates. Mosses were not recorded.

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Two teams of two students conducted the surveying each year. This arrangement facilitated
speedy recording and allowed students to consult each other in case of doubt. Each team was
armed with a flower guide (Fitter et ol., 1996) and a grass guide (Fitter et ol., 1984), along with
the species list from previous years' surveying. In addition, each team received close tuition on
plant identification during their first quadrats from the first author, who revisited them at
approximately hourly intervals during surveying to handle identification queries. All students
had previously participated in a class quadrating practical on the bank as part of the field course,
giving them some experience with plant identification. Despite this effort to minimize
identification errors, some students found it difficult to distinguish some plants by leaf shape,
particularly Common Knapweed Centourea nigro and Field Scabious Knautia orvensis, which can
have quite similar leaves to the inexperienced eye. Year-to-year fluctuation in occupancy may,
therefore, to some extent reflect year-to-year variability in identification error. However,
consistent temporal trends are still likely to represent real changes in the plant community.

The community composition of all plants within quadrats was explored graphically using Non-
metric Multidimensional Scaling (NMDS), implemented using the metaMDS function in the
vegan package in R (R Core Team, 2014). NMDS is an ordination analysis in which differences in
community composition are summarized in a small number of dimensions (normally two) for
ease of visualization. Species and quadrats close together in a plot of the NMDS axes show
closer associations in occupancy across quadrats. Occupancy data were logi0(x+l) transformed
prior to analysis.

To identify if woody scrub plants and False Brome were associated with particular plant
communities, the herb and grass occupancies within quadrats were subjected to Detrended
Correspondence Analysis (DCA), another ordination analysis, using the decorana function in the
vegan package in R. The analysed data omitted False Brome and woody scrub plants and were
logi0(x+l) transformed prior to analysis, with rare species downweighted. The extracted axis
scores for each quadrat were correlated against the mean occupancy of Hawthorn and False
Brome within quadrats, as a statistical test of association between those species and the
community of other plants.

Temporal changes in the occupancy of False Brome, of woody scrub plants, richness and alpha
diversity and of the occupancy-weighted Ellenberg indicator nitrogen scores (Hill et ol., loc. cit.)
were analysed by linear mixed effect models with repeated measures, with quadrat coded as a
fixed factor repeat-measured across year, and with year as a covariate. Analysis was conducted
in SPSS v.21. Alpha diversity was scored using Simpson's index on the occupancy data, which is
recommended for small sample sizes (Magurran, 2004), using the inverse index 1/D, where
larger values indicate a more even community, in which species have more similar occupancies
to each other; Ellenberg nitrogen scores for flora scale from 1 (extremely infertile) to 9
(extremely fertile) (Hill et ol. loc. cit.).

Results

The plant community

A total of 62 species (Appendix 2) was recorded over the six survey years, comprising 42 forbs
(of which 6 were legumes), 12 grasses, 1 sedge and 7 woody scrub plants. A rank occupancy
chart (omitting Creeping Bent) (Fig 2) shows that the most common twelve plants account for

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79% of the average occupancy and that other plants have very low occupancy. These twelve
were, in decreasing rank occupancy order, Common Bird's-foot-trefoil Lotus corniculotus,
Glaucous Sedge Carex flocco, Salad Burnet Sanguisorbo minor , Field Scabious, Rough Hawkbit
Leontodon hispidus, Quaking Grass Brizo medio, Cock's-foot, Common Knapweed, Upright
Brome Bromopsis erecto, Lady's Bedstraw Go Hum verum, Red Clover Trifolium pratense, False
Brome, Fairy flax Linum cothorticum, Ribwort Plantain Plontogo lonceolota and Yellow Oat-grass
Trisetum flovescens. Several of these are central to the entomological interest of the site, with
Common Bird's-foot-trefoil and Red Clover supporting Common Blue Butterfly Polyommotus
icorus, Six-spotted Burnet Zygoeno filipenduloe, Narrow-bordered Five-spotted Burnet Z.
loniceroe, Burnet Companion Euclidio glyphico and Dingy Skipper. Cock's-foot and False Brome
are favoured foodplants of many grass-feeding butterflies, including Small Skipper Thymelicus
sylvestris. Other recorded plants included Hairy Violet Viola hirto, which supports Dark Green
Fritillary, and Common Rock-rose Helionthemum nummulorium , which supports Brown Argus
Aricio agestis, whilst in the recent past Cowslip has held populations of Duke of Burgundy
Homeoris lucino and Milkwort Polygolo vulgaris supported Small Purple-barred Phytometro
viridorio (Sutton & Beaumont, 1989). The woody scrub plants were, in order of decreasing rank
occupancy: Hawthorn, Dog Rose Roso conino, Gorse, brambles Rubus fruticosus agg,
Pedunculate Oak Quercus robur, Blackthorn and Scots Pine Pinus sylvestris. Other plants of note
for their vivid floral displays include Common Spotted Orchid Doctylorrhizo fuchsii (see Plate I
(a), centre pages) and Felwort; and the regionally scarce Dropwort, Fly Orchid (see front cover)
and Woolly Thistle.

25 -i

Ra nk ord e r occu paney

Figure 2: Rank occupancy plot of the plant species recorded. Precise occupancy values, with

standard errors, are given in Appendix 2.

An ordination of quadrats using NMDS shows close associations between the twelve most
ubiquitous plants, as expected (Fig 3b): those with low scores on the first axis (Fig 3a) include
Sweet Vernal Grass Anthoxonthum odorotum (code F), Yorkshire Fog Holcus lonotus (code b),
Gorse (code Gg) and Wild Strawberry Frogorio vesco (code W), along with three occasionals:
Chickweed Stellorio medio (code z), Woolly Thistle (code 0) and Bearded Couch Grass Elymus
coninus (code S), whilst on the opposite end of the first axis lie Rockrose (code Z), Meadow

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101

Buttercup Rononculus acris (code u) and Blackthorn (code s), which tended not to be associated
with the former group. On the second axis, the lowest scores come from the same three
occasionals plus Hogweed Herocleum sphondilium (code a) and Dog Rose (code v). At the
opposite end of this axis are Sweet Vernal Grass (code F), Bramble (code w), and Sweet Violet
Viola odorata (Jj). The scrub plants (e.g. Pedunculate Oak, code t), with the exception of
Hawthorn (code P), tend to occur on the fringes of the core set. False Brome (code H) has a
slightly negative score on both axes and is most closely associated with Hogweed (code a),
eyebrights Euphrasia officinalis s.l. (code T) and Meadow Vetchling Lathyrus pratensis (code e)
within the core community. Plants not associated with False Brome include Wild Thyme Thymus
polytrichus, Harebell Campanula rotundifolia, Common Rock-rose and Felwort. Common
Spotted Orchid (code R) is associated with Goatsbeard Tragopogon pratensis (code Cc), Upright
Brome Bromopsis erecta (code J), and Quaking grass (code I).

A plot of the first two DCA axes (scrub plants and False Brome not included) (Figure 4) shows
similar sets of species associations. The mean occupancy of False Brome within a quadrat was
significantly positively correlated with the DCA1 score for that quadrat (rs = 0.44, n = 24, P =
0.03) but not any of the other DCA axis scores. The mean occupancy of Hawthorn within a
quadrat was significantly positively correlated with the DCA2 score for that quadrat (rs= 0.47, n =
24, P = 0.02) but not with any of the other DCA axis scores.

Figure 3. Non-metric Multidimensional Scaling (NMDS) plots of the plant communities sampled.

Figure 3a. Species (letter codes identified in Appendix 2) are filled circles and quadrats are
open circles. The central box is the area plotted below.

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NMDS1

Figure 3b. The centre of the plot denoted by the box in Fig. 3a at a finer axis scale, with
quadrats also identified by number used in Fig.l and Appendix 1, and species identified by
letter codes in Appendix 2.

Figure 4: Detrended Correspondence Analysis plot of the plant communities sampled, showing
axes 1 and 2.

DCA1

Figure 4a. Species letter codes identified in Appendix 2 are filled circles and quadrats are
open circles. The central box is the area plotted below.

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103

Figure 4b. The centre of the plot denoted by the box in Fig. 4a at a finer axis scale, with
quadrats also identified by number used in Fig. 1 and Appendix 1, and species identified by
letter codes in Appendix 2.

Changes through time

The lm2 quadrats varied in mean plant species richness from 16 to 23, with the lowest value in
any year being 14 and highest 28, a two-fold difference (Appendix 3). A linear mixed effects
model showed that richness per quadrat significantly increased slightly over time at a rate of
about 0.2 species per year (F = 10.2, df = 1,36, P = 0.003) and differed amongst quadrats (F =
5.21, df = 23, 36, P < 0.001). Richness was not correlated with the occupancy of False Brome (rs =
-0.202, n - 24, P = 0.345) nor with the commonest scrub plant: Hawthorn (rs= 0.387, n = 24, P =
0.062).

Simpson's diversity (1/D), calculated on occupancy within lm2 quadrats, varied from an average
of 9.9 to 15.0 with the highest value in any year being 18.1 and lowest value 6.5 (Appendix 3), a
nearly three-fold difference. A linear mixed effects model showed that diversity per quadrat
increased slightly but significantly over time at a rate of about 0.2 units per year (F =22.68, df =
1,33, P < 0.001) and differed amongst quadrats (F = 6.31, df = 23, 11, P = 0.002). Simpson's index
was not significantly correlated with either False Brome occupancy (rs= -0.053, n = 24, P = 0.806)
or the occupancy of the commonest scrub species (Hawthorn) (rs = 0.173, n = 24, P = 0.418).

Weighted average Ellenberg nitrogen scores per quadrat varied from 3.1 to 3.6 across quadrats,
with the highest annual figure being 3.9 and lowest 2.9 (Appendix 3). A linear mixed effects
model showed a small but significant decline (0.01 units per year) in average nitrogen scores
over time ( F - 4.22, df = 1, 54, P = 0.045) and a significant difference between quadrats (F = 9.27,
df = 23, 85, p< 0.001).

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Quadrats varied considerably in their occupancy by False Brome, from zero in all years in one
quadrat to a mean of over 21 in another, with 25 reached in at least one year by four quadrats
(Appendix 3). A linear mixed effects model shows no significant effect of year on False Brome
occupancy (F = 0.341, df = 1, 68, P = 0.561), though occupancy differed between quadrats (F =
27.5, df = 23, 92, P < 0.001). This suggests that the quantity of False Brome shows no overall
trend from year to year, though it differs from place to place.

Quadrats varied in their woody scrub species occupancy from zero in all years in two quadrats to
a highest average of 11.5, with the highest value reached in a single year being 20 (Appendix 3).
Occupancies were square root transformed prior to analysis to normalize the variance. There
was no significant effect of year on scrub species occupancy (F = 2.803, df = 1, 64, P = 0.099) but
occupancy differed between quadrats (F = 12.1, df = 23, 110, P < 0.001). This suggests that the
quantity of scrub shows no overall trend from year to year, though it differs from place to place.

Discussion

The data presented here do not suggest that False Brome or woody scrub plants have noticeably
increased across the sampled area of Ellerburn Bank over the period investigated, but both
groups of plants are well represented in the quadrats. The spread of woody scrub through
succession to a woodland climax (Tansley, 1922) is a well-known reason for the loss, and
reduction in quality, of calcareous grassland in Europe and requires careful management
(Butuye et al., loc. cit.). At Ellerburn Bank, two interventions are taken to reduce scrub invasion:
winter grazing by sheep (Leadley & Richards, loc. cit.; YWT, loc. cit.) and less frequent cutting by
hand. In addition, grazing by wild vertebrates such as deer and Rabbits Oryctolagus cuniculus
occurs. The optimal level of scrub removal is hard to gauge. At Ellerburn Bank, the central area
of pasture, including the area covered by the quadrats, is intended to be maintained as
calcareous grassland, leaving scrub to the fringes of the reserve. Hence, it would probably be
preferred if there were no scrub at all over the quadrated area. The fact that scrub maintains a
noticeable and constant presence indicates that the current level of grazing should not be
lowered in future, otherwise one could expect scrub to encroach more rapidly over the pasture,
necessitating further targeted cutting activity in order to maintain the quantity of grassland.
Higher levels of grazing may help reduce the problem of scrub encroachment from the fringes,
but may harm flora less tolerant of grazing or trampling. The scrub around the fringes is
tolerated as it provides wind shelter and a mosaic of taller vegetation, adding structural and
biological diversity to the site that is necessary for many of the grassland invertebrates.
Although the management plan (YWT, loc. cit.) calls for annual removal of parcels of scrub from
the earthwork and south-eastern boundary, in practice such interventions have been less
frequent, perhaps meaning that sources of scrub seeds that can encroach over the meadow are
more numerous than would be ideal, and that the shorter-growth regenerating areas are
actually encroaching onto the pasture rather than remaining around the fringes of the reserve.

One of the possible negative consequences of limited scrub invasion is that it facilitates the
spread of coarse grasses, such as False Brome. This was one of the plants found to have
significantly increased in calcareous grasslands in Dorset in recent decades (Newton et al.,
2011). Despite being abundant in our quadrats, this study provides no evidence for a recent
increase on Ellerburn Bank. Nine years is a relatively short timescale compared to the timescale

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of decades on which Newton's study was based, and yet the data solidly reject any rapid
increase but suggest that once established, False Brome does not inevitably continue to spread.

False Brome occupancy was significantly correlated with the first DCA axis (Fig. 4) whilst
Hawthorn was significantly correlated with the second. This suggests that both are non-
randomly distributed in relation to the other plants in the community. Non-random associations
in space could be caused by a number of different factors, including proximity to a seed or
vegetative growth source, establishment success and competitive exclusion of other plants.
Since these studies were observational rather than experimental, it is difficult to distinguish
these different possibilities, although competitive exclusion of stress-tolerating plants by
competitors would be expected (Newton et ol., loc. cit.). NMDS (Fig. 3) suggests that False
Brome tends not to be associated with several low-growing stress-tolerators such as Wild
Thyme, Harebell, Common Rock-rose, Felwort and Rough Hawkbit but that it is more associated
with Hogweed, Meadow Vetchling, Eyebright and Yellow Oat-grass. Whatever the causes of
these associations, these data do suggest that potential winners and losers were the frequency
of False Brome to either increase or decrease further, and they also suggest what is added and
lost through allowing a certain quantity of False Brome to establish on site. Given that False
Brome persists anyway around the scrubby fringes of the reserve, there is an argument for
increasing the intensity of grazing and scrub removal to avoid further establishment of False
Brome across the grassland at the expense of the shorter stress-tolerators.

Apart from scrub invasion, one further reason to expect increases in False Brome frequency is
the deposition of atmospheric nitrogen leading to eutrophication (Newton et ol ., loc. cit., van
den Berg et al., loc. cit.). Sampling from across the UK has shown that nitrogen deposition
significantly predicts reductions in floral diversity and evenness and the absence of rare plants.
Being on the North York Moors, Ellerburn Bank is expected to receive a high nitrogen deposition
load, estimated at 20.4 kgha'V 1 in 2008 (data from the Centre for Ecology and Hydrology in van
den Berg et ol., loc. cit.). This is within the range of critical loads for adverse effects on
calcareous grassland based on field experiments (Bobbink et ol., 2010). Comparisons of two long
term quadrats on the bank between 1990 and 2008 showed a slight increase in Shannon
diversity, a slight decrease in evenness and a slight reduction in richness (data from van den Berg
et ol., loc. cit., courtesy of Leon van den Berg). The majority of managed sites in the UK with a
similar nitrogen load actually increased in richness and evenness over the same time period,
whilst several decreased (van den Berg et ol., loc. cit., their Fig. 3), and those that increased
tended to have a higher soil pH. In addition, those that experienced an increase in grazing
pressure tended to experience reductions in their average Ellenberg nitrogen index. Ellerburn
Bank is sited on shallow, free-draining alkaline soil and experienced an estimated increase in
grazing pressure between 1990 and 2008 of approximately four times (data from van den Berg
et ol., loc. cit.). Grazing is likely to be the chief intervention responsible for the (slight) downward
trajectory of the Ellenberg nitrogen scores shown in the present study, and this adds to the
above arguments for at least maintaining the current level of grazing pressure. If, as expected,
nitrogen deposition remains a problem in the future, grazing should probably be increased to
compensate.

Part of the interest in maintaining the flora of the reserve is the conservation of the invertebrate
community that depends on it. The butterflies have received most attention in this regard; a

106

The Naturalist 140 (2015)

monitoring transect begun in 2010 (YWT, loc. cit.) showed that the most abundant species were
Small White Pieris ropae, Green-veined White Pieris nopi (ubiquitous species not relying on the
calcareous grassland but visiting flowers such as Knapweeds and Scabious), Meadow Brown
Maniolo jurtino and Small Heath Coenonympho pomphilus (both grass feeders). Small Skipper is
another common grass-feeder. Dingy Skipper and Common Blue are other abundant butterflies,
and both rely on the Common Bird's-foot-Trefoil which our quadrats show to be one of the most
abundant plants in the calcareous grassland. Common Rock-rose (supporting Brown Argus),
Milkwort (supporting Small Purple-barred moth), and violets (supporting Dark Green Fritillary)
are frequent but less dominant components of the community; both Common Rock-rose and
violets are more common on the earthwork than in the area of our quadrats but management
needs to take account of them given that these butterflies are less common on site and
regionally. Should the Duke of Burgundy ever return to the vicinity, it will still find abundant
Cowslips.

Taken together, our results suggest that current management of the calcareous grassland at
Ellerburn Bank is currently sufficient to offset the deleterious effects of atmospheric nitrogen
deposition, but a significant presence of woody scrub and False Brome persists, which is
probably undesirable away from the reserve fringes. There is a case for more frequent scrub
removal at the reserve fringes and increased grazing pressure on the grassland to prevent
further encroachment of scrub and the effects of atmospheric nitrogen deposition. Intermittent
monitoring will be necessary in future to gauge whether the current situation continues. To
facilitate this, we have provided details of the permanent quadrat locations in Appendix 1, a
summary of the species' occupancies from our survey in Appendix 2, and the average properties
of the quadrats in Appendix 3. Future monitoring work could also specifically target the plant
species of regional note (such as Fly Orchid and Woolly Thistle) which were not best surveyed
using the present methods, but which should feature highly in management priorities.

Acknowledgements

We thank Bill Ely and an anonymous referee for comments and corrections on the manuscript;
the Yorkshire Wildlife Trust, Forest Enterprise and the Thornton Dale Estate for permission to
work at Ellerburn Bank; Graham Young, Phil Dunning and Jim Cooper from York Metal Detecting
Club for annual assistance in relocating the quadrats; Alastair Fitter for initial identification
assistance and discussion of ideas about False Brome; Leon van den Berg for providing data on
past Ellerburn surveys; Jeremy Searle, who first decided to bring the field course to Ellerburn;
and the many students whose surveying work is recorded here and whose enthusiasm and
energy combined with the wonderful location to make it a very enjoyable enterprise.

References

Bobbink, R., Hicks, K., Galloway, J. et ol. (2010) Global assessment of nitrogen deposition effects on
terrestrial plant diversity: a synthesis. Ecological Applications, 20: 30-59.

Bobbink, R. & Willems, J.H. (1987) Restoration management of abandoned chalk grassland in the
Netherlands. Biodiversity and Conservation, 2: 616-626.

Butaye, J., Adriaens, D. & Honnay, O. (2005) Conservation and restoration of calcareous grasslands: a
concise review of the effects of fragmentation and management on plant species.
Biotechnology, Agronomy, Society and Environment, 9: 111-118.

The Naturalist 140 (2015)

107

English Nature (2004) A statement of English Nature's views about the management of Ellerburn
Bank Site of Special Scientific Interest (SSSI). Version date: 12/2/2004. Available at
http://www.sssi.naturalengland.org.uk, last accessed 4/7/2014.

Fisher, M. & Stocklin, J. (1997) Local extinctions of plants in remnants of extensively used
calcareous grasslands 1950-1985. Conservation Biology , 11: 727-737.

Fitter, R., Fitter, A. & Blarney, M. (1996) Wildflowers of Britain and Northern Europe, 5th Ed. Collins,
London.

Fitter, R., Fitter, A. & Farrer, A. (1984) Grasses , Sedges, Rushes and Ferns of Britain and Northern
Europe. Collins, London.

Frost, FI.M. (ed) (2005) The Butterflies of Yorkshire. Butterfly Conservation, Wareham.

Hill, M.O., Mountford, J.O., Roy, D.B. & Bunce, R.G.H. (1999) Ellenberg Indicator Values for British
Plants. ECOFACT Volume 2 Technical Annex. Institute of Terrestrial Ecology, Huntingdon.

Klimek, S., Kemmermann, A.R.G., Hoffman, M. & Isselstein, J. (2007) Plant species richness and
composition in managed grasslands: the relative importance of field management and
environmental factors. Biological Conservation, 134: 559-570.

Leadley, J. & Richards, J. (eds) (2012) Discover Yorkshire's Wildlife. Yorkshire Wildlife Trust, York.

Magurran, A.E. (2004) Measuring Biological Diversity. Blackwell, Oxford.

Natural England (2014) Condition of SSSI units: Ellerburn Bank (compiled 01/06/2014). Available at
http://www.sssi.naturalengland.org.uk, last accessed 4/7/2014.

Newton, A.C., Walls, R.M., Golicher, D., Keith, S.A., Diaz, A. & Bullock, J.M. (2011) Structure,
composition and dynamics of a calcareous grassland metacommunity over a 70-year
interval. Journal of Ecology, 100: 196-209.

R Core Team (2014) R: a Language and Environment for Statistical Computing. R Foundation for
Statistical Computing, Vienna.

Sutton, S.L. & Beaumont, H.E. (1989) Butterflies and Moths of Yorkshire: Distribution and
Conservation. Yorkshire Naturalists' Union, Doncaster.

Sykes, N. (1993) Wild Plants and their Habitats in the North York Moors. North York Moors National
Park, Helmsley.

Tansley, A.G. (1922) Studies of the vegetation of the English chalk II. Early stages of redevelopment of
woody vegetation on chalk grassland. Journal of Ecology, 10: 168-177.

UK Steering Group. (1998). UK Biodiversity Group: Tranche 2 Action Plans, Volume 2 - Terrestrial and
Freshwater Habitats. English Nature, Peterborough.

van den Berg, L.J., Vergeer, P., Rich, T.C.G., Smart, S.M., Guest, D. & Ashmore, M.R. (2011) Direct and
indirect effects of nitrogen deposition on species composition change in calcareous
grasslands. Global Change Biology, 17: 1871-1883.

WallisDeVries, M.F., Poschlod, P. & Willems, J.H. (2002) Challenges for the conservation of calcareous
grasslands in northwestern Europe: integrating the requirements of flora and fauna.
Biological Conservation, 104: 265-273.

Williams, J.M. (ed) (2006) Common Standards Monitoring for Designated Sites: First Six Year Report.
JNCC, Peterborough.

Wilson, J.B., Peet, R.K., Dengler, J. & Partel, M. (2012) Plant species richness: the world records.
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Yorkshire Wildlife Trust (2012) Ellerburn Bank SSSI Management Plan 2012-2017. Unpublished.

108

The Naturalist 140 (2015)

Appendix 1. Locations of permanent quadrats on Ellerburn Bank. Quadrat number refers to
identities on the map in Fig. 1 and in Fig. 3b.

Appendix 2. Complete plant list and mean (SE) occupancy (out of 25) per quadrat. Species are
assigned a letter code identifying them in the ordination plots (Figures 3 & 4).

Species (code)

Mean

Occupancy

(SE)

Species (code)

Mean

occupancy

(SE)

Species (code)

Mean

occupancy

(SE)

Achillea
millefolium (A)

0.06(0.06)

Filipendula
vulgaris (V)

0.77(0.34)

Primula veris

(q)

1.95(0.65)

Agrimonia
eupatoria (B)

2.56(0.45)

Fragaria vesca
(W)

0.70(0.25)

Prunella
vulgaris (r)

0.88(0.23)

Agrostis
capillarum (C)

0.06(0.03)

Galium verum (X)

8.17(1.78)

Prunus spinosa
(s)

0.01(0.01)

Agrostis
stolon if era (D)

N/A

Gentianella
amarella (Y)

0.87(0.20)

Quercus robur
(t>

0.03(0.02)

Anacamptis
pyramidal is (E)

0.02(0.01)

Flelianthemum
nummularium (Z)

0.08(0.06)

Ranunculus
acris (u)

0.01(0.01)

Anthoxanthum
odoratum (F)

0.03(0.03)

Heradeum
sphondylium (a)

0.56(0.24)

Rosa canina (v)

0.20(0.13)

The Naturalist 140 (2015)

109

Anthyllis 1.76(0.52)

vulneraria (G)

Holcus lonotus 0.74(0.25)
(b)

Brochypodium 7.76(1.45)
sylvaticum (H)

Briza media (I) 11.08(0.52)

| Bromopsis j 8.44(0.99)

e recto (J)

j Camponulo 0.08(0.05)
rotund if olio (K)

Knoutio 12.72(0.56)

arvensis (c)

Rubus

fruticosus agg.
(w)

Songuisorba
minor (x)

0.06(0.04)

16.96(1.44)

Carexflocco (L) 18.99(0.58)

Centoureo 10.23(0.90)
nigra (M)

Centoureo 0.45(0.16)

scobioso (N)

Cirsium 0.05(0.05)

eriophorum (0]j
Crataegus \ 1.93(0.43)

monogyno (P)

Doctylis 1 10 38(0.55)

glomeroto (Q)

Doctylorrhizo 2.46(0.44)

fuschii (R)

Elymus coninus 0.01(0.01)

US)

Euphrasia 0.13(0.77)

officinalis agg.

(T)

Festuca rubra 1.82(0.24)

uyL

Koeleria
macro ntha (d)

0.38(0.08)

Scabiosa
columbaria (y)

0.19(0.07)

Lathyrus
pratensis (e)

0.14(0.78)

r— ~~ ~ ■ *-]

Stellaria media
(?)

0.01(0.01)

Leontodon
hispidus (f)

12.17(2.01)

Taraxacum
officinale agg.
(Aa)

0.13(0.05)

Linum

catharticum (g)

4.68(0.55)

Thymus

polytrichus

(Bb)

0.31(0.15)

Listera ovata
(h)

0.02(0.02)

Tragopogon
pratensis (Cc)

0.05(0.02)

Lotus

corniculatus (i)

21.33(0.38)

Tri folium
pratense (Dd)

7.86(0.84)

Medicago
lupulina (j)

0.15(0.04)

Trifolium
re pens (Ee)

0.85(0.54)

Ophrys
insectifera (k)

0.03(0.02)

Trisetum
f lave see ns (Ff)

3.10(0.39)

Pilosella
officinarum (1)

2.10(0.30)

Ulex

europaeus (Gg)

0.10(0.06)

Pimpinella
saxifraga (m)

0.92(0.24)

Veronica

chamaedrys

(Hh)

0.03(0.02)

Pin us sylvestris
(n)

0.01(0.01)

Viola hirta (li)

1.99(0.52)

Plantago
lanceolata (o)

3.63(0.65)

Viola odorata

0.05(0.03)

Polygala
vulgaris (p)

1.50(0.34)

r— — — ™— —

Appendix 3. Changes in the lm2 quadrats across years. Quadrats are identified by their number
in Appendix 1. Numbers in each cell denote species richness, Simpson's diversity (1/D),
occupancy weighted Ellenberg nitrogen score, False Brome occupancy, and woody scrub species
occupancy.

Quadrat

2003

2004

2005

2008

2009

2011

21, 11.24,
3.85, 7, 4

23, 15.22,
3.43, 11, 7

21, 12.77,
3.59, 7, 5

24, 14.65,
3.48,4, 6

24, 12.72,
3.40, 5, 7

22, 15.25,
3.41, 11, 7

17, 6.47,
3.50, 24, 4

17, 9.56, 3.66,
25,4

21, 11.72,
3.63, 24, 3

21, 11.43,
3.35,21,8

17, 12.62,
3.69, 19, 7

19, 11.32,
3.67, 14, 9

19, 11.22,
3.67, 20, 0

17, 10.63,
3.41,21,0

21, 14.80,
3.57, 16, 0

21, 13.93,
3.48, 15, 0

22, 14.06,
3.54, 20, 0

20, 13.60,
3.43, 16, 0

110

The Naturalist 140 (2015)

22, 10.66,
3.44, 0, 11

20,9.39,3.15,
0, 10

18, 10.75,
3.41, 0, 10

19, 12.24,
3.19, 1, 11

24, 15.62,
3.43, 4, 20

23, 13.68,
3.35, 3, 7

23, 14.95,
3.61, 0, 7

23, 14.44,
3.49, 2, 3

20, 12.48,
3.55, 4,3

26, 14.71,
3.31, 4, 2

19, 14.18,
3.62, 4,4

23, 14.23,
3.41, 1, 3

22, 10.41,
3.60, 5, 3

23, 11.84,
3.38, 8, 2

22, 12.56, 20, 11.04,

3.65, 3, 2 3.65, 7, 2

24, 14.05,
3.39, 9, 2

25, 14.73,
3.64, 13, 2

21, 11.00,
3.49, 10, 0

17, 11.66,
3.11, 15,0

21, 12.88,
3.47, 11, 0

20, 13.24,
3.24, 15, 0

13, 11.19,
3.36, 18, 0

18, 12.28,
3.29, 17, 0

17, 8.48,
3.44, 22, 0

17, 9.65, 3.15,
25,0

19, 12.57,
3.42, 17, 0

21, 12.78,
3.42, 19, 0

18, 11.18,
3.23, 20, 1

23, 12.66,
3.31, 23, 1

19, 11.37,
3.29, 22, 1

15, 9.29, 3.36,
25,3

21, 12.76,
3.50, 23, 4

22, 13.56,
3.32, 15,4

21, 12.51,
3.29,18,2

15, 11.52,
3.42, 21, 3

26, 15.54,
3.40, 24, 1

17, 10.34,
3.40, 25, 0

22, 13.78,
3.31, 6,1

18, 12.36,
3.11,8,0

16, 10.82,
3.33, 10, 1

20, 12.56,
3.44, 10, 1

26, 16.19,
3.47, 10, 1

19, 10.69,
3.14, 8,0

18, 11.81,
3.64, 8, 0

19, 13.39,
3.55,17,1

25, 17.65,
3.33, 10, 2

23, 13.84,
3.44, 12, 1

21, 10.86,

3.27, 1, 2

20, 10.87,
3.21, 2, 2

24, 15.60,

3.40, 4, 0

' ' :

25, 15.15,
3.37, 9, 4

17, 11.57,
3.68, 1, 0

20, 11.78,
3.42, 2, 1

21,13.65, j 24, 17.07,
3.24,1,0 3.01,3,0

22, 13.76,
3.10, 0,0

22, 13.85,
2.94, 4, 1
28, 16.90,
3.74, 19, 5

16, 10.82,
3.33, 10, 1

22, 14.73,
2.97, 0, 2

24,15.09,
3.34, 2, 2

21, 12.70,
3.35, 0, 2

25, 12.61,
3.58, 0, 0

16, 9.93,
3.43, 1, 0

20, 14.08,
3.71, 4, 2

15 j

18, 11.61,
3.17, 0, 0

18, 12.54,
3.23, 0, 2

19, 11.51,
! 3.41, 0, 0

21, 13.05,
3.28, 7, 0

19, 11.10,
3.15, 1,0

20, 11.88,
3.17, 4, 3

I J

20, 11.79,
3.31, 8, 0

24, 15.48,
3.31, 14, 2

28, 16.54,
3.31, 7,4

26, 18.09,
3.35, 10, 2

19, 13.49,
3.35, 18, 3

25, 14.73,
3.23, 15, 1

2.97, 1,2 1,1
20,11.42, 22,12.62,

3.26,14,1 3.09,1,0

3.36, 3, 0
26, 15.87,

3.45, 9, 1

20, 10.82,
3.16, 0, 3

20, 11.57,
3.08, 0, 7

28, 17.39,
3.38, 0, 3

24, 11.78,
3.21, 0,4
14, 8.85,
3.42, 2, 1

21, 12.78,
3.42, 0, 4
20, 12.25,
3.36, 3, 5

22, 12.47,
3.55, 7, 2

19, 10.53,
3.54, 5, 2
19, 10.04^ |l7, 10.13,
3.30, 0, 0 [ 3.20, 0, 3
2l7 10.45, [l6, 9597^87,
3.03, 0, 0 0, 0

16, 11.02,
3.47, 0, 3
25, 12.79,
3.01, 6, 1
20, 12.07,
3.66, 6, 3

23, 12.10,

3.46, 2, 0
20, 12.02,
3.21, 0, 0

3.31, 0, 3
21, 12.64,
3.23, 8, 1
21, 10.98,
2.98, 1, 1
25, 15.56,
3.35, 0, 5
21, 13.91,

3.46, 0, 6
20, 11.39,

D.± / , / , U

24, 12.97,
3.38, 0, 0
17, 12.42,
3.12, 5,0

2.95, 5,0

20, 11.88,

3.14, 7, 1
20, 13.22,

3.14, 0,5

20, 12.73,

3.15, 0, 4

21, 12.16,
3.33, 1, 5
21, 13.24,

3.47, 9, 0
18,10.00,
3.24, 1, 0

2.91, 2, 1

23, 15.44,
3.20, 10, 2
27, 12.08,
3.04, 0, 4

24, 14.63,
3.10, 0, 6
19, 12.30,
3.17, 1, 0
24, 14.38,

3.47, 10, 0

19, 12.35,

3.16, 0, 1

15, 10.04,
3.10, 0,2

19, 11.27,
3.03, 0, 1

The Naturalist 140 (2015)

111

Local effects of climate change - has the date of first
emergence changed in several species of Lepidoptera in
Yorkshire during the period 1995 to 2014?

David R. R. Smith1 and Heather A. R. Smith

department of Psychology, University of Hull, Hull, HU6 7RX, UK.

Email: davidsmith.butterflies@gmail.com

Introduction

The Earth has undergone a period of sustained growth in global average temperature from the
early twentieth century onwards (Stott et oi, 2000; IPCC, 2013). The impact of substantial
climate change upon Lepidoptera includes such changes as shifts in first emergence date and
peak flight date, in flight period length and pattern, in voltinism, abundance, distribution, etc.
(e.g., Sparks & Yates, 1997; Roy & Sparks, 2000; Asher et oi, 2001; Diamond et oi, 2011;
Karlsson, 2014). Given that climate change is likely to affect all flora and fauna to a greater or
lesser extent, and that each species lies in the middle of a complex web of interdependencies
with the rest of nature, the way that climate change ramifies upon any given species may not be
simple.

Yorkshire is a particularly interesting arena to test for the effect of global temperature increases
upon butterfly emergence patterns, lying as it does at a sufficiently northerly latitude (between
53° 18' N and 54° 40' N) that many butterflies are (or were) at the northern edge of their range
where they are particularly sensitive to climate change. Interestingly, in the last few decades, we
have seen butterflies once rare or absent in Yorkshire, such as the Comma Polygonio c-olbum,
Speckled Wood Porarge oegerio and Holly Blue Celostrina orgiolus, sweep northwards to
become commonplace (Asher et oi, 2001; Fox et oi, 2007). Butterflies are poikilothermic, and
are heavily dependent on external air temperature and incident sunlight to raise their body
temperature to levels at which they are able to mate and lay eggs (at least 18-28°C). During
winter months, butterflies enter diapause - a period of physiological dormancy to survive colder
temperatures - in a variety of overwintering states (e.g., Brown Hairstreak Theclo betuloe, egg;
Meadow Brown Maniola jurtino, larva; Orange-tip Anthochoris cordomines, pupa; Brimstone
Gonepteryx rhomni, adult). As global surface temperatures have increased since the early
twentieth century onwards (IPCC, 2013), this has coincided with earlier emergence (e.g., Sparks
& Yates, 1997; Roy & Sparks, 2000), perhaps because the insects are awakened from diapause at
an earlier point of the year.

We wished to (a) quantify date of first emergence of a range of butterfly species in Yorkshire
from 1995 to 2014, (b) investigate what changes in temperature there might have been in
Yorkshire over the same time period and (c), explore the relationship between date of first
emergence and temperature. The years between 1995 and 2014 represent the longest range
over which we have detailed and plentiful records of butterflies in Yorkshire, representing a
period of time of sufficient length to potentially demonstrate historical change in phenology

112

The Naturalist 140 (2015)

(Roy & Sparks, 2000) and is the first systematic analysis of local effects of climate change upon
date of first emergence in Yorkshire's butterflies.

The criteria for the choice of species were that the butterflies should be a mix of habitat
generalists and specialists, have an early spring emergence, be easily identifiable by recorders,
should not overwinter as adults, should be from different families and be present in sufficiently
large numbers so as not to invite sampling problems. Some of these choices are motivated by
pragmatic reasons, such as a wish to increase our sampling pool, to reduce opportunities for
recorder misidentifications1 and to discount early sightings of hibernating adults due to physical
disturbance or one-off warm days. The choice of having a mix of generalists and specialists,
across a range of families, is to strengthen the applicability of our results to all butterflies.
Finally, studies have shown that earlier emergence of butterfly species is especially marked for
spring species (Sparks & Yates, 1997; Roy & Sparks, 2000). As such, we chose a 'Nymphalid'
(Nymphalidae), the Speckled Wood; a 'Blue' (Lycaenidae), the Holly Blue ; a 'White' (Pieridae),
the Orange-tip Anthocharis cordomines; and a 'Skipper' (Hesperidae), the Dingy Skipper Erynnis
tages.

Method

We searched the Butterfly Conservation database for records of our target species for the period
1995 to 2014 from the five Watsonian vice-counties (VC61-VC65) traditionally comprising the
county of Yorkshire for recording purposes. For each year we found the dates of the five earliest
records of each target species and took the mean. This provided a reasonably unbiased estimate
of each year's date of first emergence for each species.

We analysed the butterfly data using a bivariate Pearson's correlation, with the variables of year
of recording and mean date of the five earliest records for each year. We are interested in
exploring how date of first emergence and year co-vary, so we adopted a non-directional two-
tailed analysis. The equation of the best-fitting line (least squares linear regression) to the data
was used for the purposes of quantifying changes in date of first emergence. This general
statistical approach was used to analyse local and regional temperature data series (variables
year of recording and temperature), and to compare temperature and date of first emergence.

Results

Date of first emergence was significantly inversely related to recording year for Speckled Wood
(r(16) I -0.52, 95% BCa Cl [-0.824, -0.057], p = 0.027), Orange-tip (r(18) = -0.57, 95% BCa Cl [-
0.781, -0.261], p = 0.009), Dingy Skipper (r(18) = -0.45, 95% BCa Cl [-0.747, -0.180], p = 0.049),
and insignificantly related to recording year for Holly Blue (r(18) = -0.19, 95% BCa Cl [-0.680,
0.422], p = 0.422 NS). The scatter plots and lines of best-fit (linear regression) are shown in
Figure 1.:

1 Small Whites Pieris rapae and female Green-veined Whites Pieris napi being an obvious
misidentification pair

The Naturalist 140 (2015)

113

Recording Year

O Speckled Wood

y = -x + 2104.6
R2 = 0.2693

Recording Year

O Orange-tip

-1.2128X+ 2530
R2 = 0.3236

Recording Year

1990 1995 2000 2005 2010 2015

Recording Year

O Dingy Skipper

-0.7609X+ 1653.1
R2 = 0.1989

Figure 1.

Day in year of first emergence, as
a function of recording year, for
Speckled Wood Porarge oegerio,
Orange-tip Anthochoris

cordomines, Holly Blue Celastrinia
aegiolus and Dingy Skipper
Erynnis toges (pll5).

Day of first emergence in each
year is calculated by taking the
mean of the dates of the five
earliest records of each target
species. The number of records
for 1995 and 1996 were so few for
Speckled Wood that those years
were omitted from the analysis,
hence Speckled Wood is analysed
between 1997 to 2014. The best-
fitting least squares linear
regression line for 1995 to 2014
(Speckled Wood) is dotted and
shown for illustrative purposes
only. Solid best-fitting least
squares linear regression lines are
calculated for 1995 to 2014 for all
species, except Speckled Wood
where it is calculated for 1997 to
2014. Equation shown is for solid
least squares linear regression
lines. Error bars represent ± 1
standard deviation.

114

The Naturalist 140 (2015)

The day of first emergence of Speckled Wood has shifted 17 days earlier2 in the year between
1997 and 2014; equivalent to 1 day per year or 10 days per decade. Thus in 1997 the mean date
of first emergence, using the linear regression equation, would have been 18 April and this had
shifted to 1 April by 2014. For the other three species between 1995 and 2014, the shifts are 23
days earlier in the year for the Orange-tip (equivalent to 1.21 days per year); 9 days earlier in the
year for the Holly Blue (equivalent to 0.47 days per year) and 14 days earlier in the year for the
Dingy Skipper (equivalent to 0.74 days per year). This is consistent with earlier work on British
butterflies (Sparks & Yates, 1997; Roy & Sparks, 2000) which suggested climate warming of 3°C
could advance date of first emergence by two to three weeks. Table 1 summarises these
changes in phenology with additional statistical findings taking into account the variability of the
data over the recording years. If a weighted correlation, that takes into account the variability of
the data, is applied then the pattern of results is unchanged but the correlation coefficients and
level of significance increase even more. There is an undeniable shift towards earlier emergence
in the year over the last two decades for three of the butterfly species.

Table 1. Species statistical summary

Species

Day(s)/
yr shift

Start

yr3

End

yr3

Start

year

dfe4

End

year

dfe

r 5

• w

O
A Jw

i j

Speckled

Wood

Pararge

aegeria

-0.52

0.027'

1.00

1997

2014

18 April

1 April

-0.53

0.025*

Orange-tip

Anthocharis

cardamines

-0.57

0.009**

_ . j

1.21

1995

2014

20 April

March

-0.86

<0.00001

T_ , . -T-T j

Holly Blue
Celastrina
argiolus

-0.19

0.422

(NS)

0.47

1995

2014

: ... J

17 April

8 April

-0.30

0.205

(NS)

\ ]

Dingy Skipper
Erynnis tages

-0.45

0.049*

0.74

! I

1995

2014

L„__J

15 May

2 May

1 j

-0.54

0.013'

Pearson's r 2two-tailed 3Start and end year within UK BMS (BCY) database 4Date of first
emergence 5Pearson's r calculated with each year's data weighted in proportion to its variability.
*significant p<0.05 **significant p<0.01 ***significant p<0.001

What should be considered now is whether there has been an actual shift in average
temperature in the Yorkshire region in the period of study. Figure 2 plots spring and summer
temperature series covering the period from 1995 to 2014 - one from Sherburn in Elmet
(roughly in the centre of the county with Lat. 53° 47' 48" N, Long. 1° 15' 26" W, Elev. 27 m) and

2 The least-squares line of best fit has equation y=-x + 2104.6 (see Figure 1). This is the straight-line

linear form y=mx + c , where y is date of first emergence (day in year), x is recording year, c is y-axis

intercept (^constant) and m is the gradient (=1). The first year of recording period was 1997 so,
substituting into the equation, gives the answer of day in year of 1 08 (rounded to nearest whole
integer). This equates to 18 April. Similarly, substituting in last year of the recording period of 2014,
gives the answer of day in the year of 91 (=1 April). This is a difference of 17 days. See Table 1 for
further details and the other species calculations.

The Naturalist 140 (2015)

115

the other from the Central England Temperature (CET) data series. The CET series gives mean
temperature readings from the Midlands (extending back to 1659) and is taken to be a
reasonable proxy for temperature in other parts of the UK (Duncan, 1991).

& 8

1990 1995 2000 2005 2010 2015

Year

QSpring- Sherburn
•Summer - Sherburn
■ Spring - CET
□ Summer -CET

Figure 2. Temperature time series for spring (mean February— April) and summer (mean May—
July) calculated from a weather station at Sherburn in Elmet and the Central England
Temperature (CET) data set, between the years 1995—2014. Best-fitting lines (least squares
linear regression) are solid for Sherburn in Elmet and dotted for CET.

There is little discernible shift in mean spring (February— April), summer (May— July), autumn
(August— Oct)3 and winter (November— January) temperature in Yorkshire between 1995 and
2014. Yorkshire seasonal temperature data series show no significant trends over the twenty
year time period: spring temperature series r(18) = -0.18, 95% BCa Cl [-0.680, 0.357], p = 0.44
A/S; summer temperature series r( 18) = -0.043, 95% BCa Cl [-0.524, 0.423], p = 0.858 A/S; autumn
temperature series r(18) = -0.339, 95% BCa Cl [-0.738, 0.258], p = 0.143 A/5; winter temperature
series r(18) = 0.168, 95% BCa Cl [-0.317, 0.777], p = 0.478 NS. The cross-check from the CET
temperature series against the Sherburn data series shows close agreement. An analysis of the
CET and Sherburn data for spring and summer show them to be significantly correlated: spring
r( 18) = 0.96, 95% BCa Cl [0.920, 0.986], p < 0.001; summer r(18) = 0.89, 95% BCa Cl [0.746,
0.944], p < 0.001. Therefore we can be confident that the Sherburn data is representative of UK
temperatures and is not anomalous. If we are to make some link between earlier date of first
emergence and increased spring temperatures in the year of emergence (say), then it is not
obviously to be found in increased mean spring temperature because there is no increase.

We should not (and logically cannot) rule out the date of first emergence as being temperature
dependent. The problem is that what exactly is driving earlier emergence could be buried deep
within the temperature series. It is beyond the scope of this report to exhaustively chase all
possible factors. We checked for correlations with preceding season (autumn, winter, spring,
summer) temperatures and date of first emergence in the subsequent year. In no instance was

3 To avoid clutter the autumn and winter temperature series for Sherburn and CET are not shown in
Figure 2.

116

The Naturalist 140 (2015)

there a significant correlation between preceding season and subsequent year date of first
emergence (see Table 2). There are then no obvious associations between the temperature data
and changes in the dates of first emergence. Perhaps earlier trends in increased temperature
(such as the 1.5 °C increase in central England spring temperatures between 1976 and 1998 (Roy
& Sparks, 2000)) have induced a long-term change that takes years to work through the gene
pool of the population? This is beyond the scope of this short report.

Table 2. Correlation between preceding season temperature and date of first emergence in
subsequent year (statistical summary)

Species

Season

7“

95% BCa Cl3 [lower,
upper]

, . _ , ,

Speckled Wood Pararge

autumn

winter

-0.27

0.298

[-0.636, 0.202]

aegeria

-0.06

0.810

[-0.567, 0.194]

\ 1

spring

0.27

0.293

[-0.261, 0.742]

summer

-0.29

0.268

[-0.676, 0.355]

Orange-tip

autumn

0.30

0.213

[-0.257, 0.649]

Anthocharis cardamines

winter

0.03

0.896

[-0.525, 0.400]

spring

0.26

0.290

;

[-0.138, 0.639]

■ • 1

summer

-0.07

0.776

[-0.478, 0.321]

Dingy Skipper Erynnis

autumn

0.20

0.414

[-0.532, 0.640]

tages

winter

0.07

0.774

[-0.494, 0.473]

spring

0.37

0.123

[-0.127, 0.770]

summer

-0.09

0.716

[-0.702, 0.374]

Pearson's r 2two-tailed 3Bias corrected accelerated confidence intervals for the
correlation coefficient.

Another concern is a possible intervening relationship between increased observer effort and
earlier emergence dates. Basically, the more 'abundant' the butterfly (which again might be no
more than an intervening variable for having more observers around actively recording), then
the greater the chance for a given individual butterfly to be seen. Thus increased observer effort
or increased abundance can theoretically lead to a pattern of apparent earlier emergence. This
is hinted at in Figure 1 for Speckled Wood, where the mean dates of first emergence for 1995
and 1996 were both late in the year and associated with a great deal of uncertainty (shown by
the large standard deviations). The dates of first emergence were presumably late in the year
and more variable because Speckled Wood was present in very small numbers in those years
(having only just reached the southern borders of Yorkshire in a general northwards movement,
see Asher et oi, 2001), so the chance of spotting a Speckled Wood was relatively small. This
could lead to a confounding effect where the specimens seen are not necessarily newly
emerged but could be post-emergent adults by a number of days or even weeks. However, it is
hard to see how changes of up to 17 days (Speckled Wood) could be accounted for by an
increased chance to spot individual butterflies once a species has become relatively well
established in the region. Interestingly, Speckled Wood abundance peaked in Yorkshire in 2009
with recorded numbers in the last five years dropping by a factor of two but with little
discernible change in observed date of first emergence. The particularly late date of first

The Naturalist 140 (2015)

117

emergence in 2013 was attributable to the severe spring in that year. Further work on possible
interactions of increased observer effort and changes in recorded first emergence will be
needed to clarify the role of climate change upon driving changes in phenology, both at the local
and global scale.

Acknowledgements

We are grateful to the UK BMS and Butterfly Conservation and its volunteer recorders, for the
butterfly records from Yorkshire (VC61-65) upon which this research is based. Dave Ramsden
provided the temperature series for Sherburn in Elmet and the Central England Temperature
(CET) series was downloaded from www.metoffice, gov.uk/hadobs. We wish to thank an
anonymous reviewer for helpful comments.

References

Asher, J., Warren, M., Fox, Richard, Harding, P., Jeffcoate, G., & Jeffcoate, S. (2001) The
Millennium Atlas of Butterflies in Britain and Ireland , Oxford University Press.

Diamond, S. E., Frame, A. M., Martin, R. A., & Buckley, L. B. (2011). Species' traits predict
phenological responses to climate change in butterflies. Ecology, 92, 1005-1012.

Duncan, K. (1991). A comparison of the temperature records of Edinburgh and central England.
Weather, 46, 169-173.

Fox, R., Warren, M. S., Asher, J., Brereton, T. M., & Roy, D. B. (2007). The state of Britain's
butterflies 2007. Butterfly Conservation and the Centre for Ecology and Hydrology,
Wareham, Dorset.

IPCC, 2013: Climate Change 2013: The Physical Science Basis. Contribution of Working Group I to
the Fifth Assessment Report of the Intergovernmental Panel on Climate Change.

[Stocker, T. F., Qin, D., et al. (eds)]. Cambridge University Press, Cambridge, UK and New
York, NY, USA, 1535pp.

Karlsson, B. (2014). Extended season for northern butterflies. International Journal of
Biometeorology, 58, 691-701.

Roy, D. B., & Sparks, T. H. (2000). Phenology of British butterflies and climate change. Global
Change Biology, 6, 407-16.

Sparks, T. H., & Yates, T. J. (1997). The effect of spring temperature on the appearance dates of
British butterflies 1883-1993. Ecography, 20, 368-374.

Stott, P. A., Tett, S. F. B., Jones, G. S., Allen, M. R., Mitchell, J. F. B., & Jenkins, G. J. (2000).
External control of 20th century temperature by natural and anthropogenic forcing.
Science, 290, 2133-2137.

118

The Naturalist 140 (2015)

A Question of Ecology answers from biological recording

Paula Lightfoot, NFBR

The British Ecological Society Macroecology Special Interest Group and the National Forum for
Biological Recording organised a joint conference in late April 2015 at the University of Sheffield.
The event was attended by 100 delegates from a wide range of organisations involved in
collecting, managing, interpreting and using biodiversity information.

Biodiversity information is crucial to understanding ecological relationships and supporting
conservation effort in a changing climate. Use of volunteer-collected biological records by the
professional scientific community is widely encouraged and celebrated, but interpretation of
biological records is also carried out by amateur naturalists, who are uncovering new ecological
knowledge from their own records and sharing that knowledge with others. Biological recording
is not just about producing checklists, dot maps or providing 'big data' for others to analyse; it is
a way of engaging with the natural world which raises questions and provides answers to them.

A Question of Ecology celebrated achievements, highlighted opportunities and sought to
overcome obstacles regarding the use of biological records to answer ecological questions.

The conference aimed to:

• Raise awareness of how biological records can be interpreted to answer ecological

questions and lead to conservation action.

• Empower volunteer recorders and their organisations to get more out of their biological

records by highlighting effective approaches to data collection and analysis.

• Foster collaboration between the professional research community and volunteer

recording community through examples of good practice.

• Discuss barriers to the use of biological records for research and start a dialogue between

the recording and research communities about how to overcome those barriers.

The conference began with a demonstration workshop on software and tools for capturing and
interpreting biological records, including rNBN, SPARTA (Species Presence/Absence R Trends
Analyses), Scratchpads, Indicia and QGIS. This was a lively interactive session, and delegates
were inspired by the enthusiasm of the presenters and the opportunities provided by new
technology.

This was followed by a workshop in which challenges and opportunities regarding the collection
and interpretation of biological records for ecological research were discussed. Topics included
DNA techniques, Open Data and data quality. The diverse range of sectors present at the
conference and the mix of delegates' experience, knowledge and viewpoints ensured a useful
and informative debate. One delegate from the research community stated that "the most
valuable thing about the conference was the opportunity to fill the gap between my models and
the data that goes into them, by speaking directly with the people that collect those data."

The programme on the second day highlighted original work at all scales and levels, from global
to local, where knowledge of species and systems is being advanced through accurate

The Naturalist 140 (2015)

119

observation and recording. Professor Kate Jones from University College London and Bat
Conservation Trust opened proceedings with an excellent keynote address on the topic of
Technology for Nature? This was followed by a session dealing with methods for analysing 'big
data' to understand a changing environment, with speakers including Dr David Roy from the
Biological Records Centre and Dr Jon Yearsley from University College Dublin. Delegates from the
volunteer recording community and the professional research community alike were inspired to
learn how large, unstructured datasets can be analysed to provide insight into trends in
populations and ranges.

The focus then shifted to the collection and interpretation of data on a local scale, particularly
the role of local environmental records centres and natural history societies in supporting this.
Dr Teresa Frost presented examples of how Cumbria Biodiversity Data Centre empowers
volunteer recorders to get more out of their biological records by assisting with survey design,
data management and analysis. Dr Andy Millard reported on the initiatives being undertaken by
the YNU, with particular reference to the use of The Naturalist to disseminate information.

After lunch, delegates heard how biological records from museum specimens and naturalists'
diaries can be explored and interpreted to answer ecological questions. Ivan Wright from
Shotover Wildlife described how local naturalists are following in the footsteps of eminent
entomologists of the Victorian and Edwardian era, comparing current data to historic records to
establish a new benchmark of knowledge for a fascinating and diverse SSSI near Oxford. Dr Mark
Spencer from the Natural History Museum, London, explained how new approaches to citizen
science and crowd-sourcing can release a wealth of scientific information from the natural
history collections that await discovery in our museums.

Speakers from Newcastle University and the Woodland Trust went on to describe cutting-edge
citizen science projects that are engaging amateur naturalists in structured data collection to
support hypothesis-driven science, fostering collaboration between professional researchers,
the public sector and the volunteer recording community. The conference concluded with case
studies to highlight how research based on biological records by amateur naturalists is leading to
conservation action, from site-specific to landscape-scale examples. An excellent range of
posters were displayed, complementing the programme of talks. Peter and Sharon Flint
presented their investigation of the caddisflies of Malham Tarn (see page 121) in one of these.

On Saturday there was a very enjoyable and well-attended field meeting to Thorne and Hatfield
Moors, organised by Yorkshire Wildlife Trust and Natural England. Thorne and Hatfield Moors
form the core of the Humberhead Peatlands NNR, the largest lowland raised mire system in the
UK and the beating heart of the Nature Improvement Area. This was an excellent opportunity
to see examples of how science can influence conservation action and delivery on a landscape
scale - and to get out and enjoy some biological recording at a beautiful site! Species records
from the field trip have been collated and will be shared via the NBN Gateway.

NFBR and BES Macroecology SIG would like to thank the speakers, chairs, workshop facilitators,
software demonstrators, poster exhibitors and all the delegates for participating so actively and
enthusiastically and making the conference such a great success! A version of this report has
been published in the BES Bulletin.

120

The Naturalist 140 (2015)

Plate I. Ellerburn Bank. See pp 96-111.

Above left (a): View of the sward looking south. Visible are Common Spotted Orchids, Common
Bird's-foot-trefoil, Salad Burnet, Upright Brome, Quaking Grass and Cock's-foot.

Above right (b): Looking east along the reserve, a patch of invading scrub, mainly Hawthorn, sur-
rounded by the wide yellow leaves of False Brome.

Below left (c): Gorse invading the south-eastern part.

Below right (d): View of the sampled area at Ellerburn Bank, looking north-east. Small Hawthorn and
Gorse bushes are visible in the grass, with yellow patches of False Brome.

P. Moyhew

b) The Malham Sedge, mature larva and its case. S. Flint

c) The Grouse Wing Mystacides longicornis, adult male on

a rock on the north shore of Malham Tarn. S. Flint

d) Egg masses of Limnephilus politus, with spent females,

among moss-covered rocks at the high water level on the
north shore of Malham Tarn. S. Flint

e) Phryganeid egg ropes, on rocks in shallow water off the

west shore of Malham Tarn. 5. Flint

f) Grapnel sampling Stonewort Chara sp., from a boat on

Malham Tarn off the wooded shore near the East Boat-
house. P. Flint

Plate II continued

g) Larval case of Limnephilus politus; fixed to leaf of Potomogeton lucens and with the ends closed
using fragments of marl; ready for pupation.

h) Limnephilus politus, adult. S. Flint

Plate III. Ringinglow Bog, main flush area. See pp 134-145

a) vegetation dominated by Common Cottongrass, December 2014.

b) bog-moss amongst Common Cottongrass, September 2014.

c) tussocks of Hare's-tail Cottongrass, February 2015.

d) bog-moss amongst Hare's-tail Cottongrass, February 2015.

R. Goulder

Plate V. Biodiversity of moths in Borneo and China.
See pp 81-96.

Above: A few of the circa 2,500 Q-CAS voucher speci-
mens from Yunnan, China.

Right: Dr Louise Ashton with a Pennsylvania moth
trap in tropical forest understorey.

Below: Antheroeo lorissa ssp. ridlyi; Lepidoptera:
Saturniidae. A Bornean rainforest moth.

T.Whitaker

Plate IV. BES/NFBR Conference. See pll9.
Delegates on the field trip to Thorne and
Hatfield Moors.

P.Lightfoot

An investigation of the caddisfly (Insecta: Trichoptera) fauna of
the Malham Tarn NNR; with special reference to the Malham
Sedge Agrypnetes crassicornis

S. Flint and P.W.H. Flint

email: flintsentomologists@btinternet.com

Introduction

During the 1950s some of the members of the entomological section of the YNU carried out an
extensive investigation of the insect fauna of the area around Malham Tarn. For a period of
seven to ten days each year from 1954-1958 this group, led by W. D. Hincks and J. H. Flint, were
resident at Tarn House which had recently become a Field Studies Centre run by the Field
Studies Council. The then warden of the centre, P. F. Holmes, who had a strong interest in
caddisflies, took an active part in this investigation and wrote the section of the published report
(Henson, 1963) concerning them. Seventy one species of caddis flies were recorded in the study
area; which included the upper reaches of the nearby Gordale, Cowside and Darnbrook becks.
This is approximately a third of the British caddis fly list and underlines Malham as an important
area for this group of insects. Most of the caddis collecting was carried out by P.F. Holmes (who
incorporated his records from 1948 onwards into the report) and A. Brindle. During 2013 a study
of the caddisflies of the current Malham Tarn National Nature Reserve, a much smaller area
than that covered by the 1950s study, was undertaken by S. and P.W.H. Flint. A good description,
and historical review, of this high moorland site (centred on NGR SD890670; at c.380m above
sea level) set in the karst landscape of the Yorkshire Dales National Park, is given by Corey Jones
(Centre Director of the Malham Tarn Field Centre 1997-2000) in the magazine British Wildlife
(Jones, 2001) and need not be repeated here.

Aims

The aims of the 2013 study were threefold. Firstly, as there had been no concentrated work on
the caddis of the area for many years, we wanted to bring the list of the Malham caddis fauna
up to date; and into line with current taxonomy. Were the species which had been previously
recorded still present? Secondly we wanted to find out more about the ecology of these
species. What observations could we make of adult behaviour such as swarming and
oviposition? Where were the larvae to be found; and what were they doing? Thirdly we wanted
to know whether Agrypnetes crassicornis McLachlan (recently honoured with the vernacular
name 'Malham Sedge') discovered at Malham Tarn by P.F. Holmes in 1950, was still present as a
breeding species in its only known, British, locality.

Methods

Larvae were collected by pond netting in the standing and running water habitats where this
was practicable and by hand searching where use of the net was not practicable. A grapnel was
used for sampling the vegetation in the tarn both from the shore and from a boat. Collecting
commenced in mid-April and continued until the end of October.

The Naturalist 140 (2015)

121

Adults were collected from the ground vegetation, and from the lower canopy of the trees, by
the use of a sweep net. The net was also used to catch flying specimens. A malaise trap was
deployed on Malham Fen, the catch from which was collected fortnightly from the middle of
April to the end of October. The catch from an actinic light trap, situated in one of the cottage
gardens, just off the fen, and operated from the end of April until the end of July, was also
examined. Hand searching of the rocks around the tarn shore, and other likely resting places
such as the boat houses, birdwatching hide, tree trunks and post and rail fences, was also
undertaken during the day. Observations were extended well into the evenings as many species
are crepuscular and this was thought to be the best time to find the adults of A. crossicornis.

Nomenclature follows the checklist in Barnard and Ross (2012). Most caddis species do not have
vernacular names but some have been included, where appropriate, in the body of the text.
Adults were identified using both Barnard and Ross, and Macan (1973). Larvae were identified
using Edington and Hildrew (1995) and Wallace, Wallace and Philipson (2003).

Results

Table 1. List of species recorded during the 2013 survey compared with the list published in
1963.

1 1963 List

Adults

2013 List
Larvae

All All
records records

Taxa

Rhyacophilidae

Rhyacophila dorsalis
[ R. fasciata
R. munda
R. obliterata
Glossosomatidae
Agapetusfuscipes
Hydroptilidae
Agraylea
multipunctata
A. sexmaculata
Hydroptila angulata
H. forcipata
H. tineoides
H. vectis

Oxyethira falcata

■q.

'cl

Ll_

-t— >

+->

c
CD
1 1

op
' 1

■4— 1

122

The Naturalist 140 (2015)

0. flavicorn[s
Philopotamidae

Philopotamus
montonus
Wormoldio subnigra

Polycentropodidae

Cyrnus flavidus

C. trimoculatus

Neureclipsis
bimoculoto
Plectrocnemio
consperso
Polycentropus
flavomoculotus
P. irrorotus
Psychomyiidae
Lype phaeopa
Psychomyio frogilis
P pusilla

Tinodes dives

T. rostocki

T. waeneri

Hydropsychidae

Hydropsyche

instabilis

H. siltalai

Phryganeidae

Agrypnetes

crassicornis

Agrypnia obsoleto

Oligotricha striata

Phryganea
bipunctata
Lepidostomatidae
Lepidostoma hirtum
Limnephilidae
Drusus annulatus
Ecclisopteryx
dalecarlica

Chaetopteryx villosa
Anabolia nervosa

Glyphotaelius
pellucidus
Limnephilus affinis
L. auricula

L centralis

The Naturalist 140 (2015)

123

L. coenosus

L. elegans

L extricatus

L griseus

]

L. hirsutus

L. incisus

L lunatus

L luridus

L politus

L. rhombicus

L sporsus

L. stigma

L. vittatus

Rhadicoleptus

alpestris

Halesus digitatus

H. radiatus

Hydatophylax
inf u mat us

Melampophylax

mucoreus

Mesophylax

impunctatus

Micropterna lateralis

M. sequax

Potamophylax

latipermis

P. rotundipennis

| P. stellatus

Stenophylax

permistus

S. i /ibex

Sericostomatidae

Sericostoma

personatum

Beraeidae

Beraea pullata

Beraeoides minutus

Odontoceridae

Odontocerum

albicome

Leptoceridae

Arthripsodes cinereus

Ceraclea albimacula

r~-

C.fulva

C. nigronervosa

_ J

L -

124

The Naturalist 140 (2015)

Mystacides azurea

~r~ f~! T~

M. longicornis

M. nigra

r i r T

Totals

17 Jj>

14 3 11 2

(? Indicates tentative identification; + indicates species recorded outside the current NNR;
indicates single specimen)

Discussion

Of the 29 species listed in 1963 and not recorded by us, 11 occurred outside the current reserve
area and 8 were only recorded as single specimens. We have recorded 8 species which were not
listed in 1963, bringing the total number of species for the area as a whole to 79, with 68 species
being recorded within the current NNR. Thus more than a third of the 196 currently recognised
British species have now been recorded from the NNR itself.

Adults of 44 species were recorded during 2013, two of which (Agrypnetes crossicornis and
Agroyleo multipunctata) were only found by hand searching along the shore of the tarn. The
Malaise trap produced the largest number (20 species) though this was slightly less than half the
total recorded. The Light trap and sweep net sampling produced very similar numbers (17 and
15) though only four species were taken by both methods (only one species, Cinnamon Sedge
Limnephilus lunotus, was taken by all three methods). Thus the combined total for sweep
netting and light trapping was just over half of the total number of adults recorded showing that
it is necessary to use several methods in parallel to obtain anything approaching a full species
list and even then some species were missed as adults which were found as larvae. For all
except one of the species the period of adult activity was as expected. Only in the case of A.
crossicornis, see below, was adult activity observed outside previously published 'flight times'.

Larvae of 23 species were recorded during 2013; 14 of them in Malham Tarn itself, of which 8
were not found elsewhere on the NNR. The small spring-fed stream entering the east shore of
Malham Tarn produced 2 species of Micropterno, both M. lateralis (which was only found here)
and M. sequax (which also occurred in the fen runnel). The runnels carrying water through the
fen and into Malham Tarn produced 11 species, 4 of which were only found here and 1, (M.
sequax) was also found in the small stream on the East shore of the tarn; the other 6 species
were not confined to running water and were also found in Malham Tarn itself. Only 3 species
were recorded from the ponds on the fen, none of which were found elsewhere.

Examination of the Chara sp. (Stonewort) beds in the tarn (in the search for Agrypnetes)
produced large numbers of larvae and pupae of Limnephilus politus. Juveniles of this species
were found nowhere else and the adults were found all along the northern shore of the tarn,
resting and mating on the ground vegetation, and spent females, with abundant egg masses,
were found among the mossy stones at the high water level all along the northern shore. Adults
of L. politus were caught in the light trap which indicates that they were attracted to the light
some distance from the tarn but they were not caught in the Malaise trap on the fen. Another
abundant species in the tarn was Welshman's Button Sericostoma personatum whose larval
cases were found all round the edges of the tarn and in the Chara beds. The adults of this
species are day flyers and were to be seen all over the NNR from mid-June to mid-July flying in

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125

hot, bright, sunshine. Mating pairs of S. personatum were found sitting on the upper surfaces of
leaves in bright sunshine and were also taken in flight.

Adults of the Hydroptillid Agroylea multipunctota were present in enormous numbers and were
collected by 'pootering' directly off the rocks around the tarn shore, where they were sitting and
running in the sunshine. Final instar larvae, in their cases, were found among the Choro and also
on the leaves of both Curled Pondweed Potamogeton crispus, growing just off shore in Boat
House Bay, and Shining Pondweed P lucens, a large stand of which grows out in the middle of
the tarn. They were also taken by pond netting in Ha Mire Bay and we sometimes found them
attached to the (occupied!) cases of L. politus during grapnel sampling. Between mid-June and
mid-August males and females of Oxyethira flavicornis, another Hydroptilid, were taken in the
Malaise trap. Larvae identified as Oxyethira sp. (probably O. flavicornis) were found in the tarn,
attached to both Potamogeton spp. and by pond netting in Ha Mire Bay where some of them
were attached to (occupied!) cases of L. lunatus, but we did not find the adults here.

A common species on the fen was Chaetopteryx villosa whose larvae live in stony cases in the
fen runnels and adults of which were found by beating and sweeping along the fen runnels
during October when it was also found in the Malaise trap.

The fact that some species are only known on the NNR from single adult records does not
necessarily mean that they are vagrant individuals and do not breed here. Hydatophylax
infumatus , for example, is included in the 1963 list on the basis of a single adult male attracted
to a Tilley lamp on the tarn shore at the inflow and we know of no subsequent records. We have
not seen any adults but we found two larvae in the main runnel through the fen not far from
where that adult was taken. A viable, though possibly small, breeding population must therefore
exist here, possibly confined to this small area.

An abundant species round the tarn is Grouse Wing Mystacides longicornis, adults of which can
be found resting by day on fences and among the ground vegetation. They start to become
active in the early evening and large swarms, of males and females, can be seen flying as the
light begins to fade and until well into the twilight. Mating pairs can be seen flying and on the
leaves of the vegetation near the tarn shore; they can even be seen running on the water
surface in calm conditions. Larvae are abundant in the tarn and were found by grapnel sampling
the Chara.

The Malham Sedge

Grapnel sampling from a boat showed that the Chara, with which the larvae of Agrypnetes
crassicornis are associated, and from which the early instars construct their cases (large larvae
construct cases of a variety of materials) was widespread but patchy in Malham Tarn. We
examined Chara from many parts of the tarn but we were only able to find juveniles of A.
crassicornis in one small area; offshore near the East Boathouse. Three well grown larvae were
seen on the 18 June and a dead pupa (in its larval case) on the 28 August. We had also found a
live pupa in this same area of the tarn on 7 September 2012. Only a single adult specimen was
seen, on the evening of 6 August, when hand searching among rocks on the shore of the tarn in
this same area produced a female specimen. This record extends the known period of adult
activity into early August (previously it had been recorded between 30 June and 25 July) and the

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record of a pupa in the previous September indicates the possibility that adult activity might be
even further extended. Adults of this species are regarded as flightless and have been observed
running across the surface of the tarn; it is interesting therefore that this specimen was
observed to flutter a distance of at least 10cm between the tops of two adjacent rocks before
scuttling down into another, narrower, gap. A search for eggs among the rocks just offshore was
unsuccessful but it appears that this population is still surviving even if its numbers are low. The
records from the 1950s indicate that adult numbers, then, fluctuated from year to year. In the
1963 report P. F. Holmes said of A. crassicornis must be some sort of relict species here/7; it
has since been found, as a sub-fossil in peat deposits some 10 to 15 thousand years old, at two
widely separated localities (I. D. Wallace, pers. com.) so it appears that he was right.

A. crassicornis is not the only member of the family Phryganeidae in the tarn. Several adults of
Great Red Sedge Phryganea bipunctata were seen on the north shore of the tarn and very large
numbers of pupal exuviae, which we assume to be this species but which it was not possible to
confirm the identity of (other than that they were Phryganeidae but not Agrypnetes), were
found along the western shore at the same time as the adults were flying (mid- to late-June).
The adults were flying at dusk and it was noticeable that fish were rising to feed at the surface of
the tarn at the same time. No larvae were seen though they must have been present in
considerable numbers. Phryganeid egg loops were seen among the rocks off the western shore
at the same place as the pupal exuviae were found and we assume that these were also P.
bipunctata; though the only way to be sure would have been to collect and hatch them, rearing
the larvae to sufficient size to be identifiable.

Acknowledgements

We thank the National Trust and English Nature for permission to collect specimens on the
Malham Tarn NNR. We thank all the staff of the Malham Tarn Field Studies Centre for their
enthusiastic support and hospitality, and the use of their boat and other equipment. We thank
Robin Sutton for his light trap caddis captures. We thank the YNU for the use of the Malaise
trap.

References

Barnard, P. & Ross, E. (2012). The adult Trichoptera (caddisflies) of Britain and Ireland.

Handbooks for the Identification of British Insects 1 (17): iv + 192 pp.

Edington, J. M. & Hildrew, A. G. (1995). A revised key to the caseless caddis larvae of the British
Isles with notes on their ecology. Scientific Publications of the Freshwater Biological
Association 53: 1-134.

Henson, H. (1963). The Insects of the Malham Tarn Area. Proceedings of the Leeds Philosophical
and Literary Society: Scientific Section 9: ii, 15-91.

Jones, C. (2001). Malham Tarn National Nature Reserve. British Wildlife 13: 29-37.

Macan, T. T. (1973). A key to the adults of the British Trichoptera. Scientific Publications of the
Freshwater Biological Association 28: 1-151.

Wallace, I. D., Wallace, B. & Philipson, G. N. (2003). Keys to the case-bearing caddis larvae of
Britain and Ireland. Scientific Publications of the Freshwater Biological Association 61:
1-259.

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127

Notes on the dolichopodid flies of two contrasting Yorkshire
bogs

Roy Crossley

1 The Cloisters, Wilberfoss, York Y041 5RF
Email: roycrossley@btinternet.com

Askham Bog near York (SE575481, VC64) is a 44.1ha remnant of a post-glacial mire at an altitude
of c.lOm which formed behind a terminal moraine to the south, along whose summit now runs
the dual carriageway of the A64 York by-pass. It has long been known as a haunt of rare fenland
plants and insects, to the extent that in the past it was called 'the Wicken Fen of the North'.
There is an account of a visit by boys of York Quaker (now Bootham) School in 1834 in search of
water beetles and plants (Fitter & Smith, 1979), and in 1946 the site became the first, and for
almost ten years the only, nature reserve of the Yorkshire Naturalists' Trust (YNT), as the
Yorkshire Wildlife Trust (YWT) was then known.

Fen Bog (SE857982, VC62) is a valley mire of some 18.3ha at the head of the glacial drainage
channel of Newton Dale in the heart of the North York Moors National Park at an altitude of
c.l50m. It was gifted to the (YNT) in 1964 and it was well known amongst entomologists as a
site for a variety of peat-bog insects, including the Large Heath Coenonympho tullio.

Although both sites are called 'Bog' they differ markedly in structure: Askham Bog is mainly an
alkaline-neutral fen whereas Fen Bog is mostly waterlogged acid peat. The vegetation of the two
reserves is quite different but both are superb examples of their types.

These differences are also reflected in the dipterous fauna, of which the Dolichopodidae is an
important component. The combined species-list for the two sites numbers 83, of which only 29
are common to both. The total number for Askham Bog now stands at 61, of which 32 have not
been recorded from Fen Bog. The total list for the latter numbers 51, of which 22 have not been
recorded at Askham Bog.

The 29 species recorded at both sites are mostly widespread and common generalists. Examples
are: Chrysotus gromineus, Dolichopus plumipes, D. po pul oris, D. ungu lotus, Gymnopternus
oerosus, Sybistromo obscurellus, Compsicnemus curvipes, C. scombus, Sympycnus desoutteri and
Syntormon pollipes.

Dolichopodids recorded from Askham Bog only include several which are restricted in their
Yorkshire distribution. Noteworthy amongst these, in a regional context, are Argyro elongoto,
Diophorus oculotus, Dolichopus wohlbergi, Ethiromyio cholybeo, Gymnopternus ongustifrons,
Rhophium fosciotum and Lomprochromus bifosciatus.

Argyro elongoto is little known in Yorkshire: first recorded from a water trap on West Fen,
Malham (VC64) on 11/9/1980, the next was a single female at Askham Bog on 16/7/1985 (this
specimen, much damaged but still identifiable, is in my collection). Much subsequent collecting

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at the site has failed to locate any more examples. A third and more recent record by Andrew
Godfrey from Inkle Moor, Thorne (VC63) on 25/6/2012 completes the tally. I have taken single
specimens at Loch Morlich (Aviemore) and Fenn's/Whixhall Moss (Shropshire), so this
dolichopodid is widely spread nationally but apparently always scarce.

Diophorus oculatus was reported by Chris Cheetham from 'Austwick. Lawkland1 (VC64) in a note,
according to his record card, published in The Naturalist in 1919. This remained the only known
Yorkshire locality until 1985 when it was recorded at Askham Bog, at which site it has
subsequently been found frequently in the fen meadows. It has also been found at the YWT
reserves at Ashberry (VC62) and Upper Dunsforth (VC61) and also at Sand Dale on the southern
edge of Dalby Forest (VC62), and in 2008 at Ellington Banks near Ripon (VC64). These widely
scattered records over the past thirty years doubtless reflect the mobility of the present
generation of recorders and an increasing interest in dipterology. The eyes of the males in life
are of the most beautiful blue, hence the name.

Dolichopus wahlbergi is very similar in general appearance to D. plumipes , one of the most
common and widespread of dolichopodids. The males of both have distinctive pennate lateral
fringes to the basal segments of the middle tarsi and the hind margins of the wings are
conspicuously sinuate towards the bases. These are thought to be male secondary sexual
characters, and leg adornments in particular occur in a number of dolichopodid genera,
especially the species-rich Dolichopus. D. wahlbergi is found less frequently than D. plumipes,
usually singly or in very small numbers, and it appears to be absent from much of the upland
western parts of Yorkshire.

Ethiromyia chalybea was first recorded in Yorkshire in 1987 from the towpath of Doncaster
Canal and thereafter from Gypsy Marsh, a wetland site in the south of VC63, in 1992 and 1997
and Holbrook Marsh in 1993. In 1996 a single female was found in a fen bordering Hornsea
Mere (VC61) and subsequently at half a dozen further sites, mostly in VC61. The first to be
recorded at Askham Bog was in 2010 and further examples were found in the same area of carr-
woodland near the entrance to the reserve in 2014.

Gymnopternus angustifrons (Lower Risk = Nationally Notable) is a tiny black fly for which there is
a record from Askham Bog (date unclear but pre-1953). That remained the sole record until
1971, when it was found at Hotham Carrs (VC61). The next was in 1984 from Askham Bog,
where it still occurs, and in the same year from Allerthorpe Common (VC61). Thereafter it has
been reported from several other damp, peaty sites in lowland Yorkshire: Skipwith (VC61) and
Strensall (VC62) Commons, Thorne Moors and sites in the Lower Derwent Valley (VC61).

Rhaphium fasciatum has a long association with Askham Bog, the first record being in 1953. This
was the only known Yorkshire site until it was found during the Malham survey undertaken by
the Entomological Section of the YNU between 1954/58 (Henson, 1963). Since then it has been
recorded from about a dozen widely scattered localities in the county but surprisingly from only
one in VC63: Inkle Moor, 2012. At Askham Bog several specimens of both sexes were found in a
small area of carr-woodland near the entrance to the Reserve between 1-19 May 2014.

Lamprochromus bifasciatus was first recorded in Yorkshire in 1996, when a single male was
found at Sand Dale, and there have been several subsequent records from the same locality.

The Naturalist 140 (2015)

129

There was a further VC62 record from Ashberry in 2008 and then from Askham Bog in 2013 and
2014.

Dolichopodids reported from Fen Bog but not Askham Bog include Diophorus nigricans,
Dolichopus longitarsis, Tachytrechus consobrinus, Hydrophorus albiceps, Schoenophilus versutus,
Rhaphium longicorne and Syntormon zelleri.

Diaphorus nigricans was first reported in Yorkshire from Thorne Moors in 1975. Since then it has
been found at about a dozen widely scattered sites in the county, mainly, but not exclusively, on
peat in both lowland and upland localities.

Dolichopus longitarsis records are mostly from upland localities, especially calcareous sites on
the North York Moors (VC62). There are three known Pennine sites and an isolated coastal one
from Easington (VC61). One of the Pennine sites (Bingley Bog) is the only one for VC63 and there
are no records from VC65.

Tachytrechus consobrinus (Lower Risk = Nationally Notable) was first recorded in Yorkshire at
Fen Bog in 2002 and it has been found on several subsequent occasions at this site. Elsewhere
on the North York Moors it has been found at Tranmire Bog and Bonfield Gill. The only other
Yorkshire record is from the shore of a sandy lagoon on Hatfield Moor (VC63) in 2004.

Hydrophorus albiceps was first recorded in Yorkshire at Helwith Moss, Austwick in 1921, and
subsequently there have been numerous reports from Pennine localities including Warley Moor
Reservoir and Studley Pike (both in VC63). There are two further sites on the North York Moors
in addition to Fen Bog - Tranmire Bog and Bonfield Gill. The majority of localities are acid bogs
dominated by Sphagnum spp. As is the case with some other insects once regarded as being
restricted to upland bogs, H. albiceps also occurs in similar situations in the lowlands, and there
are records from Skipwith Common and Thorne Moors; specimens were reported as being
'abundant' at the latter site in 1969.

Schoenophilus versutus is a tiny (2mm) dull grey fly of undistinguished appearance which can
easily be overlooked in the field. The first Yorkshire record was in 1948 at Gristhorpe Bay (VC62)
on the coast south of Scarborough and then followed reports from Great Close Mire and Ha
Mire in the 1954/8 survey at Malham (Henson, loc. cit.). There was a further record for the
former site at Malham in 1993. The only other reported County site is Fen Bog, where single
specimens were found in July 2013 and 2014.

Rhaphium longicorne is a well-known member of the dipterous fauna of peat bogs, mainly in the
uplands but also in similar lowland places (Crossley, 2014), and this spectacular dolichopodid is
frequent at Fen Bog.

Syntormon zelleri has been known in Yorkshire since first being recorded in 1982 at Catcliffe
Flash (VC63). Three years later it was found at Sug Marsh at Timble (VC64), and since then there
have been reports from five sites on the North York Moors in addition to Fen Bog, where it was
first recorded in 1988. There is a recent (2009) lowland record from the tiny conservation area at
the Yorkshire Air Museum, Elvington (VC61) near York. This is only c.lOkm from Askham Bog and
the fly may well be there, awaiting discovery!

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In the foregoing account nomenclature follows Chandler (1998 with updates) and threat
statuses are in Falk & Crossley (2005).

Acknowledgements

As always, a huge debt is owed to past and present entomologists who have given of their time
to submit, and in some cases to maintain, the records of the Yorkshire Naturalists' Union,
without whose efforts articles such as this could not be written. It is my privilege to maintain the
dolichopodid records at the present time and full details concerning any species noted in this
paper will gladly be provided on application.

All records pertaining to reserves of the Yorkshire Wildlife Trust are deposited with the Trust,
and it is a pleasure to record my thanks for the support received from Officers of the Trust in this
and other studies over more years than I now care to remember!

References

Chandler, P. J. (1998) Checklists of Insects of the British Isles (New Series), Part 1: Diptera.
Hondbk.ldent. Br. Insects 12 (1): 1-234.

Crossley, R. (2014) Notes on the distribution and habitat associations of dolichopodid flies in
Yorkshire. The Naturalist 139: 108-111.

Falk, S. J. & Crossley, R. (2005) A review of the scarce and threatened flies of Great Britain. Part
3: Empidoidea. Species Status 3: 1-134. JNCC. Peterborough.

Fitter, A. FI. & Smith, C. J. (eds) (1979) A Wood in Ascam- A Study in Wetland Conservation, v.-viii.

1-164. Wm. Sessions, York, and Yorkshire Naturalists' Trust.

Flenson, H. (1963). The Insects of the Malham Tarn Area. Proceedings of the Leeds Philosophical
and Literary Society: Scientific Section 9: ii, 15-91.

Erratum

The Spurn Bird Observatory was established in 1946 under the auspices of the Yorkshire
Naturalists' Union at Warren Cottage on the Spurn peninsula. It is now a separate trust. The
Yorkshire Wildlife Trust (then the Yorkshire Naturalists' Trust) bought Spurn from the Ministry of
Defence in 1959 but did not create the bird observatory as I erroneously stated on p.36 of my
recent article (Moore 2015). I thank Jan Crowther and Barry Spence for bringing this to my
attention.

P.G. Moore

Reference

Moore, P. G., (2015). Michael Clegg DSc (Hon.), FMA, MBOU (1933-1995): a biography and
bibliography. The Naturalist 140: 33-40.

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131

Geological and land use influences on Badger sett densities
across South Yorkshire

Colin Howes

7 Aldcliffe Crescent, Doncaster DN4 9DS
Email: colinhowes@blueyonder.co.uk

Introduction

Badgers Meles meles generally prefer to excavate their setts into a steep or even vertical surface
(Paget & Middleton 1969, 1974); it is therefore understandable that their distribution should
reflect the prevailing topography and geology. From data available in the archived Mammal
Society Yorkshire Badger Sett Survey files and data from Mike Dyson, one-time member of the
South Yorkshire Badger Group, it has been possible to examine the geographical distribution of
208 setts known to be active from 1974 to 1988 across the Metropolitan County of South
Yorkshire. This has revealed an intriguing association with the exposed solid geology, the history
of human exploitation of mineral resources, the management of arable agricultural landscapes
across the Holocene drift geology and the fluctuating performance of the Sherwood sandstone
aquifer in the east of the region.

By counting the numbers of 1 km squares of the Ordnance Survey National Grid north and south
along a line of 64 1 km squares from west to east of the Metropolitan County, those squares
known to contain setts were expressed as a percentage of the total at each easting. By plotting
the percentage scores along the line of the 64 eastings, a pattern of sett preference was
revealed in Figure 1.

Examination of the solid and drift geological maps of the British Geological Survey for the
Sheffield, Barnsley, Doncaster and Goole regions has shown a rationale for the uneven
distribution. Geological divisions which run conveniently in five belts from west to east,
commence with the Carboniferous Namurian Millstone Grit, the Westphalian Lower coal
measures, the Middle coal measures, often overlain by alluvium of the Don and Dearne valleys,
the Permian Magnesian Limestone and marl ridge, the Triassic Sherwood sandstones overlain by
the 25 ft drift of the bed of the post-glacial Lake Humber and by deposits of morainic and glacio-
fluvial debris.

The tough Millstone Grits of the Pennine Peak District (between OS easting 12 and 18) are
occupied, but relatively sparsely compared with the Westphalian Lower coal measures (within
OS eastings 19 to 32) which provide more friable strata including sandstones, coal, shale and
clay. These form the 'exposed' coalfield, where strata have been worked at the surface. Over the
centuries, the landscape has been pitted by hundreds of disused shallow mine workings
(including large numbers of 'bell pits', now abandoned and wooded), quarries and clay pits. This
industrial heritage has provided numerous artificially excavated structures which badgers have
taken over and developed as sett systems. These occupied sites are particularly resistant to
illegal badger digging and are therefore disproportionately represented across South Yorkshire
(Mike Dyson pers. comm.).

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The Noturolist 140 (2015)

OS Eastings

Figure 1: Relative frequencies of Badger setts across the geological zones of South Yorkshire
(1974-1988).

The alluvial river washland landscapes of the Don and Dearne valleys overlie parts of the
Westphalian Middle Coal measures (within OS eastings 33 to 44) to the west of the Permian
Magnesian Limestone ridge. This was commercially exploited by deep mine technology offering
fewer opportunities for sett excavation and those that are present in the landscape are subject
to high levels of illegal badger digging.

Figure 2. A badger sett under the Permian limestone, which is visible
immediately above the entrance. P Simmons

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133

A second favoured area for sett creation, from OS eastings 45 to 58, is provided by the Permian
limestone and marl ridge. The underlying basal Permian sand, revealed largely along the
western escarpment of the ridge, is particularly favoured by badgers for the excavation of setts
(Fig. 2), as are the numerous fissures and crevices exposed by quarries and disused railway
cuttings.

Across the low-lying bed of the former Lake Humber, from OS eastings 59 to 75, setts are largely
absent. This is probably due to a relatively high water table and a notably flat landscape. Islands
of occurrence do occur in a series of slightly elevated areas of moraine and glacio-fluvial erratic
debris, between OS eastings 69 and 72 as at Lindholme Island on Hatfield Chase. Human
constructions in the form of flood embankments and the faces of networks of Internal Drainage
Board ditches and drains are increasingly being utilised (Mike Dyson pers. comm.). Exploitation
of these sites appears to be a post 1970s phenomenon, coinciding with the lowering water
table, through agricultural irrigation and over-exploitation for public supply of the underlying
Sherwood sandstone aquifer.

Acknowledgements

I am greatly indebted to Ron Deaton of the Harrogate Naturalists' Society for making available
the Mammal Society Yorkshire Badger Sett Survey files and Mike Dyson for his exhaustive
knowledge of former badger setts across South Yorkshire.

References

Ordnance Survey Londronger Series 1:50,000 scale sheets 109; 110; 111; 112; 118; 119 and 120.
Paget, R.J. and Middleton, A.L.V. (1969) National Badger Survey. The Naturalist 94: 81-82.

Paget, R.J. and Middleton, A.L.V. (1974) Badgers of Yorkshire and Humberside. Ebor Press, York.

Increase in bog-mosses Sphagnum and other changes in the
vegetation of Ringinglow Bog (Southern Pennines) since the
1940s

R. Goulder

5 Bishops Croft, Beverley HU 17 8JY
Email: r.goulder@hull.ac.uk

Introduction

It is well known that bog-mosses typically thrive on cold, nutrient-poor upland bogs where they
can out-compete vascular plants and are responsible for the bulk of peat deposition (e.g. van
Breemen, 1995). By the mid-20th century it was recognized that the blanket bogs of the
Southern Pennines were atypical: bog surfaces were often dominated by cotton-grasses

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(Common Cottongrass4 and/or Hare's-tail Cottongrass) while bog-mosses were few or absent
(e.g. Pearsall, 1950; Anderson & Shimwell, 1981). Nevertheless, studies of stratigraphy showed
that much of the peat deposits underlying the current vegetation had been formed from bog-
mosses (Tallis, 1964) while Pearsall (loc.cit.) reported an account of upland blanket bog in the
area with great abundance of bog-mosses as recently as 1813. Potential reasons for the loss of
bog-mosses include changes in drainage, grazing pressure and moor-burning regimes but the
likely cause that has been most stressed is deposition of atmospheric industrial pollutants. The
industrial conurbations of Sheffield and Manchester are close by and Tallis (1965, 1985) showed
that blanket bog in the Kinder-Bleaklow area had surface deposits, <10-cm deep, of soot-
contaminated humified peat overlying peat that was formed from bog-mosses; the discontinuity
appears to have occurred since about 1800 and is aligned with the development of
manufacturing industry and a general increase in coal burning.

Experimental evidence supports the hypothesis that atmospheric pollution has been responsible
for loss of bog-mosses. For example, Ferguson et al. (1978) used laboratory experiments to
demonstrate inhibition of extension growth, chlorophyll synthesis and photosynthesis in diverse
bog-mosses at concentrations of bisulphite (HS03_) and sulphate (S04_ -) that were
commensurate with concentrations in Sheffield and Manchester rain in the 1970s. Subsequent
field experiments (Ferguson & Lee, 1980) showed that bog-mosses at an unpolluted site in
North Wales were inhibited by application of artificial rain that contained bisulphite and
sulphate at 1970s Sheffield-Manchester-region concentrations. It has also been suggested that
high concentrations of combined inorganic nitrogen in acid deposition may disrupt nitrogen
metabolism and retention by bog-mosses (Press & Lee, 1982; Press et al., 1986) and lead to
disruption of the nitrogen regime of bog surfaces and upset of interactions between bog-mosses
and vascular plants, to the detriment of the bog-mosses (Woodin & Lee, 1987; Gunnarsson et
al., 2004).

This article is concerned with an area of blanket bog in the Southern Pennines that was given
the name "Ringinglow Bog" by Conway (1947). Ringinglow Bog forms part of the Eastern Peak
District Moors SSSI and lies at altitude c.400m within National Grid monads (1km x 1km squares)
SK2583, 2683, 2783, 2584, 2684, 2784 and 2682; Upper Burbage Bridge (SK261830) is at the
south-west corner of the bog. The bog is bordered to the south by the Hathersage-Ringinglow
road, from which it extends northwards for c. 1500m, and to the west by Burbage Brook, from
which it extends eastwards for c. 1400m. Conway (loc.cit.) summarized the vegetation of the
bog: she described Heather, Common Cottongrass and Hare's-tail Cottongrass as dominant/co-
dominant, Wavy Hair-grass as widespread and Soft-rush as locally dominant. Bog-mosses were
not conspicuous, with the exception of a wetter area towards the north-western part of the bog
where Flat-topped Bog-moss was abundant amongst the cotton-grasses. She stressed, however,
that unconsolidated remains of bog-mosses are revealed over much of the bog if the superficial
5cm or so of deposit is scraped away and suggested that the dominance of bog-mosses was lost
perhaps about 100 years before this 1940s study. She acknowledged the possible role in this of
atmospheric pollution but concluded that cutting of drainage channels was a more likely cause.

4 Scientific names which are included in the tables are not repeated in the text. Nomenclature of
vascular plants follows Stace (2010) and that of bryophytes Atherton, Bosanquet & Lawley (2010).

The Naturalist 140 (2015)

135

A view more aligned to recent work, reviewed above, is that deposition from atmospheric
pollution was probably the key factor at Ringinglow Bog. Evidence of extreme deposition of
industrial pollutants specific to Ringinglow Bog is provided by metal concentrations in the peat
(e.g. cadmium, copper, iron and lead) that greatly exceed those at unpolluted bog sites (Markert
& Thornton, 1990; Jones & Hao, 1993; Gao et al., 1999).

Vegetation of the main flush area of Ringinglow Bog in the 1940s and 2014

Conway (1949) described the vegetation of Ringinglow Bog as it was in the 1940s. Part of that
study focussed on one of the wetter parts of the site that was referred to as the "main flush".
This lay in the central and eastern part of the bog and, from the sketch map provided had an
area of c.31ha. Conway emphasized that the term 'flush' was used to describe an area in which
surface water accumulated and that its use did not imply that there was upwelling of inorganic
nutrients. She described two vegetation types within the main flush: (1) an area with much
Common Cottongrass (Plate Ilia, centre pages) along with dominant Hare's-tail Cottongrass
(Plate lllc, centre pages) found in the central area of the main flush; (2) wide bands to the north
and south of the central area dominated by Hare's-tail Cottongrass but where Wavy Hair-grass
was also important.

Conway recorded plants in lm2 quadrats at intervals of 10 paces along a c.HOOm transect A-B
that crossed the main flush, taking in both vegetation types, and extended onto Heather-
dominated heath beyond. Her Table 1 includes the percentage frequency for occurrence of each
plant in 1-m2 quadrats in both vegetation types of the main flush. The site was species poor;
only ten plants (five vascular plants and five bryophytes) were recorded in 20 quadrats placed in
the Common Cottongrass-Hare's-tail Cottongrass vegetation type, the most frequent vascular
plants were Hare's-tail Cottongrass 100%, Common Cottongrass 45% and Wavy Hair-grass 15%.
Thirteen (seven angiosperms and six bryophytes) were found in 25 quadrats5 placed in the
Hare's-tail Cottongrass-Wavy Hair-grass vegetation type, including Hare's-tail Cottongrass 100%,
Wavy Hair-grass 100%, Common Cottongrass 52%, Crowberry 24% and Cranberry 24%. Bog-
mosses were not recorded in any of the quadrats placed on the main flush.

The author of the present article participated in an undergraduate field course to the site in May
1963 (led by Dr D. J. Boatman); my notes record abundant cotton-grasses with bog-mosses
confined to ditches. A more recent visit to the main flush area was made in September 2014.
The zonation of vegetation observed by Conway was in general terms still discernible from the
Hathersage-Ringinglow road; the reddish leaves of Common Cottongrass gave the vegetation
that it dominated a reddish tinge, making it clearly distinguishable from the greener Hare's-tail
Cottongrass-dominated vegetation. Also, it was observed while walking approximately
northwards from the road (beginning at SK26938307) that an initial Heather-dominated slope
gave way after c.80m to Hare's-tail Cottongrass-dominated tussocky blanket bog with Wavy
Hair-grass. There were abundant bog-mosses (Flat-topped and Fringed Bog-mosses) amongst
the tussocks (Plate Hid, centre pages). At c.l30m from the road the vegetation changed to
Common Cottongrass dominance with Hare's-tail Cottongrass and Wavy Hair-grass. Bog-mosses

5 The relevant column in Conway’s Table 1 states that percentage frequencies were determined from
20 quadrats but the values given are all integers that are multiples of four hence it is likely that they
are from 25 quadrats.

136

The Naturalist 140 (2015)

were also present including Flat-topped and Fringed Bog-mosses. Also present were red plants
from Sphagnum section Acutifolia. During a subsequent visit in February 2015 it became
apparent that these comprised Lustrous Bog-moss, identifiable in the field by red capitula with
green centres, and Acute-leaved Bog-moss. At c.730m from the road the Common Cottongrass
was lost and Hare's-tail Cottongrass became dominant again, with Wavy Hair-grass and some
Purple Moor-grass; bog-mosses were abundant. At c.800m Heather again became dominant. An
elongated pool orientated about east-west was found to the south-east of the main flush; it
contained abundant Feathery Bog-moss with Fringed Bog-moss and Common Cottongrass (Plate
lllb, centre pages). This was presumably a remnant of the ditch complex shown on Conway's
sketch map.

The plants found in the main flush area in September 2014 are listed in Table 1; also included
are those found in quadrats on the main flush by Conway in the 1940s (taken from her Table 1).
The principal change since the 1940s is clearly the reappearance of Sphagnum. The vascular
plants found in 2014 were much the same as in the 1940s; all seven of those recorded in the
1940s were still there in 2014.

More systematic recording of the vegetation of the main flush was undertaken in November-
December 2014. Plants were recorded in 1-m2 quadrats using the Braun-Blanquet abundance
scale (Kent, 2012); i.e. + = sparse, 1 = common but <5% cover, 2 = 5-25% cover, 3 = 25-50% cover,
4 = 50-75% cover and 5 = >75% cover. Acute-leaved and Lustrous Bog-mosses were not
separated; liverworts, always inconspicuous, were not included in the recording. Quadrats were
located, so far as possible, as described by Conway; that is, they were placed at 10-pace
intervals along a transect from the Hathersage-Ringinglow road that began at SK27148316 and
crossed the main flush at an alignment 40 degrees west of north. Twenty quadrats were placed
across the central area of the main flush (between c.l90m and 340m from the road) in
vegetation that was perceived to be dominated by Common Cottongrass, and 20 quadrats were
placed in the zone between the marginal heath and the central area of the main flush (between
c.50m and 190m from the road) in vegetation that was perceived to be dominated by Hare's-tail
Cottongrass with Wavy Hair-grass (two of these latter quadrats were placed 15m west of the
transect).

The aims of the recording were:

• To test whether the subjective discernment of two vegetation types in the main flush is
supported by an objective analysis of records. To this end the Braun-Blanquet scores for
each species in each quadrat were re-coded and simplified; i.e. not recorded = 0; + & 1 =
1; 2 & 3 = 2; 4 & 5 = 3. The re-coded scores were used to compare quadrats on the basis of
species present and their abundance using de-trended correspondence analysis
(DECORANA) (Kent, 2012); Community Analysis Package 3.0 was used (Henderson &
Seaby, 2008).

• To look for statistically significant change since the 1940s in the percentage frequency
of each species recorded, within both vegetation types in the main flush. Analysis of
association (Campbell, 1967) was used to achieve this; a 2 x 2 contingency table was
prepared for observed frequencies (number of quadrats with or without the species) in

The Naturalist 140 (2015)

137

the 1940s and 2014 and another for expected frequencies. The 1940s data were taken
from Table 1 of Conway (1949). This was done for each species and each of the two
vegetation types; a two-tailed chi-square test was used to look for significant differences
between frequencies in the 1940s and 2014.

The Braun-Blanquet scores for 20 quadrats in the Common Cottongrass-dominated vegetation
towards the centre of the main flush are available as additional electronic material (Appendix
la). The scores for some quadrats suggest more than 100% cover but this is because of overlap
of species. Nine vascular plants were recorded. The dominant Common Cottongrass was present
in all quadrats and was recorded as 50-75% or >75% cover in 18 out of 20 quadrats. Hare's-tail
Cottongrass and Wavy Hair-grass were also recorded in all quadrats but with much lower
percentage cover; because recording was in December the Wavy Hair-grass foliage was largely
withered and dead. Cranberry was inconspicuous but was found in 18 quadrats; Cross-leaved
Heath was found in 11 quadrats and its cover was estimated as 25-50% in two of these. Bog-
mosses were recorded in 13 out of 20 quadrats; five of these records were for cover of 25-50%
or 50-75%. Species found were Acute-leaved/Lustrous, Feathery and Fringed Bog-mosses.
Bryophytes were otherwise largely inconspicuous; Heath Plait-moss was recorded in three
quadrats, Common Haircap in two quadrats (albeit at 25-50% in one of these) and Springy Turf-
moss in one quadrat.

The Braun-Blanquet scores for quadrats on the vegetation dominated by Hare's-tail Cottongrass
with Wavy Hair-grass are also available as additional electronic material (Appendix lb). Eight
vascular plants were recorded. The dominant Hare's-tail Cottongrass was in all 20 quadrats with
cover estimates of 25-50% in seven quadrats, 50-75% in eight quadrats and >75% in five
quadrats. Wavy Hair-grass was also in all quadrats, although largely withered and much less
conspicuous. Cranberry, although inconspicuous, was recorded in 16 quadrats; Common
Cottongrass was also frequently encountered being recorded in 14 quadrats, although with low
values (5-25% in one quadrat, otherwise <5% cover). Bog-mosses were recorded in 17 quadrats
and two species were found; Flat-topped and Fringed Bog-mosses. The latter was the most
abundant, being found in 14 quadrats and conspicuously abundant in some of them (25-50% in
three quadrats, 50-75% in two quadrats and >75% in one quadrat). Flat-topped Bog-moss was
found in seven quadrats, being notably abundant in a few of them (25-50% in one and 50-75%
in another). Otherwise, amongst bryophytes, Common Haircap was recorded in eight quadrats
(at 25-50% cover in two of these) while there were occasional records of Rusty Swan-neck Moss,
Silky Forklet-moss, Heath Plait-moss and Springy Turf-moss.

The DECORANA ordination plot for quadrats (Fig. 1) largely supported the initial subjective visual
categorization of the main flush vegetation into two distinct types. Although there is an element
of subjective judgement in the drawing of the cluster boundaries, it appears that 17 of the
quadrats (prefix a) from the Common Cottongrass-dominated area were in a cluster towards the
lower left of the plot while 15 quadrats (prefix v) from the Hare's-tail Cottongrass (with Wavy
Hair-grass) vegetation type occupied a cluster towards the upper right of the plot. There were
eight intermediate or outlying quadrats which is concomitant with the observed patchy nature
of the vegetation. The species plot is available as additional electronic material (Appendix 2).
This suggested that Common Cottongrass, Cross-leaved Heath, Acute-leaved/Lustrous and

138

The Naturalist 140 (2015)

Feathery Bog-mosses were important in pulling quadrats into the lower left cluster while Hare's-
tail Cottongrass, Flat-topped and Fringed Bog-mosses were important in pulling quadrats into
the upper right cluster.

Figure 1. Ringinglow Bog, main flush area, November-December 2014; DECORANA ordination
plot for quadrats. Quadrats al-20 are for the vegetation dominated by Common Cottongrass
with Hare's-tail Cottongrass ; quadrats vl-20 are for the vegetation dominated by Hare's-tail
Cottongrass with Wavy Flair-grass. Points for quadrats a5 & al7 are co-incident as are those for
quadrats al2 & al4. The scale on the axes indicates relative difference between the quadrats.

In the central area of the main flush, dominated by Common Cottongrass, this plant had
increased significantly since the 1940s to be present in 100% of quadrats in 2014 (Table 2, pl44).
Three other vascular plants had significantly increased in frequency (Wavy Hair-grass, Cross-
leaved Heath and Cranberry). Amongst bryophytes, bog-mosses (Acute-leaved Bog-
moss/Lustrous Bog-moss, Feathery Bog-moss and Fringed Bog-moss) had increased significantly
(from zero). Nodding Thread-moss had apparently been lost.

In the Hare's-tail Cottongrass (with Wavy Hair-grass)-dominated area of the main flush only
Cranberry had increased since the 1940s (Table 3, pl45); otherwise the vascular plants showed
no significant change in frequency. Amongst bryophytes, Flat-topped Bog-moss and Fringed Bog-
moss had increased significantly (from zero). Common Haircap had also increased significantly
while Nodding Thread-moss had apparently disappeared.

The Naturalist 140 (2015)

139

Discussion

The substantial increase in bog-mosses at Ringinglow Bog since the 1940s (Tables 2 & 3) can be
seen in the context of the report on the recent status of bog-mosses on the Peak District
moorlands that has been published by the Moors for the Future Partnership (Carroll et al.,
2009). That report accepted that gross sulphur dioxide and acid deposition from around the
1850s onward was the principal cause of the widespread disappearance of bog-mosses from the
region. The report also suggests that bog-mosses are now returning to these moorlands and
relates this to greatly reduced atmospheric pollution over the past 40 years, a reduction that is
part of a Europe-wide improvement; since 1980 total European land-based emissions (i.e. not
from shipping) of sulphur dioxide have fallen by 84% and those of nitrogen oxides by 46%
(Agren, 2013). Furthermore, Caporn et al. (2006) observed a marked increase of bog-mosses
between the early 1980s and 2005-6 on blanket bog at Holme Moss, also in the Southern
Pennines; this they concluded is likely to be due to amelioration in atmospheric pollution. Thus
the increase in bog-mosses at Ringinglow Bog is liable to be, at least in part, related to reduced
incidence of pollution even though it is likely that a legacy of pollutants remains in the peat.

The increase in bog-mosses in the main flush area of Ringinglow Bog has not taken place in a
landscape otherwise devoid of bog-mosses. Carroll et al. (loc.cit.) reviewed Peak District records
between 1989 and 2007 and showed that bog-mosses are widely distributed, albeit not
necessarily abundant. Twenty were found, the most frequently recorded being Flat-topped,
Fringed and Blunt-leaved Bog-mosses. Even in the 1940s bog-mosses were to be found at
Ringinglow Bog (Conway, 1949), although not recorded in quadrats across the main flush.
Conway recorded Flat-topped Bog-moss at frequencies of 75-85% in quadrats across an area of
c.l5ha that she called the "north-western flush". Blunt-leaved Bog-moss was present in one
quadrat; Fringed and Lustrous Bog-mosses were also in this part of the bog although not in the
quadrats and a patch of Papillose Bog-moss Sphagnum papillosum was found in the central part
of the bog.

As is emphasized by Carroll et al. (loc.cit.) there are other environmental and management
factors in addition to atmospheric pollution and its amelioration that might have contributed to
the loss and recovery of bog-mosses in the Southern Pennines; these include changes in erosion,
burning, grazing, trampling and drainage. Conway (1949) inferred that loss of bog mosses
through erosion at Ringinglow Bog was unimportant; erosion was proceeding only slowly, in
contrast with the summits of Kinder Scout and Bleaklow, and she attributed this to local
topography. Nor was erosion especially obvious in 2014. Conway (1949) mentions periodic
burning at Ringinglow Bog; no evidence of burning was observed in 2014. Conway also mentions
sheep on the bog. Sheep grazing generally in the Peak District increased substantially in the 20th
century; the number of sheep on the hills trebled between 1930 and 1976 (Anderson & Yalden,
1981). Since 2000, however, this trend has been reversed with numbers of breeding ewes in the
South Pennines decreasing by 3.6% between 2000 and 2010 (Silcock et al., 2012). I do not have
information about whether sheep numbers have decreased at Ringinglow Bog; there were
about ten sheep on the bog on 4 September 2014 and 23 on 26 February 2015. Yalden (2004)
found that slow regeneration of Flat-topped Bog-moss took place over the years 1980-2004
when sheep were excluded from eroded blanket bog in the Southern Pennines, with no other
treatment. He concluded that grazing and trampling had hitherto hindered regeneration.
Conway (1949) concluded that artificial drainage was important in the loss of bog-mosses at

140

The Naturalist 140 (2015)

Ringinglow. Since the 1940s the ditches that she described are likely to have become more
occluded, hence there may now be a higher water table. Furthermore, at least one watercourse
towards the west of the bog has been blocked by a series of dams, although it is not clear to me
whether this has raised water levels in the main flush area. Such gully blocking has recently
become a useful technique in the management of blanket bogs (Evans et a!., 2005).

The preliminary visit to Ringinglow Bog in September 2014 suggested that the vascular-plant
flora of the main flush area has not changed much since the 1940s because all seven of the
vascular plants found by Conway (1949) were still there (Table 1). Furthermore, the overview of
the site from the Hathersage-Ringinglow road suggested that Conway's separation of the main-
flush vegetation into (1) Common Cottongrass with Hare's-tail Cottongrass in the central area
and (2) Hare's-tail Cottongrass with Wavy hair-grass towards the margins of the flush, still holds
good. DECORANA (Fig. 1), using the data from quadrats collected in November and December
2014, confirmed that there genuinely are two distinct vegetation types in the main flush area,
albeit with some patchiness and overlap.

It seems, however, that there has been some recognizable change shown by vascular plants
since the 1940s. This is most obvious in the central area of the main flush, where Conway
described Common Cottongrass as abundant but Hare's-tail Cottongrass as dominant. Since then
Common Cottongrass has increased in frequency from 45% to 100% of quadrats. Moreover, the
Braun-Blanquet abundance scores for Common Cottongrass were always greater than those for
Hare's-tail Cottongrass (Appendix la). Common Cottongrass had clearly become dominant by
2014. Other evidence of change is provided by the significant increase in frequency of Wavy
Hair-grass, Cross-leaved Heath and Cranberry (Table 2, pl44). The possible changes in habitat
described above that might have led to the regeneration of bog-mosses (e.g. reduced pollution,
perhaps less grazing and trampling) are also potentially relevant to the increase in species
richness shown by vascular plants. Change has apparently been less in the Hare's-tail
Cottongrass-Wavy Hair-grass vegetation towards the margin of the main flush. Here there were
no significant changes in frequency except for Cranberry, which increased from 24% to 80% of
quadrats (Table 3, pl45).

Two vascular plants that were only occasionally seen in the main flush area in 2014 are also
worth comment. Firstly, Purple Moor-grass was noted in September 2014 (Table 1, pl43) and
was later recorded in one quadrat (Appendix la). This plant was not recorded in the main flush
by Conway (1949) and only sparingly in the north-western flush area. In contrast, by September
2014 there was extensive Purple Moor-grass in the north-western flush area with Hare's-tail
Cottongrass and Wavy Hair-grass. Vegetation dominated by Soft-rush observed by Conway in the
north-western flush persisted in 2014. Secondly, a sapling of Rowan was observed in the main
flush in September 2014 (Table 1, pl43) and another (height 76 cm) was later recorded in a
quadrat (Appendix la). This incipient colonization by trees might indicate reduction in grazing.

The decrease in frequency of Nodding Thread-moss since the 1940s was the only significant
change found for mosses other than increase in bog-mosses (Tables 2 & 3 ppl44, 145) and
Common Haircap (Table 3, pl45). Indeed, Nodding Thread-moss was not recorded in 2014;
Atherton et ol.( 2010) suggest that this species has declined generally in recent years.

The Naturalist 140 (2015)

141

Acknowledgements

I am grateful to Natural England and to land holders for permission to work on the bog and to

Professor G. W. Scott for his helpful comments on the manuscript.

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Table 1. Plants recorded in the main flush area of Ringinglow Bog in September 2014 and by
Conway in the 1940s

Recorded by Recorded September
Conway in the 2014

1940s*

Vascular plants

Calluna vulgaris Heather + +

Deschampsia flexuosa Wavy Hair-grass + +

Empetrum nigrum Crowberry + +

Erica tetralix Cross-leaved Heath + +

Eriophorum angustifolium Common Cottongrass + +

Eriophorum vaginatum Hare's-tail Cottongrass + +

Juncus effusus Soft-rush - +

Molinia caerulea Purple Moor-grass - +

Sorbus aucuparia Rowan (sapling) - +

Vaccinium myrtillus Bilberry - +

Vaccinium oxycoccos Cranberry + +

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143

Mosses

Compylopus flexuosus Rusty Swan-neck Moss + +

Compylopus introflexus Heath Star-moss - +

Dicranella heteromall a Silky Forklet-moss + +

Hypnum jutlandicum Heath Plait-moss - +

Pohlia nutans Nodding Thread-moss +

Polytrichum commune Common Haircap + +

Sphagnum capillifolium/subnitens - +

Acute-leaved/Lustrous Bog-mosses

Sphagnum cuspidatum Feathery Bog-moss - +

Sphagnum fallax Flat-topped Bog-moss - +

Sphagnum fimbriatum Fringed Bog-moss - +

Sphagnum palustre Blunt-leaved bog-moss - +

Liverworts

Calypogeia azurea Blue Pouch wort +

Cephalozia bicuspidata Two-horned Pincerwort + +

Gymnocolea inf lata Inflated Notchwort + +

(+) = present, (-) = not recorded. *Data from Table 1 of Conway (1949).

Table 2. Percentage frequency of plants in 1-m2 quadrats in the vegetation dominated by
Common Cottongrass with Hare's-tail Cottongrass in the main flush area of Ringinglow Bog in
the 1940s and in November-December 2014.

Percentage frequency

1940s

2014

Vascular plants

.unn,.,,,

Calluna vulgaris Heather

Deschampsia flexuosa Wavy Hair-grass

100

<0.01

Empetrum nigrum Crowberry

Erica tetralix Cross-leaved Heath

<0.01

Eriophorum angustifolium Common Cottongrass

100

<0.01

Eriophorum vaginatum Hare's-tail Cottongrass

100

Molinia caerulea Purple Moor-grass

Sorbus aucuparia Rowan (sapling)

Vaccinium oxycoccos Cranberry

<0.01

Mosses

Hypnum jutlandicum Heath Plait-moss

Pohlia nutans Nodding Thread-moss

<0.05

Polytrichum commune Common Haircap

Rhytidiadelphus squarrosus Springy Turf-moss

Sphagnum capillifolium/subnitens
Acute-leaved/Lustrous Bog-mosses

<0.01

Sphagnum cuspidatum Feathery Bog-moss

<0.05

Sphagnum fimbriatum Fringed Bog-moss

<0.05

1940s records are from Conway (1949). Values are derived from 20 quadrats; NS=P>0.05.

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Table 3. Percentage frequency of plants in 1-m2 quadrats in the vegetation dominated by Hare's-
tail Cottongrass with Wavy Hair-grass in the main flush area of Ringinglow Bog in the 1940s and
in November-December 2014.

Percentage frequency

1940s

2014 P

Vascular plants

Calluna vulgaris Heather

25 j

Deschampsia flexuosa Wavy Hair-grass

100

Empetrum nigrum Crowberry

Erica tetralix Cross-leaved Heath

Eriophorum angustifolium Common Cottongrass

Eriophorum vaginatum Hare's-tail Cottongrass

Too

100

Galium saxatile Heath Bedstraw

Vaccinium oxycoccos Cranberry

<0.01

Mosses

Campylopus flexuosus Rusty Swan-neck Moss

Dicranella heteromalla Silky Forklet-moss

Hypnum jutlandicum Heath Plait-moss

Pohlia nutans Nodding Thread-moss

<0.01

Polytrichum commune Common Haircap

<0.01

Rhytidiadelphus squarrosus Springy Turf-moss

Sphagnum fallax Flat-topped Bog-moss

<0.01

Sphagnum fimbriatum Fringed Bog-moss

<0.01

1940s records are from Conway (1949).

Values for the 1940s are from 25 quadrats while those for 2014 are from 20 quadrats;
NS=P>0.05.

Appendices la, lb and 2 can be downloaded from The Naturalist page of the YNU website .

Additions and corrections to the Yorkshire Diptera list (part 6)

Andrew Grayson

56 Piercy End, Kirkbymoorside, York, North Yorkshire, Y062 6DF

A small number of Diptera enthusiasts steadily continues to discover new flies in Yorkshire.
Some of them are within underworked and difficult groups and were previously overlooked
whereas others are recent arrivals to the county, e.g. the horsefly Tabanus autumnalis which
was found at Fairburn Ings during July 2014 (Brothers & Grayson, 2014). Many of the additions,
etc., given here are due to the work of Ian Andrews (IA), John Coldwell (JDC), Roy Crossley (RC)
and Bill Ely (WAE). Most additions result from Bill Ely's prolific and longstanding recording in the
Rotherham area. Ian Andrews' recent fieldwork at such regionally important sites as Allerthorpe
Common and Calley Heath has produced a good number of additions to the county and VC61
lists. John Coldwell continues to investigate underworked Diptera assemblages in the Barnsley

The Naturalist 140 (2015)

145

area, which has resulted in many additions to the county and VC63 lists over many years. Roy
Crossley has recently restricted his studies to Dolichopodidae, yet he continues to make new
and interesting discoveries.

An updated version [dated 22.1.2015] of A Simplified Provisional List of Yorkshire Diptera is now
available via the YNU's web-site. This list names various species as being 'excluded' from the
county list, or 'queried' - usually by the determiner. It is inevitable that many of these 'queried'
species would be excluded from any definitive county list, therefore the previous paper in this
series (Grayson, 2014) provisionally excluded many of them. The current paper continues that
process and also continues the policy of only including additions which have not been published
elsewhere.

Taking into account all adjustments due to additions, corrections, species lost to synonymy and
provisionally excluded, etc., the provisional Yorkshire Diptera list at 20.2.2015 contained 4,296
species, including 1,578 in the sub-order Nematocera. This is a net increase of 24 species to the
list at 3.3.2013 (Grayson, 2014). This modest increase would be far greater if so many 'queried'
species were not now considered 'provisionally excluded'. In the list below, (CJtf) or (99)
denotes that the precise number of males or females was not recorded (W.A. Ely, pers. Comm.).

Additions to Yorkshire Diptera List

CECIDOMYIIDAE

Obolodiplosis robinae (Haldeman, 1847): VC63 Elmfield Park, Doncaster. Gall on False Acacia
Robinia pseudoacacia T. Higgin bottom.

PSYCHODIDAE

Telmatoscopus ambiguus (Eaton, 1893): VC63 woodland around Woodall and Killamarsh Ponds
(SK477807) 28.6.2000 (OU) WAE.

SI MU LI I DAE

Simulium ( Simulium ) posticatum Meigen, 1838 [= austeni Edwards, 1915]: VC63 stream and marsh
at Birch Wood (SK435977) 18.10.2005 (99) WAE; Kilnhurst Ings (SK466977) 7.8.2006 (99)
WAE.

CERATOPOGONIDAE

Serromyia ledicola Kieffer, 1925: VC63 Old Spring Wood (SK535811) 23.5.2001 WAE; The
Deans, Listerdale (SK465922) 30.6.1991 WAE.

CHIRONOMIDAE

Parachironomus monochromus (van der Wulp, 1874): VC63 Harthill Lower Reservoir (SK489802)
2.6.2005 (dtf)WAE.

Paraphaenocladius penerasus (Edwards, 1929): VC63 Sheffield Airport (SK414885) 12.8.2001 (Otf)
WAE.

Paratanytarsus inopertus (Walker, 1856): VC63 Nor Wood, Roche Abbey (SK538904) 10.8.2001 (Otf)
WAE; Quarry Hills, Roche Abbey (SK5490) 28.6.1984 (Otf) WAE.

Psectrocladius ( Psectrocladius ) limbatellus (Holmgren, 1869): VC63 Thurcroft Colliery tip
(SK503906) 27.9.2003 (CfcT) WAE.

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The Naturalist 140 (2015)

HYBOTIDAE

Platypalpus incertus (Collin, 1926): VC63 Dalton, Huddersfield (SE157169) 30.5.2014 (2$)
Gavin Boyd det. RC, teste Adrian R. Plant. Both specimens are now deposited in the
National Museum of Wales in Cardiff (RC, pers. comm.).

DOLICHOPODIDAE

Chrysotus collini Parent, 1923: VC61 Reighton Cliffs, (clay cliffs) (TA1476) 26.6.2013 (3Cf) RC.

Medetera bispinosa Negrobov, 1967: VC62 Cayton Bay 5.7.1990 ((f) RC. Male genital examination is
necessary to separate this species from M. nitida (Macquart, 1834), which was added to the
Yorkshire list by Skidmore (1985). British records of M. nitido may all refer to M. bispinoso
(RC, pers. comm.).

PHORIDAE

Megaselia brunneipennis Costa, 1857: VC65 Thorpe Perrow Arboretum (SE2585/2685) 17.7.1982
(Cfcf)WAE.

M. collini (Wood, 1909): VC63 Don Canal towpath, Holmes Lock(SK415923) 13.8.2000 ((f(f) WAE.

M. stichata (Lundbeck, 1920): VC63 Birch Wood (SK437978) 18.10.2005 (OtT) WAE;

Herringthorpe Wood (SK458919) 11.10.2005 ((f6) WAE; Old Spring Wood (SK533810)
30.8.2000 (CfcT) WAE.

Triphleba smithi Disney, 1982: VC63 Quarry Hills (SK541901) 6.5.2000 (Otf)WAE.

PIPUNCULIDAE

Chalarus gynocepholus Jervis, 1992: VC63 Old Spring Wood (SK535811) 19.7.2000 ((ftf) WAE.

Eudorylas kowarzi (Becker, 1898): VC63 Barrow Colliery (SE3503) 5.6.2013 (Cf [dissected]) JDC.

AGROMYZIDAE

Agromyza lithospermi Spencer, 1963: VC63 Wath Wood Drive, Swinton (SK439991) 1.6.2011,
leafmine in forget-me-not Dean Stables; VC64 Hayton Wood near Aberford (SE445381)
7.2008, leafmine in Common Gromwell Lithospermum officinale, Chris S. V. Yeates.

A. sulfuriceps Strobl, 1898: VC64 Newton-in-Bowland (SD6950) 1.8.2011, leafmine in
Meadowsweet Filipendula ulmaria WAE; Swinsty Moor Plantation (SE1843) 17.9.2011,
leafmine in Raspberry Rubus idaeus WAE.

Cerodontha (Dizygomyza) iridis (Hendel, 1927): VC63 Sheffield Airport (SK414885) 12.8.2001 ((Qtf)
WAE.

C. (D.) morosa (Meigen, 1830): VC63 Maltby Low Common (SK544914) 5.6.1983 (CJtf) WAE.

Liriomyza eupatoriana Spencer, 1954: VC63 Greasbrough Street, Rotherham (SK426931)
7.8.1991 WAE.

L flavopicta Hendel, 1931: VC63 Treeton Wood (SK445867) 25.5.2000 (CfcT) WAE.

Melanagromyza cunctans (Meigen, 1830): VC63 Dodworth (SE3105) 3.7.2013 ((f) JDC; Edderthorpe
Ings (SE4106) 29.6.2013 ((f) JDC; Haigh (SE3011) 17.11.2013 ((f), 25.7.2014 (C?) JDC; Old
Moor (SE4202) 19.6.2013 (4(f) JDC; Rabbit Ings (SE3711) 26.6.2013 ((f), 21.9.2013
(C?[dissected]) 3.9.2014 (Cf) JDC; all taken where Bird's-foot-trefoil Lotus corniculatus occurs
at these Barnsley area sites (JDC, pers. comm.).

M. eupatorii Spencer, 1957: VC63 Maltby Common (SK548914) 8.6.1991 WAE; Shireoaks Quarry
(SK5481) 5.6.1985 WAE.

Ophiomyia collini Spencer, 1971: VC63 Thundercliffe Grange (SK379937) 3.8.1997 WAE.

O. melandricaulis Hering, 1943: VC63 Quarry Hills (SK5490) 18.5.1986 WAE.

O. orbiculata (Hendel, 1931): VC63 Edderthorpe Ings (SE4106) 29.6.2013 ((f [dissected]) JDC.

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147

Phytomyza (Phytomyza) artemisivora Spencer, 1971: VC63 Swinton Lock Adventure Park
(SK464991) 26.6.2003, leafmine in Mugwort Artemisia vulgaris WAE.

P. (P.) conii Hering, 1931: VC61 Catwick (TA1345) 25.7.2011 WAE; VC63 Broad Lane, Sykehouse
(SE6317) 21.5.2011 WAE; VC64 Thwaite Mill, Leeds (SE3231) 27.7.2011 WAE; all recorded
from leafmines in Hemlock Conium maculatum.

P. (P.) fulgens Hendel, 1920: VC63 Dodworth, by Whinby Road (SE3105) 2013, leafmines quite
common in Old Man's Beard Clematis vitalba JDC (teste Andy N.R. Godfrey [ANRG]).

P. (P.) pastinacae Hendel, 1923: VC63 Rainborough Park (SK402995) 28.7.1988 (C^) WAE.

SARCOPHAGIDAE

Metopia staegeri Rondani, 1859: VC61 Allerthorpe Common (SE755480) 17.7.2013 (Cf) IA det.
Daniel Whitmore; Calley Heath (SE751498) 4.6.2014 (Cf), 15.6.2014 (Cf) IA.

TACHINIDAE

Dufouria chalybeata (Meigen, 1824): VC61 Allerthorpe Common (SE755480) 30.5.2014 (Cf) IA
(teste Chris Raper [CR]); Calley Heath (SE751498) 4.6.2014 (C?) IA.

Gonia picea (Robineau-Desvoidy, 1830): VC61 Allerthorpe Common (SE755480) 26.3.2012 IA
(teste CR).

Phryxe heraclei (Meigen, 1824): VC61 Allerthorpe Common (SE755480) 28.7.2014 (Cf) IA.

Subclytia rotundiventris (Fallen, 1820): VC63 Haigh (SE3011) 17.6.2013 (9) JDC.

Re-instatements to Yorkshire Diptera List

CECIDOMYIIDAE

Neurolyga truncata (Felt, 1912) [= hammi (Edwards, 1938)]: VC63 Lindrick Golf Course (SK544824)
1986 Richard J. Hall; Roche Abbey (SK5489) 1985 John Pearson. Both these records are ex
Rotherham Data Bank sub nom. Cordylomyia hammi and were of galls on Lady's Bedstraw
Galium verum (WAE, pers. comm.). N. truncata was provisionally excluded by Grayson
(2006c).

Rhopalomyia palearum (Kieffer, 1890): VC64 Ling Ghyll (SD8078) 8.8.1987, gall on Yarrow
Achilea millefolium WAE. This [sub nom. Misospatha palearum] was excluded from the
Yorkshire list by Grayson (2005) as the only record known to him at the time was a
transcription error.

CHIRONOMIDAE

Cricotopus ( Isocladius ) ornatus (Meigen, 1818): VC 63 Chesterfield Canal (SK507823) 18.6.2000 (Otf)
WAE; Thrybergh Tip (SK460960) 12.7.2001 (Cfc?) WAE. This non-biting midge was formerly
excluded by Grayson (2009).

DOLICHOPODIDAE

Dolichopus ( Dolichopus ) caligatus Wahlberg, 1850: VC65 Marske, by river (SE113994) 13.7.2008
ANRG. An earlier record was stated to be a transcription error by Grayson (2006d).

SYRPHIDAE

Volucella zonaria (Poda, 1761). This very large hoverfly can here be excluded from the county list
and re-instated again by virtue of the following notes. V. zonaria was recorded from the
York area by Fife & Walls (1973); however, the record is suspect and can be discounted
without serious misgivings. All Diptera identifications in Fife & Walls (loc. cit.) are
questionable, some being obviously erroneous and others very doubtfully authentic. The
next published mention of V. zonaria in Yorkshire was by Stubbs (2005), followed by

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enumeration of the relevant record by Grayson (2006a). These referred to a sighting of V.
zonorio on the Yorkshire side of the River Rother in Rother Valley Country Park by WAE in
2004; however, WAE (pers. comm.) subsequently reported that the River Rother was re-
routed since Watson defined his vice-county borders and his sighting was on land in the
neighbouring VC57 (Derbyshire). The re-instatement of V. zonorio is due to Brian Smith's
photographs of a 9on Butterfly Bush Buddleio dovidii in his garden in Hilda Street, Goole,
on 26.8.2013. Two good photographs were forwarded to me by Peter Kendall for
verification of this record.

LONCHAEIDAE

Lonchaea scutellaris Rondani, 1874: VC65 Thorpe Perrow Arboretum (SE2585/2685) 17.7.1982
WAE. This fly was recorded from Yorkshire in MacGowan & Rotheray (2008) but provisionally
excluded by Grayson (2014).

Exclusions from Yorkshire Diptera List

MYCETOPHILIDAE

Mycetophila bohemica (Lastovka, 1963). Falk & Chandler (2005) stated "A record from Studley
Royal Park, Yorkshire, requires confirmation". This refers to a 9 taken at SE287691 on
19.6.1989 by Peter Skidmore and tentatively identified by him with the note "queried as
this species". There is no specimen standing under M. bohemico in Doncaster Museum and
Art Gallery. On this basis, M. bohemica is best provisionally excluded from the Yorkshire list.

Phthinia humilis Winnertz, 1863. This fungus gnat is best provisionally excluded from the
county list pending verification of its occurrence in Yorkshire. The three Yorkshire
records are from 1980-1982; therefore, specimens require re-examination, as they are likely
to be P. miro (Ostroverkhova, 1977) (JDC, pers. comm.).

CECIDOMYIIDAE

Contarinia acetosellae (Rubsaamen, 1891). The gall of this midge would appear to have been
erroneously recorded from Yorkshire due to a transcription error. It was listed by Grayson
(2007) on the basis that it was recorded from Yorkshire by Bagnall & Harrison (1918) as their
species no. 266, according to John Robbins (pers. comm.). However, there is no such record.

CERATOPOGONIDAE

Forcipomyia ( Thyridomyia ) monilicornis (Coquillett, 1905). This biting midge was tentatively
recorded as "? this sp." from Hatfield Moors by Skidmore (2001). There is another
Yorkshire record on the Malham Tarn cards, viz. "5.8.1978, North Wing 9 (runs t0
F.polustris in Edwards, 1926: no literature on rest of sub-genus, Thyridomio available)
RHLDisney". F. (7.) polustris sensu Edwards (1926) is synonymous with F. (7.) monilicornis
(Coquillett) but there are two other British Forcipomyia in the sub-genus Thyridomyia, of
which F. (7.) rugosa Chan & Le Roux, 1970, was recorded from Yorkshire by Boorman (1974)
[from the Rothwell area of Leeds, det. M.W. Service]. Provisional exclusion of F. (7.)
monilicornis from the county list would appear logical, given the degree of doubt about its
occurrence in Yorkshire.

AGROMYZIDAE

Cerodontha ( Butomomyza ) eucaricis Nowakowski, 1967. The record from Skipwith Common in
Grayson (2006b) was a misidentification for C. ( B .) scutellaris (von Roser, 1840) (J.H. Cole,
pers. comm.).

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149

Phytomyza ( Phytomyza ) pauliloewii Hendel, 1920. Spencer (1972) mentioned leafmines on
Burnet-saxifrage, stating " Pimpinella saxifraga L. Yorks.: nr. Settle, 30.vii.62 (G.C.D.
Griffiths). Whitish blotch mines (brown when old) of a Phytomyza sp., possibly referable
to Phytomyza pauli-loewi Hendel, 1920 (fig. 360)". This identification was tentative and
there have been no further Yorkshire records; hence, P. [P.) pauliloewii is best provisionally
excluded from the county list, pending verification of its occurrence.

SPHAEROCERIDAE

Trachyopella ( Trachyopella ) atomus (Rondani, 1880). This lesser dung fly' is probably best
provisionally excluded from the county list as Pitkin (1988) knew of only one British site,
and the record in Payne (1957) probably referred to T. (T.) lineafrons (Spuler, 1925);
however, this is not entirely certain as Payne ( loc . cit.) intriguingly stated his specimen from
Copmanthorpe on 22.9.1956 was "very small".

TACHINIDAE

Leiophora innoxia (Meigen, 1824) [= procera sensu auctt., nec (Meigen, 1824)]. This parasitic fly
is best provisionally excluded from the Yorkshire list. Belshaw (1993) stated that the only
record in northern England was an unconfirmed literature record from Yorkshire. Chris
Cheetham's record card [for Hypostena procera] probably alludes to the same record, as
Cheetham wrote "Ripon dist: C Morley in litt. Hincks".

Phebellia villica (Zetterstedt, [1838]) [= ingens (Brauer & von Bergenstamm, 1891)]. A 2 was
listed from Frog Hall in Durham, 27.8.1929 by van Emden (1954). Probably this was in error
for Frog Hall at the edge of Allerthorpe Common in Yorkshire; but regardless, the
review by Belshaw (1993) considered Q p- villica was indistinguishable from 2 P vicina
(Wainwright, 1940).

Further Notes

A Simplified Provisional List of Yorkshire Diptera [on the YNlTs web-site] is a 'work-in-progress'
which lists many species as being 'excluded' from the county list or 'queried' by their
determiners. Grayson (2014) regarded many of these 'queried' species as warranting provisional
exclusion from any definitive county list. No records have been published for any of the
following 'queried' species, which also warrant provisional exclusion: PSYCHODIDAE: Psychoda
erminea Eaton, 1898: DOLICHOPODIDAE: Dolichopus (Dolichopus) mediicornis Verrall, 1875:
AGROMYZIDAE: Phytomyza (Phytomyza) murina Hendel, 1935 [=brevicornis sensu Brit, auctt.,
nec Hendel, 1934]: and EPHYDRIDAE: Parydra ( Chaetoapnaea ) hecate (Haliday, 1833).

ASILIDAE

Ken Payne's record lists [in 13 folders now in the YNU archives] contain an intriguing
unpublished record of the robber-fly Eutolmus rufibarbis (Meigen, 1820), which may be correct.
However, it would represent a remarkable find and no voucher specimen has thus far been
located. As the asilid involved may have been a Machimus, then provisional exclusion from the
Yorkshire list is desirable. Ken's record was of a C? from Snake Hill Plantation, North Cave
(SE8634) 4.7.1981, with the note that it was initially recorded as Philonicus albiceps (Meigen,
1820).

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The Naturalist 140 (2015)

DOLICHOPODIDAE

Writing on the fauna of Beacon Lagoons Nature Reserve near Spurn, Cook (2009) mentioned "a
Dolichopodid fly (Tachytrechus hyollipennis), not previously recorded in Yorkshire." This refers to
Tochytrechus insignis (Stannius, 1831) taken from an old drainage ditch by RC (RC, pers. comm.).
There is no such species as T. hyollipennis. The first published Yorkshire record of T. insignis is
contained in Chandler (2002).

Acknowledgements

This paper would not have been possible without information and co-operation from the
following: Ian Andrews, John Coldwell, Jon Cole, Roy Crossley, Bill Ely, Andy Godfrey, Tom
Higginbottom, Peter Kendall and Chris Yeates.

References

Bagnall, R.S. & Harrison, J.W.H. (1918) A Preliminary Catalogue of British Cecidomyidae (Diptera)
with special reference to the Gall-midges of the North of England. The Transactions of the
Entomological Society of London, [65] (1917): 346-426.

Belshaw, R. (1993) Tachinid Flies: Diptera: Tachinidae. Handbooks for the Identification of
British Insects, 10 (4a i).

Boorman, J. (1974) Forcipomyia ( Thyridomyia ) rugosa Chan & Leroux (Dipt., Ceratopogonidae) from
Britain. The Entomologist's Monthly Magazine, 109 (1973): 183.

Brothers, P. & Grayson, A. (2014) Large Marsh Horsefly Tabanus autumnalis new to Mid-west
Yorkshire. Soldierflies and Allies Recording Scheme Newsletter, 2: 10.

[Chandler, P.J.] (2002) Dipterists Day Exhibits 2001 - compiled by Editor from exhibitors' notes.
Dipterists Digest, (Second Series) 9: 19-21.

Cook, P.J. (2009) Erosion & Accretion - Threats and Ephemeral Opportunities. YNU Bulletin, 52
(Suppl): 33-36.

Edwards, F.W. (1926) On the British Biting Midges (Diptera, Ceratopogonidae). The Transactions of
the Entomological Society of London, 74: 389-426.
van Emden, F.l. (1954) Diptera: Cyclorrhapha: Calyptrata. (1) section (a): Tachinidae and
Calliphoridae. Handbooks for the Identification of British Insects, 10 (4a).

Falk, S.J. & Chandler, P.J. (2005) A review of the scarce and threatened flies of Great Britain. Part 2:
Nematocera and Aschiza not dealt with by Falk (1991). Species Status 2. Joint Nature
Conservation Committee. Peterborough.

Fife, M.G. & Walls, P.J. (1973) The River Foss. From Yearsley Village to York: Its History and Natural
History. Sessions Book Trust, York.

Grayson, A. (2005). Additions and corrections to the Yorkshire Diptera list (Part 1). The Naturalist,
130: 129-136.

Grayson, A. (2006a) Volucella inanis and V. zonaria in Yorkshire, plus notes on other species.
Hoverfly Newsletter, 41: 9.

Grayson, A. (2006b). New Yorkshire Mycetophilidae and Agromyzidae (Diptera). YNU Bulletin, 46:
37-42.

Grayson, A. (2006c) Additions and corrections to the Yorkshire Diptera list (Part 2). The Naturalist,
131:83-93.

Grayson, A. (2006d) Additions and corrections to the Yorkshire Diptera list (Part 3). The Naturalist,
131: 131-138.

Grayson, A. (2007) A brief progress report on the forthcoming provisional list of Yorkshire Diptera.
YNU Bulletin, 48: 60-61.

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Grayson, A. (2009) Additions and corrections to the Yorkshire Diptera list (Part 4). The Naturalist,
134: 24-29.

Grayson, A. (2014) Additions and corrections to the Yorkshire Diptera list (part 5). The Naturalist,
139: 14-22

MacGowan, I. & Rotheray, G.[E.] (2008) British Lonchaeidae: Diptera Cyclorrhapha, Acalyptratae.

Handbooks for the Identification of British Insects, 10 (15).

Payne, K.G. (1957) Diptera in The Yorkshire Naturalists' Union: Ninety-fifth Annual Report:
Entomology. The Naturalist, [82]: 21-25.

Pitkin, B.R. (1988) Lesser Dung Flies. Diptera: Sphaeroceridae. Handbooks for the Identification of
British Insects, 10 (5e).

Skidmore, P. (1985) Diptera Report: 1977-84. The Naturalist, 110: 111-117.

Skidmore, P. (2001) A provisional list of the insects of Hatfield Moors. THMCF Technical Report 7.

Doncaster: Thorne and Hatfield Moors Conservation Forum.

Spencer, K.A. (1972) Diptera: Agromyzidae. Handbooks for the Identification of British Insects, 10 (g).
Stubbs, A.[E.] (2005) Flies. In Wildlife reports. British Wildlife, 16: 437-438.

YNU VC63 Field Excursion to Thorpe Marsh 14th June 201 4
Bryology Report

Colin Wall

The old railway embankment bisecting the reserve from east to west was probably the most
productive area for the bryologist. With a substrate rich in acid clinker/ash and basic slag ballast,
it exhibited a high degree of opportunity for both calcicoles and acidophiles, and a variety of
common examples from both persuasions had become established, interspersed with more
neutral species. Within a footfall of acid-loving Campylopus introflexus and Brachythecium
albicans could be found calcicoles such as Plagiomnium undulatum and Didymodon insulanus,
against a general background of neutral and ubiquitous Brachythecium rutabulum and
Kindbergia praelonga. Small acrocarps such as Pseudocrossidium hornschuchianum, Barbula
unguiculata and B. convoluta had become established where the substrate had become hard-
packed. The embankment, shaded by mature oaks and hawthorns, had Fissidens taxifolius and
F. bryoides with a little Mnium hornum and Polytrichum juniperinum.

Despite the virtual absence of suitable host trees for epiphytes (only two Ash trees were noted),
the oaks supported a number of mosses that were rare only ten years ago but are now thriving
due to the reduction of atmospheric sulphur dioxide levels. These included Ulota bruchii, U.
phyllantha, Cryphaea heteromalla, Orthotrichum affine, O. diaphanum and O. lyellii . The latter is
still scarce in the Doncaster area, so it was pleasing to see it on Ash, though it was first found on
the reserve in 2011 on oak. Other epiphytes included the liverworts Frullania dilatata and
Metzgeria furcata, both on oak.

On the day 43 species of bryophytes were recorded, compared to 49 recorded on visits in 2009
and 2011. There were, however, five that had not been previously recorded. Among these was

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The Naturalist 140 (2015)

the thalloid liverwort Pellia epiphyllo, the flap-like involucre in evidence. The most surprising
addition to the reserve's list was a very small amount of the tiny Leskea polycorpo on Elder at
the foot of the embankment.

Botanical Report for 2014

Phyl Abbott, Richard Middleton, Gill Smith & Linda Robinson

Email: phyl.a@virgin.net

VC61 South-east Yorkshire

The greater part of all botanical recording activity during 2014 was directed towards achieving
good coverage for the forthcoming Atlas 2020. This has resulted in a great many new records
but mainly for more common-place plants in areas which have been less well recorded. All
records for rare and scarce plants have been published in the 3rd Edition of the Rare Plants
Register (Middleton & Cook 2015). There was, of course, the occasional surprise. Among these
was a clump of Monk's-hood Aconitum napellus on the verge at Thwing; obviously an
introduction but matching a herbarium specimen collected by Professor Ron Good from this
locality in 1956! The 'square bashing' also provided useful records for Spiny Restharrow Ononis
spinosa and Lesser Hawkbit Leontodon soxotilis on the Humber banks at Pauli Fort. Less
welcome was the large quantity of New Zealand Pigmyweed Crassulo helmsii in the village pond
at Fimber (John Killingbeck).

Over the previous winter, storm surges re-modelled the landscape around Kilnsea and Spurn.
Large amounts of sand were moved and it was feared that much damage would have been done
to the flora. The large patch of Intermediate Polypody Polypodium interjectum south of the
Warren Cottage seems to have succumbed but, rather surprisingly, Sea-holly Eryngium
moritimum now seems to be thriving on the dunes between Kilnsea and Easington and the
upper strand-line in this area has sprouted a long line of the UK BAP species Prickly Saltwort
So I solo koli subsp. koli.

There were two records this year for Small-flowered Catchfly Silene gollico, the first this
millennium, from farms in the Vale of York. Although neither is thought to have been a
deliberate introduction, caution must be exercised in interpreting these occurrences as they
were both near areas where other cornfield flowers had been seeded.

The late May Botanical Section meeting in Millington Dale was notable for a large well-spread
colony of over 100 flowering spikes of Frog Orchid Coeloglossum viride, found by Martin
Stringer; this is a site for which there seem to be no previous records. The cliffs between
Reighton and Speeton provided several sites for Grass-of-Parnassus Parnossio polustris, a plant
which is now confined in the vice-county to Filey Bay. Alerted to its presence by Sarah White, it
was good to see it growing so well. It was with Felwort Gentionello omorello in areas that had,
earlier in the year, produced a profusion of Pyramidal Anocomptis pyromidolis and Fragrant
Orchids Gymnodenio conopseo.

The Naturalist 140 (2015)

153

Two notable sedge records were Pale Sedge Corex pollescens, found by Gabrielle Jarvis and
Rohan Lewis near Houghton Hall and considered rare in the vice-county, and a patch of Divided
Sedge Corex divisa at Skeffling, last recorded by Eva Crackles in 1956 and now re-discovered by
Peter Cook.

The South-east Yorkshire Rare Plants Register (RPR) is available for free download from the VC61
BSBI pages: http://www.middletonl2.karoo.net/.

Richard Middleton

VC62 North-east Yorkshire

These notes are heavily biased to Ryedale as I received no other records, nor did I botanise in
other parts of the vice-county myself. 2014 had a remarkably mild start, with plenty of sunshine
in Ryedale.

Snowdrops Golonthus nivalis were more or less fully out by 1 February and at their best to 24
Feb. White and purple Sweet Violets Viola odorata were flowering nicely near Whitwell by 12
March. Blackthorn Prunus spinosa was out by 9 April, and Wild Cherry Prunus avium and Plum
Prunus domestica were opening. On 24 April both golden saxifrages were flowering together
near Hovingham, but there was no sign of the Greater Chickweed Stellaria neglecta. I counted
24 Early-purple Orchids Orchis mascula on the roadside just south of Gilling with a few hybrid
False Oxlips Primula x polyantha nearby.

Herb Paris Paris quadrifolia was doing very well in Gilling with 50+ plants. The Baneberry Actaea
spicata was weak though, with one plant of the three broken/eaten, probably by deer, and only
one showing flowers and on only one spike. Those on the east side of the road at Ashberry were
faring poorly but the plants on the west side were strong and healthy, though again with only
one flower spike.

On the Ryedale Naturalists' trip to Raindale on 11 May it was good to re-find Intermediate
Wintergreen Pyrola media in the same spot (SE806924) where it had been seen 50 years before
by Gordon Simpson. He also showed us Dwarf Willow Salix herbacea at SE813923. Bob Dicker,
via Nan Sykes, reported quite a good population of the wintergreen not far from the 'fire tower',
or Red Dyke (at c. SE892887), on 19 June.

Globeflower Trollius europaeus was reported in woodland at Beadale, Wrelton. This is an old
site but it hadn't been seen for several years, so it's great to have it back. In Spaunton Quarry
(SE7287) on 20 May we found a swathe of Adders-tongue Ophioglossum vulgatum - 100 or
more plants under bracken , as well as some Fly Orchids Ophrys insectifera .

Hairy Rock-cress Arabis hirsuta and Knotted Clover Trifolium striatum were both in flower at
Hutton Common (SE7088) on 5 June. There was a small patch of the former of about 20 stalks,
and three small clumps of Knotted Clover.

As a result of some wet weather all the roadside vegetation grew incredibly tall and lush.
Southern Marsh-orchids Dactylorhiza praetermissa at Castle Howard Arboretum were
spectacular on 11 June. Horse-radish Armoracia rusticana on a roadside verge at Whitwell
produced a flower spike - the first I can remember seeing.

154

The Naturalist 140 (2015)

Roy Crossley recorded Lesser Water-plantain Baldellio ronunculoides in flower on 16 June on
the shore of a recently created pod/scrape near a track within the Strensall Common MOD
Danger Area at ca. SE650595.

A small colony of Bee Orchids Ophrys apifera in a clearing in Gilling Woods, which I have been
watching for a few years, produced 13 spikes this year, some very tiny with only one flower, but
the highest number so far. There were two Bee Orchids at Bull Ings , which is good as I didn't see
any there last year. Eight Fragrant Orchids Gymnadenia conopseo and Pepper-saxifrage Siloum
silaus were just coming into flower on 4 July.

In late June/early July something very nasty affected willows, notably Goat Willow Solix caprea,
around Gilling. A combination of rust and beetle attack meant that many trees looked dead by
mid July, although some, at least, tried to put out new shoots. In the second half of July there
was perfect summer weather. A plant of Welted Thistle Corduus crispus was found near Gilling
on a field edge (SE6276) on 20 July.

It was a poor acorn year, though good for beech mast, hazel nuts, plums, haws and some
blackberries - but plenty of knopper galls. The end of October to mid November was remarkably
warm and quite wet. There was poor autumn colour, as there was no frost.

A further species list is available from the YNU website www.ynu.org.uk.

Gill Smith

VC63 South West Yorkshire

Report not available.

VC64 Mid-west Yorkshire

Many interesting plants were seen during field meetings of the Bradford Botany Group. Among
an extensive display of Bluebells Hyocinthoides non-scripto at Rougement Carr in April, a few
had pink (var rosea) or white (var alba) flowers. Also in the woodland were a few spikes of
Toothwort Lathraea squamaria and in the nearby Weeton churchyard at SE283465 the
Goldilocks Buttercup Ranunculus auricomus was surprisingly frequent. During the visit to Park
Rash on 7 June, in monad SD9774 Mountain Everlasting Antennaria dioica and Fragrant Orchid
were found and in SD9874 Pyrenean Scurvygrass Cochlearia pyrenaica, Bird's-eye Primrose
Primula farinosa, and Globeflower.

On the naturally revegetated Sun Lane tip at Burley-in-Wharfedale, SE1546, on 23 July, we saw
Small Teasel Dipsacus pilosus in one of its seven sites in VC64, Water Violet Hottonia palustris,
and several alien species including Stinking Iris Iris foetidissima in its third site in VC64 and
Filbert Corylus maxima, a new plant for the vice-county. On 26 July the Attermire and Langcliffe
reserve in SD8365 was particularly species-rich. Finds there included Moonwort Botrychium
lunaria , Dioecious Sedge Carex dioica , the hybrid between Tawny Sedge, and Long-stalked
Yellow Sedge Carex x fulva, Autumn Gentian Gentianella amarella, Herb Paris, Grass of
Parnassus Parnassia palustris, Spreading Meadow-grass Poa humilis, Holly Fern Polystichum
lonchitis, Lesser Clubmoss Selaginella selaginoides, Limestone Fern Gymnocarpium
robertianum, Mountain Pansy Viola lutea , and Green Spleenwort Asplenium viride, some of
whose fronds were forked making it A. viride var. multifidum.

The Naturalist 140 (2015)

155

Visits to Fairburn Ings RSPB reserve, by Leeds Naturalists' Club on 16 July and by Bradford
Botany Group on 21 August, boosted the total number of plants recorded in SE4527 to 327. The
most intriguing plant was a dwarf Centaury, about 3 inches tall and with bright pink flowers,
growing in patches alongside the Common Centaury Centourium erythreo at the edge of the car
park. This was identified, from photographs sent to Dr F. Ubsdell, the BSBI referee for
Centaurium, as Lesser Centaury Centourium pulchellum. Most of its British sites are in southern
England with a few in coastal areas further north. The only previous records in Yorkshire were
from Redcar in 1892 and 1930. It is new to VC64.

John Webb has found two new sites for Narrow-leaved Water-plantain Alismo lonceolotum , in
the Leeds and Liverpool canal near Silsden at SE053447 and near Morton at SE092415. A steeply
sloping, north-facing field in Littondale, SD9569, had a good diversity of plants including Marsh
Helleborine Epipactis palustris, Autumn Gentian, Grass of Parnassus, Sea Plantain Plantago
maritima, Creeping Willow Solix repens , and Saw-wort Serrotulo tinctorio. While monitoring
plants on the Malham Tarn estate we found Jacob's-ladder Polemonium coeruleum in a new site
to the west of the tarn and, although Alpine Bartsia Bartsia olpino was quite plentiful, there
were no flowers this year

During the YNU meeting in SD7666, at Austwick Moss we could find only a small amount of
Cranberry Vaccinium oxycoccos amongst the dominant Purple Moor Grass Molinio coeruleo. In a
damp hollow there was a beautifully flowering patch of Round-leaved Sundew Drosero
rotundifolia at SD762666. Lawkland Moss had more plants of interest, including Meadow
Saffron Colchicum autumnole at SD767666, Dyer's Greenweed Genista tinctoria (SD767666),
Slender St John's-wort Hypericum pulchrum (SD767666), Blunt-flowered Rush Juncus
subnodulosus, Bogbean Menyanthes trifoliata, (SD761664), Marsh Cinquefoil Comarum palustre
(SD763667) and Saw-wort Serratula tinctoria (SD767666).

Rare and scarce plants in new tetrads:

Scientific name

Vernacular name

Location

Recorder

Dryopteris submontana

Rigid Buckler Fern

Attermire, SD8264

Bradford Botany Group

Alchemilla glaucescens

l !

Silky Lady's Mantle

Sleets Gill, SD958688

P. Abbott, B. Brown, C.
Florner

Primula farinosa

Bird's-eye Primrose

Attermire, SD8264

Bradford Botany Group

Species new to the vice county

Scientific name

Vernacular name

Location

■

Recorder

Betula populifolia

Grey Birch

Allerton Bywater
SE414284

P. Abbott, K. McDowell

r"1"" - — — ~~ *i

Centaurium pulchellum

Lesser Centaury

Fairburn Ings, SE4527

Leeds Naturalists' Club,
Bradford Botany Group

Corylus maxima

Filbert

! i

Burley-in-Wharfedale

SE1546

Bradford Botany Group

Phyl Abbott

156

The Naturalist 140 (2015)

VC65 North West Yorkshire

Yellow Star-of-Bethlehem Gogea luteo was found near the Round Howe below Richmond at the
end of March by Jan Owen and LR, last recorded from here in the 1960s. It was also found in
Iron Banks Woodland in April, and downstream from Richmond in a few new sites by Trevor
Lowis (TL) and LR. Further sightings were on the riverbank near Brompton-on-Swale and in
profusion in the riverside woodland near Catterick village in April by Trevor Lowis and LR, all new
sites. There is an old 1800s record for the plant near Asenby.

Henbit Deadnettle Lomium omplexicoule was found in April on dry banks between Catterick
Bridge and Scorton by Brian Burrow (BB) and LR, and more has been spotted since then in arable
fields near Ainderby Steeple and Danby Wiske later in the year. These are the first records for
VC65 since the 1960s.

Juniper Juniper communis - a seedling found among the heather at Uldale Head by Tim & Eileen
Laurie (T&EL), TL and LR in August was the second seedling spotted well away from any Juniper
stands and bodes well for the plant now that grazing on the fells has reduced. The first seedling
was noted just over the border into Cumbria.

Pyrenean Lily Lilium pyrenaicum - a few plants naturalised on the edge of the Tees about 200
yards downstream from Wynch Bridge are probably garden escapes. Spotted by LR in June.

Dwarf Mallow Malva neglecto was seen in profusion in the village of Danby Wiske by Allison and
LR in August, it was also growing alongside tracks through arable fields around the village.

Vernal Sand-wort Minuartio verna, a beautiful double-flowered form was spotted by Dave
Hickson and LR in June on the Ballowfield Nature Reserve in Wensleydale.

Royal Fern Osmundo regolis - one plant was seen by Chris Irvine and LR in the gorge at the Fairy
Glen Waterfall near Holwick in Teesdale in June.

Scottish Goat Willow Salix coprea subsp. sphacelota has been found in previous years in upland
gills and scars in Swaledale, Teesdale and Wensleydale. T&EL and LR spotted another ancient
specimen in Hebblethwaite Gill above Hebblethwaite Hall near Sedbergh. It must have been an
integral part of the original woodland which is now just hanging on in these upland gills and
scars. I believe that these unique scar woodlands need protection as examples of the original
'wildwood' with its unique DNA, present after the ice melted 12,000 years ago. Planting of
shrubs and trees from 'foreign' sources should be prevented to protect their unique
provenance.

Hairy Stonecrop Sedum villosum was seen by Brian Burrow (BB) and LR in June whilst walking
down Arten Gill as a small patch of around 34 plants on a flushed stream bank.

Tomato Solanum lycopersicum plants were found naturalised on the shingle banks on the Swale
near Great Langton by LR in August. This is a new record for VC65.

During a Field trip to Morton-on-Swale we came across a drainage ditch with old fenland plants
including Common Reed Phrogmites australis, Bottle Sedge Carex rostrata, Brown Sedge Carex
disticha, False Fox-sedge Carex otrubae, Common Meadow-rue Thalictrum flavum and Purple-
loosetrife Lythrum salicaria still growing on the ditch edge. Nick Morgan got permission to visit

The Naturalist 140 (2015)

157

a garden bordering the old fen site below Ainderby Steeple the following week, where we found
a remnant about quarter of an acre in size of this fen vegetation still intact bordering the old
'bottoms7. The owner of the garden had built a pond just above this remnant and the fen
vegetation was spreading nicely round it. Plants found here were Skullcap Scutellaria
galericulata, Common Meadow-rue, Bottle Sedge, Purple-loosestrife, Amphibious Bistort
Persicaria amphibia, Common Reed and Greater Pond-sedge Carex riparia, all mentioned in a
Yorkshire Naturalists' Union Report of a Field Meeting to Ainderby Steeple on 22 June 1946. The
wet field beside this remnant fen has only just been successfully drained in the last two or three
years and it's a pity that funding couldn't be found to purchase this field and re-wet it. It would
make a wonderful Nature Reserve and preserve an example of the now-lost botanically rich fens
and carrs, once common in this area before they were drained for arable crops in the 1950s.

Linda Robinson

YNU Notice

YNU Annual General Meeting

Notice is hereby given that the 153rd Annual General Meeting of the Yorkshire Naturalists' Union
will take place at the 'Lakehouse', Ron Cooke Hub, University of York on 14 November 2015.
The meeting will be preceded by a meeting of the Natural Sciences Forum and followed by an
address from outgoing YNU President Dr. Geoff Oxford.

The full programme for the day is as follows:

9:30 Registration

9:45 Short guided walk around the Heslington East development, led by Professor Chris

Thomas FRS (expert on butterflies and climate change). Chris has been on the committee
planning the environment of Heslington East from the start and will explain the thinking
behind the extensive landscaping features, which include lakes, hay meadows and
woodland.

10:30 Refreshments
11:00 Natural Sciences Forum
12:30 Cold Buffet Lunch
13:30 Group photo

13:45 AGM, hosted by the Yorkshire Mammal Group

14:45 Presidential Address: 'A roll of the dice: the unnatural history of Large House spiders in
the British Isles.

15:30 Refreshments

16:00 Meeting close and departure

The charge for the day will be £14.00, which includes lunch.

Members may book online at www.ynu.org.uk, or with a cheque to the YNU Treasurer, Barry
Warrington, Hessle Mount Farm, Jenny Brough Lane, Hessle, HU13 0JZ (treasurer@ynu.org.uk).

Details of how to get to the site can be found at: http://www.vork.ac.uk/about/maps/. Parking is
free on campus at weekends, and there is a car park a short walk from the venue, signposted
'Ron Cooke Hub'. Inside the building, go to the second floor, following signs to the Lakehouse.

158

The Naturalist 140 (2015)

Book review

Butterflies of Lesbos and Dragonflies of Lesbos, ebooks by John Bowers. These ebooks will be
available as free downloads when the Friends of Green Lesbos website www.greenlesbos.com is
rebuilt. In the interim they are available from the authorj.k.bowers@icloud.com

The subject matter of these ebooks lends itself well to this modern treatment. The author's aim
is to enable general naturalists and members of the public to be able to recognise the island's
butterflies and dragonflies with a minimum of need to catch specimens, other than with a
camera. He provides good photographs of all of the species (including upper and lower surfaces
when necessary) and each is annotated with clear identification features. Similar species are
compared side by side. There are some groups of butterfies, such as Meadow Browns and
Graylings where this treatment doesn't work, and the author admits that these can only be
identified to genus level. He points out that greater confidence cannot be achieved by anything
other than microscopic examination of dead specimens, something that he does not feel is
justified for mere casual identification.

The general biology of both groups of insects is included where it will assist with finding them.
Status, habitat and distribution notes are given for each species and there are introduction
pages for each of the major groups and sub-groups (e.g. Whites, Graylings, Damselflies etc).
Photographs and descriptions of the main habitat types on the island are useful features and a
list of the larval food plants of butterflies will aid the search for a particular species, as well as
identifications. Given that Lesbos can appear a hot and dry island in summer, he gives good
information, with maps, of where water is present year-round and will have likely places to find
dragonflies.

For many readers a difficulty of the Dragonflies book could be that, though a table of English
names is given at the end, all the species descriptions mention only their scientific ones. Whilst
there is unfortunately no general agreement over many of the English names, a good number
(e.g. Emperor Dragonfly, Broad-bodied Chaser) are well-established and could perhaps have
been included in the main text.

This said, the real joy of both of these ebooks is their effective use of a simple technology trick -
the hyperlink. A comprehensive index and a link symbol on each page allow the user to move
from page to page with a simple tap or key press. The layout is clear and consistent, though
perhaps not 'polished', and there are a few textual errors which will no doubt be removed as the
books evolve. The author's informal style encourages involvement and interest and because it
works well on ipads and mobile phones as well as on a laptop it should allow any of us to
identify the butterflies and dragonflies seen on Lesbos (and many of those seen on neighbouring
islands) with confidence.

RPS

The Naturalist 140 (2015)

159

YNU Calendar 201 5

Up-to-date information can also be found on the YNU website at:

w w w.y n u . o rg. u k/e ve nts/ge n e ra I

Sept 5 Conchological Section Field Meeting. 10:30. Fridaythorpe, Driffield, East Riding of

Yorkshire. For further details contact A. Norris via AdrianXNorris@aol.com.

Oct 3 Conchological Section Field Meeting. 10:30. Murton Wood, North York Moors

National Park. For further details contact A. Norris via AdrianXNorris@aol.com.

10 Bryology Section Field Meeting, Kilburn. Meet at 10:00 in the White Florse car park
at SE514811.

24 YNU Executive Meeting. 10:30- 12:30 St Chad's Parish Hall, Headingley, Leeds

31 Conchological section AGM. 13:00 - 16:00. For further details contact A. Norris via
AdrianXNorris@aol.com.

Nov 14 AGM, York. Preceded by Natural Sciences Forum (see details on pl59).

2016

Mar 19 YNU Conference - advance notice. Theme: 'Names, knowledge and natural

history - the importance of modern taxonomy to the amateur naturalist'. At the
National Science Learning Centre, University of York.

Endpiece: Cowslip Primula veris, by Dorothy Bramley (see The Naturalist 140, 68-70)

160

The Naturalist 140 (2015)

Yorkshire Naturalists' Union

c/o NEYEDC, St William College, 5 College Street, York YOl 7JF
Tel: 01904 641631 Email: membership@ynu.org.uk
Website: www.ynu.org.uk
Registered Charity No. 224018

The Naturalist

This publication is issued free to individual members of the Yorkshire Naturalists' Union and to
Affiliated Societies. The Editorial Board of The Naturalist is currently:

J. Bowers, W. Ely, A. Henderson, A. Millard, P. Simmons

Notice to contributors

Contributors should indicate whether they wish their manuscripts to be subjected to anonymous peer review. All other
manuscripts will be reviewed by the Editorial Board who at their discretion may send them to third parties for comment
and advice.

Please note change of email address for submission of articles, which should now be sent as an MS Word document
to Dr A. Millard at editor@ynu.org.uk

Please look at a recent issue of the journal for a general idea of how to present your article. Also see The Naturalist
Guide to Consistency on p77 of The Naturalist 1079 and please avoid the following:

• using any paragraph formatting and line spacings other than single.

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Good quality, high resolution images are very welcome and should be sent as .jpg files, with a separate MS Word file
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If electronic submission is not possible, contributions should be sent to Dr. A. Millard, Woodland Villas, 86 Bachelor
Lane, Horsforth, Leeds LS18 5NF (Tel. 0113 258 2482).

Contributors should ensure the accuracy of reference citations. The Editorial Board and Council accept no responsibility
for opinions expressed by contributors.

Copy Dates:

April issue - 14 February; August issue - 14 June; December issue - 14 October

Single copies may be made of single articles in this journal provided that due acknowledgement is made and the copies
are for non-profit making educational or private use. Copying of more than one article or multiple copying of a single
article is forbidden unless special permission has been obtained from the Yorkshire Naturalists' Union. Permission is
granted for the use of brief quotations in published work provided that acknowledgement of the source is clearly stated
but the use of substantial sections of text and any illustrative matter requires the express permission of the Yorkshire
Naturalists' Union.

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