1. Art and Diseño

(—;
MONKS
EF
S9
2(063)
C.) N4I_t
WOOD
EXPERIMENTAL
STATION
No. I
SYMPOSIUM
March 23rd-25th,
1965
THE
CONSERVATION
OF
INVERTEBRATES
THE
NATURE
CONSERVANCY
•
I
STATION.
VAPERIMENTAL
WOOD
IIIONKB
EGOLORT.
TERREBTNIAL
OF
INSTITUTE
[UPTON.
ABBOTS
HUNTINGDON.
THE
OF INVERTEBRATES
CONSERVATION
Station
heldat MonksWoodExperimental
A Symposium
March23rd- 25th1965
Section
Research
by theConservation
Organised
editedby
Proceedings
Dr. EricDuffeyandDr. M. G. Morris
tIONK8
WOOD
WERIMENTAL
8TATION,
• INOTITUTE.O
TEORESTD1.
F
ECOLOGY,
11.
ABBOTO
RIPION,
HUNTIMIDON.
5q2_(061)
SYMPæIUM on the
CONSERVATION
of INVERTEBRATES
C ONTENTS
Page
Programmeand Summary
List of Participants ..
..
..
..
5
OpeningAddress- Invertebrate
Conservation
in relation
to theConservancy's
commitmentand
s .the
developmenof
t its researchprogram:no.
E. Duffey.. .. .. ..
The Fate of areasof invertebrate
intereStas listedin
1949. J. F. D. Frazer ..
..
..
..
..
16
Extinctionand
s Invasions
- somecasehistories
and
conclusions.
J. E. Satchell ..
..
..
19
Invertebrate
Conservation
and Management
Plans. E. Duffey 29
StatusandConservation
of theLargeBlueButterfly,
Maculinea
arionL.
0. D. Hunt ..
35
The Conservation
of Dragonfliein
s Britain.N. W. Moore
45.
Basicecological
knowledge
requiredfor invertebrate
conservation.
E. Broadhead ..
..
..
52
Why we needinvertebrate
surveys.M. G. Morris .
65
Organization
andPracticeof intensive
surveys.
W. O. Steel
.. ..
•
.. ..
71
SomaFeaturesof Conservation
Interestarisingfrom surveys.
A. M. Massee .. .. .. .. .. ..
77
00
00
00
06
1
00
00
The Role of Invertebrate Collections in the Conservancy.
T. Huxley ..
..
..
..
..
..
..
..
General Discussion ..
.
83
89
á
PROGRAMMEand SUIVARY
This meetingwas the firststaffsymposium
to be held at
MonksWood and was attended
by 32 representatives
fromneaplyea
the regionalofficesinEngland',
Walesand Scotland;theResearch
Stationsand Specialist
Branches.In addition,
5 invitedecologists
attendedto.readpapersor.takePart in the discussions
and .the
organisers
are gratefulto themfor theirhelp and interest:-Mr.
CharlesEltonF.R.S.,Dr. A. H. Masses,Dr. E. Broadhead,
Mr. 0. D.
Hunt-apdMr. W. O. Steel. Elevenpaperswere read duringthe three
days and two fieldexcursions
arranged, to Woodwalton
Fen N.N.R.
and to CastorHanglands
N.N.R.andBarnackHillsandHoles5.5.8.1.
Discussions
on the importance
of invertebrate
studiesfor
ecological
researchin relationto conservation
problemshavebeen
held from_time
to timeby Conservancy
scientific
staff. Some of the
problemswere definedby CaptainDiver in 1954 in a paperto the
Scientific
PolicyCommittee.The purposeof thisSymposium
was to
discussthepracticalaspectsofinvertebrate
conservation
under
fourMain themes whichhad been arranged
with Mr. T. Huxley:(i) to examinethe Conservancy's
recordand its achievements
in
this subjectsince
1949; (ii)toanalyse
the typesOf problemS-in
the conservation
of invertebrate
animalson N.N.Rs.and thefl
contributionmade to themby currentresearch;(iii) to assessthe
dbjectsand valueof invertebrate
surveysand (iv) to discuasthe
purpose,use and organisation
of invertebrate
collections
in the
Conservancy.
The prograumwas ai follows:Tues
March23rd
ThemeI
Chairman.Dr. E. Duffey
OpeningAddress- Invertebrate
Conservation
in relationto ti:a
Conservancy's
commitfrents
and the developrent
of its researchprograMme.
Dr. E. Duffey.
The.fateof areasof invertebrate
interest
in Enc,land,
Walesand Scotlandknownin
1949.
Dr. J. F. D. Frazer.
Extinctions
and invisions:solre case
histories
and conclusions.Dr. J. Satchell.
Visit to Woodwalton
7en ledby Dr. E.
Duffey-and
Dr. M. G. Morris.
-2-
Wednes
March24th
ThemeII
Thurs
2 th
N.z.trch
ThemeIII
March25th
Thurs
• •
continued
ThemeIV
Chairman:Dr. K. Mellanby
and management
conservation
Invertebrate
plans. Dr. E. Duffey.
of theLarge-Blue
The.ConservatiOn
Butterfly.Mr. 0. D. Hunt.
s Nature
of Dragonflieon
The Conservation
Reserves.Dr. N. W. Moore.
.requiredfor
knowledge
Basicecological
Dr. E. Broadhead.
conservation.
invertebrate
Visit to BarnackHillsandHolesS.S.S.I.
N.N.R.led by Dr. M.
and CastorHanglands
George.
Chairman:Dr. E. Duffey
surveys.Dr. M.
Why we need invertebrate
G. Morris.
and practiceof intensive
Organisation
surveys.Mr. W. O. Steel.
interest
Some featuresof conservation
arisingfrom surveys.Dr. A. N. Nassee.
s
collectionin
The roleof invertebrate
theNatureConservancy.Mr. T. Huxley.
of ThemeIV. Summingup
Continuation
by GeneralDiscussion.
followed
.
con,'s work in invertebrate
ThemeI - A reviewof the Conservanc
servation.
greatlyhelpedby a 20-pageaccount
s
The discussionwere
the work of theNature
preparedby CaptainC. Diverdescribing
inverteCommitteein the 1940sconcerning
ReservesInvestigation
Command
preparing
to theCommittee
bratesandthe factsavailable
primarily
18 siteswere listedas P.N.N.Rs.
7122. In thisdocument,
reasons.So littlewas known
(mainlyentomological)
for zoological
s
and populationof
requirements
ecological
abOutthe distribution,
thatthe safestsolutionto theproblemof conservation
invertebrates
was presenton
was to ensurethat the greatestvarietyof habitats
at that timethat
the prqposedN.N.Rs. However,it was recognised
typeswouldfail to savemany
based solelyon vegetation
selection
was made duringthe Reference
s.
population
localised
and
rare
by theRoyal
of thisthemeto an undatedlistprepared
discussion
Society(in the early1940s?)of 158 sitesworthyof
Entomological
grounds. This listis not referredto
on entomological
preservation
at 1949,therehad been
in Cmd. 7122. The meetingagreed.tnsince
on problemsof invertey
verylittleprogressin Conservancthinking
in the lightof
A re-assessmen
t necessary
was
brateconservation.
conservation
of the Britishfauna. Invertebrate
modernknowledge
- 3-
of.other
couldnot.bedeveloped'without
dueregard-for-the
needs
organisms
and thewholeshouldformpartof a national
conservation
policy.
ThemeII - Invertebrate
conservation on NatureReserves
An analysis
of 550 research
projects
(in 69 Management
Plans)'showed
thatonly47 were concenned
withstudies
on invertebrates.Lessthanhalfof thesewerebeingconducted
by Conservancy
staffand fewwereplannedin relation
to management
requirements.
The vastnuMbers
of invertebrate
species
and ahortage
of specialists
meantthatmanagement
studies
muStbe carefUlly
selected.It was
proposed
thatconservation research
on invertebrates
should
be greatly
expanded
on twomainlines:(a)
studies
on the faunasof habitats
of special
importance,
eitherbecausesuchhabitats
arewidespread
on related
NatureReserves,
or haveto be maintained artificially
(e.g.fendykes; hazelcoppice)
or an) subject
to regularinterference
(e.g.sheep-grazed
grassland;
rabbit-disturbed
Breckheath):
(b)
the selection
of a f'ecvy
or perhaps-only one.,
species
ef -special
interest
whichcanbe studied
intensively
in relation
to
habitatand theenvironmental
factors
influencing
numbersand
behaviour.The workon theLargeCopperwas.quoted
as an example
of sucha studywhichopenedtheway to workon related
problems
of
management
e.g.on watertablecontrol,
involving
theco-operation
of otherdisciplines.
The meetingsUggested
thatmanagement
studies
of thistype,theresults'ef
whichwouldhavewideapplication,
should
be definedas Partof thenational
conservation
policyreferred
to in ThemeI.
ThemeIII - The valueof invertebrate
surves andhow the should
be developed.
Therewas generalagreement
thatthe-present.inadequate
knowledge
of theinvertebrate
faunaon NatureReservet
was a serious
handicap
to conservation
and thattheoriginal
intention
in Cmd.
7122to promote
basicsurveyworkof thistypeshould
be givenmore
support.Mr. Steel,Drs.MasseeandMorrisreferred
to thework-of
pastandexisting
surveyteams,theirachievemen%
theproblems
of
organisation,
finding
enoughspecialists
and adequate
finance.
Surveyworkon invertebrates
hadbeenconducted
in manydifferent
ways
withvaryingresults
but it was agreedthatthebestreturnformoney
spentwas obtained
froma specialist
or groupsof specialists
working to an agreedplan. Suchteamshad madeimportant
contributions
to a national inventory of wildlife
on NatureReserves.An analysis
of recordsfrom125N.N.Rs.,
F.N.Rs.,
andL.N.Rs.showedthatonly
threespecies
of Britishbutterflies
had notbeen foundon om or
moreof them,while28 out of 40 Britishbreeding
dragonflies
were
represented
on N.N.Rs.The meetingalsonotedthatcertaintypes
of surveyworkoutside
NatureReserveswas important.
4
and their
collections
of invertebrate
ThemeIV - The or anisation
use.
speciesand theircomThe largenumberof invertebrate
for consemtion staffto
made it difficult
plicatedtaxonomy
by Mr.
An investigation
documentrecordsand maintaincollections.
wheresomeattempt
T. HuxleyShowedthattherewere eightStations
but the extentand qualityvaried
was beingmade to formcollections
thata properlyorganised
nbly Therewas agreement
consider
and that•his
servicewas requiredin theConservancy
invertebrate
researchand invertebrate
Was a naturalextension
of the management
at least,two cenin ThemesII and III. Initially
surveydiscUseed
Edihburghand MonksWood,wherecuraiedinverte,treswere required,
couldbe builtup. It was acceptedthat
collections
bratereference
for NatureReservesalthoughsppciwere_required
vouchercollections
groUpe. The Conservationmenswere not neededfor all invertebrate
for theproposedcollection
be responsible
ResearchSection.would
RecordsCentre
of theBiological
at MonksWood and the co-operation
soughtfor documentation.
of the meeti are:The mainconclusions
After16 years,a carefulreviewshouldnow.bemade of
(a)
of Reservesfor
policyon selection
the Conservancy's
for researchinto
and of its commitments
invertebrates
on the Reservesalreadyestablithed.
menagement
researchon invertestudiesand fundamental
ManageMent
(b)
and plannedas
bratesshouldbe more closelyco-ordinated
policy.
conservation
part of a national
to the
contributions
survuysmadeimportant
Invertebrate
(c)
knowledgeof wildlifeon NatureReserves.thiswork should
for
by providingmoremoneyand facilities
be expanded
teamsof.specialists.
propurlyorganieed
was requiredin the Conservancy
.service
An invertebrate
(d)
at least,two centresshouldbe established
and initially
and vouchercollecwherecuratedinvertebrate-reference
tionswouldbe.he1d.
5
List of Particiants
BRIANEr. M. V., Furzebrook
ResearchStation,Wareham,
Dorset.
BROADHEADr.
D E., ZoologyDepartment
UniVersity
of Leeds.
CMISKTI,
Mr. R. N., The NatureConservancy,
Edinburgh.'
COILINGMr. A. W., The NatureConservancy,
London.
COPLANDMr. W. O., Furzebrook
ResearchStation,WarehaM,,Dorset.
tamus Dr. B. N. K., ToxicChemicalsnndWildlifeSection,Monks
WoodExperimental
Station.
DEMPSTEDr.
R J., ToxicChemicalsand WildlifeSection,MonksWood
Experimental.Station.
DUFFEYDr. E., ConservationResearch
Section,MonksWoodExperimental
Station.
EGGELINGDr. W. J., The NatureConservancy,
Edinburgh.
ELTONMr. C. S., F.R.S.,Bureauof AnimalPopulation,
Airzliversity
of.Oxford.
FRAZERDr. J. F. D., The NatureConservancy,
London.
GEORGEDr. M., The NatureConservancy,
Norwich.
GOODIERMr. R., The NatureConservancy,
Bangor,North-Wales.
HUNTMr. 0. D., "Wynstay",
The Fairway,NewtonFerrerO,
Plymouth.
HUXLEYMr. T., The gatureConservancy,
Edinburgh.
KERR Mr. A. J., Woodlands
ResearchSection,MonksWoodExperimental
Station.
LABERNMr. M. V., The NatureConservancy,
Wye,Kent.
MASSEEDr. A. M., 0.B.E.,"Acarina",
East Mailing,Kent.
MELLANBY
Dr. K., MonksWood Experimental
Station.
MERRETTDr. P., Furzebrook
ResearchStation,Wareham,Dorset.
MORGANDr. N., The NatureConservancy,
Edinburgh.
6
MOOREDr. N. W., Toxic Chemicaland
s Wildlife
Section,
MonksWood
Experimental
Station..
MORILRTY
Dr. F., ToxicChemicaland
s Wildlife
Section,
Monks.Wood
EXperimental
Station.
MORRISDr. M. G., Conservation.Research
Section,
MonkSWood
Experimental
StatiOn.
PENDLEBURY
Mr. J. B., MonksWoodEXperimental
Station!
PERR1NG
Dr. F., Biological
RecordSCentre,MonksWoodExperimental
.Station.
PHILLIPS
Mr. J..13.,
TheNatureConservancy,
Wye,Kent.
POLLARDMr. E:, ToxicChemicaland
s Wildlife
Section,
MonksWood
Experimental
Station.
Dr. J. E., MbrlewooResearch
d
Station,
Grange-over7Sands,
Lancs.
SKELLAMNi:
TheNatureConservancy,
London.
STEELMr.W. 0., Imperial
College,
Silwood
Park,Sunninghill,.Berks.
STEELEMr. R. C., WoodlandResearch
s
Section,
MonksWOodExperimental
Station.
TANTONDr.M.,WoodlandResearch
s
Section,
MbnksWoodExperimental
.Siation.
THOMPSON
Mr. J. A., TheNatureConservancy,
ShrowSbury.
UNGLEYMr D C., Furzebrook
Research
Station,
War.oham,-Dorset.
.JELCH
Dr. C., WoodlandResearch
s
Section,
MbnksWoodExperimental
Station.
Nr..T.C..E.,Conservation
Research
Section,
MonksWood
ExperiMcntal
Station.
-7
OPEN-ING
ADDRESS
INVERTEBRATE
CONSERWTIONIN RELATION
TO THE CONSERVANCY'S
COMAITIMS AND THE DEVELOPMENT
OF ITS RESEARCHPROCRAV1E
E. Duffey
Staffsymposiato discussConservancy
scientific
commit-.
mentsand problemsduring•the
last15 yearshave onlybeen-organised
in relationto Woodlande(Bangor,llth-13th
April1961) and Soils.
Conservation
work in whichinvertebrate
studiesare of importance
havebeen discussed
by scientific'staff
from time to timebut usually
came to nothing.because
of 'thedifficulty
of appointing
sufficient
taxohomic
specialists.In 1964, thisqUestion
was raisedagainby
Mr. TomHuxleyand•theConservation.Research
Sectionat thisStation
agreedto organisea staffsymposium
to discussthissubjectin the'
lightof theConservancy's
recordand moderndevelopments.
The themesof the Symposium
concernthe practical
problems
arisingout of managingdiversefaunason NatureReservesand other
areasand-theorganisers
have not included
papersdealingwith
ecological
theoryexceptwhereit is relevantto the application
of ecological
knowledgeto wildlifeconservation.
The programme
beginswith a surveyof theConservancy's
plansand thinkingabout
invertebrate
conservation
in its earlyyearsand its achievements
since1949.Thisis followed
by an examination
of invertebrate
conservation
problems,
both on NatureReeerves
anden unprotected
areas,followed
by a sessionto assessthe objectsand valueof survey
work. Finally,we Shalldiscussthepurposeand use of invertebrate
collections
in theConservancy.
Duringthe finalmeetingon Thursdayafternoon,
we hope
everyonewill takeadvantage
of the discussion
to pool theirideas
and suggestions
for futurework, particularly
in relationto the
Conservancy's
scientific
work and the contribution
it is making
to invertebrate
conservation.
In the following
account„Iwant to_review
some of the
historical
aspectsof theConservancy's
work in thisfielaand to
examinethe contribution
invertebrate
studiescan make to broader
problemsof wildlifecanserv-tion
as part of the totalresearcheffort
byvourscientific
staff:
As a basisfor thisreview,I am indebtedto-Captain
Diver
for a seriesof noteson the effortsto get the requirements
of
invertebrates
includedin the proposed
NationalNatureReserves
beforetheConservancy
was broughtintobeing. In 1943,theBritish
Ecological
Societyproduceda listof 66 majorhabitattypeswhichit
thoughtoughtto be includedin reserved
'areas.This .hasbeen largely
accomplished
and wouldprobablymeet;in mestcases,the main entom-,
ological
requirements.
In 1945,the finalreportof the Nature
ReservesInvestigation
Committeetriedto strikea balancebetween
the claimsof the NOTiolls
scientific
interests
and.18siteswore
listedprimarilyfor entomological
reasons:CadtorHenglands
BrauntonBurrows
ChiPpenhafl'en
HawesWater(SilverDale)
Monksjtioed
Ham StreetWoods
Blec;n.
Woods
Basingstoke.Canal
Wye and CrundaleDowns
EglwysegMountain(Dehbigh)
DennyWood and Bog
Deal Sandhills
MordenBeg
WickenFen
HeathsfromStudland'to
Arne Woodwalton
Fen
BirdlipWoods
WindsorGreatForest'
So littlewas knownaboutthe distribution,
ecological
requirements
andpopulations
of inVertebrates
thatthe onlyway of
dealing,
with the problemof/conservation
was to maintainthe greatett
varietyof habitatson NatureReserves.However,the Nature-Reserves
Investigation
CoMmittee,viere
awarein 1945 thatseleetion
based,:
solelyon vegetation
types'would
fail to save manyrare andlecalised
populationá..The 1945 reportled tothe firstofficial
bedy whichwas
appeinted
by the Ministryof.Tpwnand CountryPlanningwithSit
JulianHuxleyas Chairman...Ali
the 18 entomological
sitesmentioned
were inclUdedand'theoriginal
list of.N.N.Rs.
was expanded..1i.•
largenumberorscientificand naturalhistorybodieswere inVited
to suhmitevidence,
but for somereason,one of the few which
declinedwas.theRoyalEntomological
Society.CaptainDiver thinks
thiswoS.parily.due
to the entomologists
not forminga Coherent
integrated
body of opinion; therewere-too/many
diyerseinterests
and therewas a fairlygeneralfeeling,at.the.time
thatinverte-.
brates.cou74
look-after
themselves.However,thereis an undated
list of'158sitesdrawnup.bytheRoyalEntomological
Societywhich
»is not montioned
by theHuxleydoMMittee..Manyof..the
158 sites
are
included'in
the presentN.N.R.totalof 111, in spiteof
the fact thatonly18 have any reference
to invertebrate
interest
in the Reasonsfor Establishment.
Selection
of sitesfor invertebrates
21,000stpecies
of inSectsof which
-- Thereare approximately
about5;000are'determinable"
in the Sensethatthereare Modern
or useablekeys. Some groUpsare-verymuchmorepepularthanothers
so thatentomological
NatureReserveshavebeen proposedlargelyon
thebasisof recordsreceivedfromamateursor professionals
behavingas amateurs.Thesegroupsare mainlyLepidoptera
or
Coleoptera
withHymenoptera,
Odonata,Orthoptera
and Neuroptera
wellbehind. The completeness
of invertebrate
knowledgefor Nature
- 9
Reservesalsovariesverymuchand is probablymuch-better
for sites
in the.South,
Sputh-Zastern
and EastAnglianregionsthanfor most
placeselsewhere.
The Woodlands
Symposium
unfortunately
onlydealtuith
treesand did not attemptto analysewoodlandentomological problems.However,11 of the 50 woodlandNatureReservesand Forest
Nature.ResLxves
have an entomological
interestquotedfor them.
Othermajorhabitatsdo not comeoff so well e.g. thereare 7
ChalkN.N.Rs.but onlyone has a referenceto entomology;112
S.S.S.Is.
on chalk,but only11 have a 'reference
to invertebrates;
perhnpsevenmore striking
is the fact that thereare 21 chalk
S.S.S.Is.
in a countylikeKentwhichis well workedentomologically
and not one of thesehas any reference
to an invertebrate
interest.
Similarly,
thereare 18 ChalkS.S.S.Is.
in Wiltshireand no referenceto invertebrates.
-Dr.-Frazer.
willbe surveying
the actualrecordand fate
of sitesoriginally
proposedfor entomological
purposesbut it is
clear.
that-the
Nature-Conservancy
has no properlydefinedpolicy
towardsinvertebrate
conservation,
nor has it any clearideaof the
typeof researchrequiredto ensurethatNatureReserNemarmgement
giveeadequateattention
and resourcesto thisproblem.Beyonda
generalguideto conserving
habitatvariety,theConservancy
probably.
does not knowwhat it is tryingto conserveor what-itshould:be
conserving:.However,
untila-clearanswerto thiscan be foundi
mostecologiStS
will agreethatit is unlikelythatAherewillbe
any seriousomissions
or errorsif the 66majorhabitattypesproposed
by theBritishEcological
Societyare represented
on the national
seriesof NatureReservesand thatwithineachhabitattype,,
there
is a uide range_cif
variation.
The Huxley'Committee
intendedthatReservesehoUldbe
managed.Thiswe mustassume,includedinvertebrates
as well as
otherformsof wildlife.Their1947 WhitePaper includedthree
mainproposals:(1)
to establish
a seriesof N.N.Rs.: thishas moreor less
been accomplished.
(2)
to establish
a biological
service.
to .esteblish
researchinstitutes
in terrestrial
ecology.
(3)
.
This has alsobeen achieved,
at leastto someextent.
Theseinstitutes
were to be plannedon academiclinesand
to be independent.
The biological
servicewas intendedto accomplish
thefollowing:-(1) the settingup of an administrative
headquarters
thisis BelgravaSquare; (2) the establishment
of regionalorgan-.
isations- thishas alsobeen accomplished-.
- 10 . The biological
servicewas to includethreegroupsof
specialists
who wereneVerappointed.(1) •Scological
specialists
in majorhabitats- Woodlandsspecialists
were apPeinted
but these
were pert of the researchinstitutes'
staffand thusseparate
from'
the biological
service,(2) specialists
in groupsof organisms
of whichnonewere appointed,
and (3) specialists
in othersubjects
suchas soils,climatology;
physiography
- most of whom are now
in post. It is not generally
realisedwhat a significant
effect,
the failureto appoint specialists
in habitatsand groupsof organ-ismshas had on the development
of conservation
policyat national leveland management
policyon a NatureReservelevel. Thesespecialistswere not to be attachedto researchinstitutes
and in the
WhitePaperare referredto as "mobile".Theywere to workclosely
with the regional'staff
specifically
on practical
conservation
problems doingthe survey,studyand fieldwork necessary
to formulate
conservation
policy,and to produceecological
information
required
by regionalstaffwho were to be directlyresponsible
forNature
Reservemanagement.Invertebrate
stUdieawouldobviously
haveplayed
a:majorpartin the activities
of thisgroupof scientists
and many
of the.greup
spedialists
wouldpresumably
havebeen invertebrate
zoologist's.
The ConServancy!has'tried,:to
soma-extent,
to fill the
gap by thebuildingof MonksWood'Expuriffiental
StationWhiph
fically:concerns
itselfwith the fieldof appliedecologY.The-Co/1.
servation
ResearchSection-has
termsof reference
veryClOseto
these.which
we Cam assumewere intendedfor thanUssingspecialists
i.e. to,:exemine
the differenttypeaof NatureReservemanagement
problemS;.to'co-Ordinate•field
studiesOriNatureReservemanagement;
to do.research
on Practical
Conservation;
'•-to
providean.adVisory'
•
service'
:for±heconservation
staff. This coversmost of the sUbjects
concerninitheconservation-of
NåtureReserve.types
with the exception
:ofwoodlands,
and it is madeup of'two different.but
cOmptementaryfunctions:7
planning,
initiating
and doingreSearchrelevant,
to.
(4.5.
conservation
and forReservemanagement;
(b)
the settingup of a co-ordinating
and advisory'group.of
peoplewho would-bring
togethercommonproblemS•otmanage'mentstudies,
indexexistingrecordsand datareleVantto
consi:xv,tion,and
management,
and adviseon the application
of ecological-knowledge
to theConservancy'S
commitments.
_StUdies.and
Surveyahich'theConservancy.must
developin
relatiOnto'itScoMmitments,for
invertebrate
conservation
and
management
On NatureReserVesand Othersitesof conservation
interest
canbe.grOuped
basicpoints:- .
. aroundthe following
to prepareanationelinventory
of animalsrecordedon
(a)
NatureReserves; to,bringtogetherexisting
datawhichare very
scattered
and:promote
more surveywork on.the linesof.thatalready
bcing organised.The Huxley,
Committeerecommended
thatthe national
seriesof NatureReservesshouldincludeas Manyexamplesas possible
of all the manydifferent
groupsof liVingorganisms
whichformpart
of Our natural.fauna
and flora. We cannotmeasurethe successof
this.without,knowing
whatproportionof the'fauna
is represented
on
our NatureReserves.this information
is .1:Isebasic
to.management
.:programmes.
derivedfrom
information
a nationalindexof ecological
(b)
aspectsof
all
covering
unpublished,
and
published
work
existing
of
control
and
maintenance
the
to
relevant
knowledge
ecological
popuof
management
the
s
and
invertebrate
for
habitatsimportant
lationsand soeciesof animalson naturereservesor otherareasof
value. This wouldalsoform thebasisof the
conservation
staff. It wouldhelp to show
serviceto the conservation
.advisory
and forma guideto future
are,
knowledge
our
in
gaps
main
wherethe
ecologists.
s
applied
Conservancy'
by the
research
studieson the main factorswhich.have influenced
historical
(c)
habitatsin thiscountrytoday,e.g.use
major
of
the development
areassuchas theChalkdowns,Breckland
of
by man
and exploitation
and woodlands.Without
moorlands
pastures,
upland
fens,
heaths,the
the patternof
understand
to
possible
not
is
it
thisknowledge,
techtraditional
the
nor
today
survives
it
as
distribution
habitat
of
types
particular
maintain
to
centuries,
for
perhaps
niquesused
on
influence
profound
a
has
obviously
this
All
vegetation.cover.
s well as otherwildlife.
invertebrateas
(d)
problems
research
conservation
the nuMberof potential
is so greatand the numberof researchstaffto work on themwillbe
research
for a longperiodto come so thata Conservancy
inadequate
musthave an order
commitments
conservation
policybasedon practical
a seriesof
can be doneby recognising
of priority.I suggest.this
aroundthemain habitats
orientåted
conservation-projects
national
so thatundereachheadingare many sitessharingcommonproblems
as follows:-
habitatsof conservation
les of the mainland formations,and
on National
i ortance most of whichhave somere resentation
NatureReserves
heathS.
LowlandgrasSlerd'and
woods.
(a) Chalksites,excluding
s similarareasin southern
(b) New FOrestheathlandand
England.
•
and East Suffolk.
heatherheathsof Breckland
(c) Grassland
(d) SMellerunitsSuch as CannockChaseand Charawood
Forest.
2.
Wetlands.
(a) NorfolkBroads.
and lakes(e.g.the LakeDistrictand
(b) Reservoirs
ScottishLakes).
(c) Lowlandfensand mershes.
Englandand:
.(d) Bogs,mossesand Miresin northern
_
SCotland.
3.
Woodlands.
types.
woodsof different
deciduous
(a) Surviving
s
(b) Nativeconiferouwoods.
New Forest,
e
(c) Forestareasof specialimportance.g.
and otherman-madeforests.
Chasein Breckland
Thetford
Coastlands.
., shingleformations.
(a) Sand-duneand
(b) Salt-marshes.
barrageson
(u.g.effectsof proposed
(c) Estuaries
wildlife).
c
specialscientifiinterest
s
(d) Rockycoastlinewith
colonies).
(e.g.sea-bird
Uplands.
WalesandHighlands.
(a) Limestoneareasin Pennines,
(b) Grousemoorsand sheeppasturesinWales,northern
Englandand Scotland.
schemps,
d hydro-electric
(c) Mountainareasinfluenceby
tourism.
and
afforestation
6.
Agricultural.
(a) Hedgerows.
(b) Pondsand ditches.
(c) Crops.
-13-
The following
tableattemptsto showthatecological
studieson
NatureReservemanagement
are not isolated
projectsrelevantonly
to the particular
siteor regionwherethe work is beingdone:
Management
problemsof the same typemay be foundon a seriesof
verydifferent
NatureReserves-or
on typesof countryof contrasting
landscape.Seventypesof country-in
whichNatureReservesare
presenthavebeen takenfram the preceding
tableto Showthat the
sameland-usefactorsinfluence
conservation
problemsin many areas.
The resultsof researchon any one of thesewill therefore
have
wide application.
Conservation-Management
Problems
0
Grazingas a management
technique
++
+
+
+
(++ = well-establishand
ed widespread:+ = in someareasonly)
114
in the tablesrequirebasicsurveywork
All the sitesmentioned
studies.It is possible
as part of the management
on invertebrates
to the'problems
urgency
or
priority
however,to assigndegreesof
or modification
change
causing
factors
the
to
in relation
of management
different
at
change
undergoing
to the environment.The habitats
as follows:ratesmay be described
changeis natural,
.
where
stableformations
Ecologically
(1)
by humanor animal.
unafteeted
slaalyand relatively
proceeding
someduneand
saltmarshes,
Examplesare mountains,
interference.
and somewoodlands.
Shingleformations
degreeof vegetational
Habitatswhichhave a considerable
(2)
and some.
,
moorlands
grasslands
as
.such
uniformity
or structural
than
ly
ecological
compleX
less
.are
category
n-this
heaths. Reserves.i
be
may
burning
or
grazing
as
such
factors
Certain
othersalthough
uniformity.Publicuse not
the characteristic
in:retaining
important
signifiCant..
r
unstable.oheavily-ex
complex,relatively
Ecologically
(3)
wherechangeis, or mdghtbe, rapid. These
ploitedformations
whiehbecemeNatureReservesafterheavy
inclUdesoyewoodlands
trade
fen areaswherea fallingwater-tabland/or
exploitation;
o
pollution;
itionaluse is no longerpractised;hydroseres_subject-t
moorsbeing
ather
y
Scruhf-heneglected_grasslands-invaded-b
and
heaths,grasslands
by over-grazing;
convertedto poor grassland
dune sYstemssubjeetto intensepublicuse.
•
returnfor
obtaina more valuable
y
The Conservancwill
mainly
directed
studiesare
its researcheffortif its management
cases,
beth
In
(3).
(2).and
with categories
s.associated
to problem
greatly
be
coUid
field
the
in
work
al
studiesand experiment
ecological
certain
of
's"
"indicator
as
species
of invertebrate
aidedby making'use
of
nature
the
on
knowledge
, developing
conditionsof
environmental
g
classifyin
of
methods
the
of
s
animalcommunitieand
invertebrate
animalhabitats.
--summApxcan be of greatvalue
studieson invertebrates
Conservation
1.
s.They can
exPoriment
field
researchand ecological
in management
conscientific
national
a
of
part
y
ap
onlybe plannedeffectivel
servation.policy:
workwhich mustbe done is
The typeof appliedecological
2.
apartfromMonks
stations
research
of the
not the responsibility
in theRegions
staff
on
Conservati
the
by
Wood. It cannotbe done
for the present
subject
a
vast
too
is
It
s.
who have othercomadtment
and can
impaet
effective
any
make
to
ResearchSection
Conservation
to the
ecologists
applied
more
t
of
appointmen
the
by
onlybe solved
s
working
specialist
"outside"
of
use
greater
y making
Conservancand
to an agreednationalplan.
- 15 on naturereservesand other
commitments
The Conservancy's
3.
researchcan be
and the associated
areasfor wildlifeconservation
groupedundersix mainheadings.Thesesharea nuMberof common
modify,causechargeor maintaincomponent
factorswhichinfluence,
partsof the habitat.The sites,in relationto thosefactors,
urgency,and wouldenable
can be assignedto an orderof management
policy.
conservation
to framea scientific
theConservancy
constitute
we shalldealwith at thisSymposium
The subjects
will no
and complexproblem.A gooddeal of discussion
a difficult
but I wouldrepeatWinstonChurchill's
afterwards,
doubtbe necessary
advisers;"Let'snot talkof.the
to his scientific
instruction
Let us
will speakfor themselves".
the difficulties
difficulties;
ideaswhichwill enhanceand
spendour timelookingforconstructive
researcheffort.
developtheConservancy's
Summar of Discussion
It was pointedout thatlittlewas knownaboutinverteexceptthatman was
brates.inthe earlydeys of the Conservancy
in-whichtheylived-.The Conservancy
thewholeenvironment
upsetting
with onlyvery
programme
however,had to carryout an acquisition
And theydid thebest thatwas possibleat the
limitedknowledge
not
time. Dr. Nasseesaidthatthe reasonwhy the ontoMelegiata.did
Committee
submitevidenceto theNatureReservesInvestigation
in the 1940swas becausetheirnuMberswere madeup chieflyof
Theseactedas a kind of closedfreemasonry;they
Lepidopterists.
was kept
but thisinformation
knewwherethe greatraritiesoccurred
for British
Committee
to themselves.It was not untiltheProtection
W..G. Sheldonthattheybegan to let others
Insectswas formed-by
thu firstlistsof insectsto be protected
sharetheirknowledge'and
were drawnup in 1946.
by otherrcMbersthat surveywork
The viewwas expressed
or
administration
was not givenhighpriorityby theConservancy's
to bc intellectually
becauseit was.not considered
scientista,
e
surveys-on
thr,t-invertebrat
satisfying.It is pointedout howevur-,
whemplan-i
value
land are of particular
certaintypesof agricultural
are.2na of the commonest_
ningresearchprojects.Althoughhedgerows
aboutthe animals.
isknown
verylittle
habitatsin our countryside,
to have a
needed
we
that
emphasised
MeMbers
whichlivein them.
beenengaged.
had
Conservancy
The
Paner.
freshlookat thc 1947:Miteyet taken-a
not
has
it
and
years
fifteen
of
a.period
for
on research
to ,
need
We
achievement.
scientific
of
record
its
on
lookback
White
the
in
outlined
were
plans-which
ord
the proposals
re-examine
in-the
do
to
out
set
we
what
achieving
we..are
whether
Paperand see
of the;
the effectiveness
beginning.SeveralmeMbersquestioned
existence
in
been
LiaisonCommitteewhichhas now
Entomological
for sevenyears. Therewas a generalfeelingthatits recordwas
over
not verygood and thatit had failedto takeenoughinitiative
wildlife.
invertebrate
our
concerning
issues
important
- 16 INTEREST
THE'FATEOFAREAS:OFINVERTABNATE
1949
AS LISTED.IN
J. F. D. Frazer
London.
The NatureConservancy,
liesin the reportsof the
The basis Jf thisknowledge
heldby theConservancy.Thesereports
localN.R.I.sub-committees
accomvaryin qualityand are devoidof the mapswhichoriginally
missingbut muchof it
paniedthem. The originalquestionnaire'is
can be deducedfromthe replies. It is apparentthatcommittees
educational
for habitat,species,
were askedto suggestareassuitable
onix)thnationaland localscales. In addition,
and amenityreserves,
entomological,
whetherthesesitescoveredbotanical,
they'li'sted
or mammalinterests.Somerepliesare
ornithological
geological,
whileothersaro briefer,and a few
statements
'considered
carefUlly
Lancs.,and
more listsof names. The reportson Regions6 (South
are missing:Only one siteis given
Cheshire)and 16 (Oxfordshire)
Roman
interestotherthanontomol:Dgical.(the
invertebrate
as'having
Snail,Helixpomatia).
Theserenortsand Cmd. 7122have formedthe bksisof the
at
Table. Where areasare listedas havingno statutory.protection
present,the causesare varied. Soma of the woodshavebeen clearfelledwhileono has gone down to housing. Some areasare ploughed,
otherspublicopen spaces. Some are at presentunderscientific
and othersproposedas S.S.S.Is.Thereis a smo11
investigation,
neithertheRegionalstaffnor localnaturnuMberof sites.where
or
have anY knowledgeof existingentomological
.alists
interest,
interestat all.
evenof'anyscientific
WhileWalesis coveredby-theN.R.I.C.reports,only
interestare mentioned.Scotlandis
six sitesof entomological
but in Cmd. 7814,fourareaS.are
not coveredby thesereports,
("Cerinterest.Theseare the Cairngorms
listedas of invertebrate
insectsare peculiarto the
of •
tainspecies
or arebetterrepresented
plateau
Cairngorms
here thanelsewhere");
theBlackWood of Rannach,now an S:S.S.I.("Thearea is of general
interest;its insectlifeparticularly
so");AlitVolagir("The
nowan
also of greatinterest")
life is
insectwherethe interestis listedas purelybotanical;and
5.1.
3.5.
Craigs'andRacksMoss (Itsinsectlife is especially
.worthyof
now denotified
part.wasformerlyan S.S.S.I.,
.vdtere
furtherstudy")
as substandard.
- 17 Table- showingfateof Englishsiteslistedby N.R.I.C.committee
or Cmd. 7122as of entomolo.ical
interest
Summar of Discussion
It was statedthatverylittleseemedto be knownabout
the invertebrate
interestof naturereservesin Scotlandand that
reference
to suchan interestin management
planswas veryscanty.
Each yearnew raritieswerebeingdiscovered
in Scotland,
so thatit
was difficult
to make a properassessment
of the entomological
interest.In no casedoes a Scottishmanagement
plan makea recommendation
for conserving
invertebrate
speciesor populations,
or for
conserving
an area for thatend. It was felt thatScotlandshould
be regardedas an areawhereit is moreprofitable
to conserveand.
do surveysbecauseScottishnaturereservesare not subjectto the
publicpressures
foundin southern
England. The post of Historical
Geographer
is also one in whichScotlandhas considerable
interest
becausea trueunderstanding
of populations,
both botanical
and
zoological,
is very dependent
on what the past treatment
of the land
has been. It was pointedout thatthe Scottishentomology
in
the mid-1940s
was a lot betterknownthanis reflected
in the management
plansof Scottish
NatureReserves.Perhapsif Scotlandhad agreed
in 1947 to combinetheirrecommendations
with Englandand Walesin
Cmd. 7122,the zoological
interestof theirsiteswouldhave
receivedmoreattention.It was said thatin Wales,onlyaboutsix
siteshad any entomological
interestrecordedfor them. Almostall
countiesin Waleshad scantyentomological
information
with the
possibleexception
of Glamorgan
whichis the best-known.It was
addedthattherewas no provisionfor the management
of the entomologicalintereston any of the WelshReservesat the presenttime.
A meMbersaidthatin Scotland,
most of the entomological
recording
had been doneby Englishentomologists.
Many of the recordsare not
localised
to specificareasand in many casesonly the rarities
havebeen recorded.Dr. Masseepointedout that a greatdeal of
information
aboutthe insectfaunaof Scotlandcan be foundin the
diariesof a Mr. Harwoodwho livedin AviemorejustbeforetheWar.
— 18—
abouttheColeoptera
His diariescontaina vastfundof information
of theHighlandsfrom1923onwards.He saidthat the diarieswere
and on
Brownof Bournemouth,
of Mr. Scarsdale
now in the possession
Oxford. If the
his death,wouldbe handedto theHope Department,
Easseeofferedto make
wouldlike to borrowthem,-Dr.
Conservancy
arrangements.
the necessary
of species
thenturnedto the conservation
The discussion
on the edge of theirrange. Dr. Satchellsaid thatin his talk,
.of
on the edge
is-conserved
he wantedto ahow thatif a.species
its range,theproblemof ensuringsurvivalis muchgreater
a largedegreeof interference..
becauseit is unableto withstand
a good
near thecentreof the speciesdistribution,
But elsewhere,
withoutdanger. He
canbe,tolerated
interference
deal of huhaah
of NatureReservesfor
whetherthe establiahment
questioned
specieson the edge-oftheirrangewas worthwhile.OthermeMbers
and gave as an exaiple,the
point.of.miew
expresseda different
an insectwhich shouldbe conserved.although
LargeBlue—butterfly,
it is on the edge of its rangein thiscountry.
-1 9-
- S6ME
INVASIONS
EXTINCTIONS'AND
AND CONCLUSIONS
CAS HISTORIES
J. E. Satchell
ResearchStation
Merlewood
SUMMARY
faunaas a etatic
The conceptof a Britishinvertebrate
Evidenceof
history.
its
bf
with the facts
entityis inconsistent
fossil
by.the
provided
composition'is
its
of
nature
the transitory
the-last
hin
.distributions-wit
recrd andby changesin species
century.
or near
of extinction
of the casehistories
Inspection
wide
suggests
documented
adequately
species
few
of the
extinction
contin-.
arctic/alpihe:and
betwebn
balance
changesin recentyearsin the
becruse'they
sbecies,
continental
Britain,
In
entalcomponents.
to clim-.
vulnerable
are on the edge of theirrange,are particularly
risk.
conservation
good
a
not
are
term,
aticchangeand,in the long
measures
conservation
Whereby
sted
approach_is_Sugge
An international
.the-centre
toWerds
tie-taken
protection.wouid
for a species.needing
responsbe a.special
of-its,.range.yIrctic-alpine-species.would
Britain.
of
ibilitY
I. INTRODUCTION
may be distinguiShed:
conservation
Two'aimsof invertebrate
of particular
(2) the protection
of diVersity;
(1) the maintenance
willbe conattention
aim,
primary
the
is
former
the
If
sPecies.
changesin
large:scale
of
invertebrates
on
effect
the
on
centrated
controland the brelanduSe,generalmeasuressuchas inseeticide
seriesof plantassociations.Inpracof a representative
serVation
to
willbe largely incidental
conservation
invertebrate
tice,
reserves
nature
of
Management
the
to
and
preservation
'amenity'
mainlyforbetanicalreasons.The lihitedpUbliclappeat
selected
may dictatesucha policy.
of invertebrates
speciesrequiresa
particular
A policyof preserving
and criteriafor judgingwhere
definedbasisfor theirselection
.This
measuresare mostlylikelyto be effective.
conservationpaperoffersevidenceon theselatterpointsfrom specieswhibhin
so and from otherqbeCie's
Britainhavebecomeextinctor netirly
of theirhabitats."
whichhave survivedseveredisturbance
- 20 -
II CHANGESIN THE COMPOSITION
OF THE INVERTD3RATE
FAUNA
THE FOSSILRECORD
The fossilshellsof Molluscaprovideevidenceof
invertebrate
historyin Britainsincethe late-Glaciafl
(A)
The illustration
given(Fig.1)showsthe speciessucessionfrom the sub-Boreal
to historictimesfroma siteon the North
Downs. The succession
startsin the Mesolithic
witha community
indicative
of wet closedwoodlandwith shade-loving
speciese.g.
Car chiumtridentatum
and Discusrotundatus,
and marshSpecies,
e.g.Limneatruncatula
and Succineaoblongadominant.Duringthe
Neolithicand apparently
associated
with forestclearance
the woodland speciesgaveway to grassland
species,at firstof a kind that
avoidsbare downlandand then,duringa dry period,to xerophils
suchas Valloniaexcentrica
and the nowextinctMonachacartusiana.
Helix asperse,fumy/commonspecies,arrived-probnbly
from the
Europeanmainlandat:about250-300A.D.
MOnachacantiana,
now
generallydistributed,
arrivedapparently
in theRomanperiod.
L similarpatternof chargeis evidentfrom fossil
Coleoptera.
(B)
(1)•
RECENTCHINGES
Ches
associated
with climatictrends
(4
The,nearextinction
of the wood ant,Formicalugubria•
in Ireland.
e storyo • ugu ris in re.an provi
.es an
exampleof the extinction
of a speciesfor reasonsapparently
unconnected
with'human
activity.On the mainland,
thiswood ant is
an uplandand northernspeciesgenerally
confinedto ereas.over
1000
feet abovesea levelin Wales,thePennines,theCleveland
Hills,
theLake Districtand the ScottishHighlands.In Irelandit occurs
onlyin the southwherealsoit is foundmainlyin uplandareas(Fig.
2). It has been recordedfrom17 sites,the earliestrecordsdating
from the mid-nineteenth
centuryand the majorityfromthe 1920s.
Sincethen,the specieshas virtually
disappeared,
onlyone colony,
the most northerly
one,remaining
in 1963. On one site,the.wood
thenbeen felledwhilethe remaining
fifteenwereunchanged
(Collingwood,
pers.comm.).No colonyextinctions
havebeen reported
fromBritain.
hh.cl
It willbe seenfromFig.2thatall the extinctcolonies
lie on the warucrsideof the 42° Januaryisotherm
whilethe surviving
Irishcolonyand theBritishcoloniesare on its colderside. The
extinctcolonieswere in the part of the speciesrangein theBritish
IslesWhichhad thewarmestwinters. Moreover,
duringthe period
when the coloniesdied out,wintertemperatures
were aboveaverage
for a longer'period
thanat nmy time sincethe1730sandbeforethat,
sincethe 'littleclimaticoptimum'of the medieval
period(Fig.3,
Laub1965,-Monley
1965). The alliedspecies,'Formica
rufa,
Sub
Boreol
400
Circa 3000 B.0
Mesolit h ic
Neolithic
grassland
woodland
wet closed
CLEARANCE.
Damp grassland
ond scrub
Dry open
grassland
Damp open
grassland
herbs
with
(?)toll
Open
A.D.
Zonation
Environment
Bronze Age
Iron Age
RomanoBri tish
Clrco
ond
General
1.s.,
0
Fig.1
1111111111/J111i
0
60
40
20
0
0
20
'0
adopted
1_1_1 11,11111111
Sesuence of itkollusca at Devil's Kneading Trough
0000
402
715
1031
246
370
396
412
416
515
745
562
441
512
520
0
1.11-1.1.4 °/0
I
349
361
from Kerney et ol
1964.
á
Of
Distribution
the Wood Ant Formica tug4ris
cf
0
00
o
38°
Woodford
•:Q
0
42°
c
450
6
4 2°
means 1901 30
uplands over t000 feet approx.
vice County distribution
0 Surviving Irish colony.
Extinct colonies
Isotherms
°F January
Fig. 2
-
L700
3-65
1200
1730
1300
50
1400
1900
1500
1600
50
1700
1800
Mean winter temperature in England since A.D. 1150
1670 onwards instrumental.
1150 - 1670 conjectured.
-4
-3
50
(Dec. - Feb)
19001910 1920
1900
Fig 3. Number of Successive winters with mean winter temperature
obove the overage far 1661-1963
1580
2
.3
4
5
6
7
9
10
11
-
- 21 to producesexualformswhen kqptunderisothermal
conditions
(Gosswald
andBier 1954). Collingwood
(1958)suggeststhata similar
reproductive,failure
caused the disappear-nce
of F. lugubriSin Ireland
and gkotesin:SUppert
an observation
of Stelfoxthathe foundno
sexualfernsin coloniesin Wicklow*.sited
in the years19217726.1f thisexplanatiOn-isCorrect,
it,SeemsthatF. lUmUbrismust havearrivedin Irelandsincethe medieval
periodas it couldhardlyhave
surVivedtbe mildwintersof the twelfth,thirteenth
and fourteenth
centuries.
(b) C
es in the distribution
of certainLe idomtera.
A possiblycomparable
case is thatof the Mountain
Ringlet,Erebia
emimhron,
described
(Ford,1946)as 'our onlytrulyAlpinebutterfly'. It was recordedin Sligoand Mayo at the turnof the century
but is now apparently
extinctin Irelandand survives
onlyin the
LakeDistrictand Scotland.
.The.moth,
Hyppa-rectilinea,
a northernspecieswith a
rangeextehding-to
Siberia,-wad
at ohe'tinie-Tönhd-in'Yerkbut
shire
has likewise,withdrawn
northwardsnnd
is not now foundsouthof
CuMberland..
Duringthe sameperiod,severalspecieswith a southern
and continental
distribution
haveextendedtheirrangenorthwards
e.g. thette Admiral,Limenitiscamdlla(Fig4) or have established
theirfirstrecorded
breedingpopulations
in Britain(Fig.5)..
(c)--Cha es in the diStribution
of littoralHs
Changes-in
sea-water-temperatutes.linked-rvith.the
changesin'air-temperature:referred
to aboveare'deScribed.bymanfaUthers. Lamb
and JOhnson(1959),from an examination
of all the available
data.
on Arcticice,reporteda longperiodof minimumamountin the north
Atlanticfromabout1920 to the late1940s. Crisp(1965)calculated
a shiftof about80 kilometres
in the isotherms
of theEnglishChannel
betweenthe 1930sand 1950s. Corresponding
with thesesea temperaturechangesthe.distributions
of northernlittoralspecieshave
contracted
and.southern
epedieshave'ektended
horthWards.
Betweenthe 1930sand 1950s,the kelpLaminariaochroleuca
and the herMit.crab
Clibannrius
misanthromus
extendedtheirrange
from.the-coaSt
of Franceto the southof Britain. The topshell
Monodonta
lineataeXtendednorthwards
fromCardigan-Bay
to Anglesey
while the,northern
dog-whelk
Nucellalapillusretreated
northward
from the.wrMer.parts
of the Gulf of Gascony.
Between1934and 1951,the arctic-boreal
barnacleBalanus
balanoides
allbut disappeared
from tbe•south-west
Shoresof Britain
and its placewas takenby the more southernspeciesCthalamusstellatus
(Southward
1958). Withinthe territory
commonto both species,they
monopolised
almostAbe wholeintertidal
zoneof exposedcoastsand
competedstrongly
for smace. Extremetemperatures
cppearto have
had littleeffecton the abundanceof eitherspeciesbut whereas,
- 22 B. balanoides
can breed onlyafterit has been pre-conditioned
for several-weekS:at
temperatutes.below.10°C.,
C. stellatus
requires
temperatures
exceedingabout
15°Cin orderto.breed.Thuswham
conditions
favour.Cthalamus
and.cold
conditions
Balanus,,
wherever
they are in competition
(Crisp1965). The examplefurtherilluS
tratesthe risk of extinctien
of speciesat the edge of theirrange.•
(2)
Changesassociated
with excertional
weather
The distribution
of manyinvertebrates
thoughcontrolled
in partby climate,may alsobe determined.by
shortspellsof extreue
weather0C-burring
very-infrequently
but persisting
in theireffects
for many'years.The winterof 1963,in theEnglish-Lowlands
the
coldest.since
1740 (Laub1963),providesa good exampleof such
effectson invertebrate
populations.
(a) The effectof winter1963 on littoralinvertebrates.
The effectsof the extreuecold of the earlymonthsof 19 3 on the
littoralfaunaof.the coastsof Dorsetto Hampshire,
theBristol
Channeland Caernarvonshire
havebeen rePertediby
CrisPand coworkers(Crispet al 1964). Llthoughmany speciessuffered
heaVY
mortality,
few sufferedsignificant
and pronounced
changesin their
range. In general,sessileformswere ableto tolerateverylow
temperatures
and thosewhichwere not able to toleratelow temperatureswere able to moveinto the warmersub-littoral.
Exceptionally
heavymortality
was nevertheless
rdcordedin the oysterL.nomia
ephiprium,
thetopshellsMonodonta
lineataand GibbulauMbilicalis
and the crabPorcellana
lo ,icornis.The largetopshell,
Monodonta
lineatawas-completely
destroyed
in LymeBay in SouthWalesand in
Tremadoc.Bay
by th6suddenonsetof-severe
frosts,but as the populationsof the more maritimepromontories
of southDevon,PeMbroke
and North-Wales
surVived,
thereis no reasonto supposethatthe'
bays will not eventually
be re-populated
fromtheseseurces.
(b) The effectof winter1963on freshwater
molluscs.
The data.OfFig.6are fromunpublishedrecordsof a naturalist
with
many yearsknowledgeof the Molluscaof the Southwell
district.
They referto Sine freshwater
bodiesincluding
threecanals,two
ponds,a lake and a riverand comparethe speciesof molluscs
recorded'tetween.-1949
and1963 tith thoserecordedin 1964. Between
1949and 1964,two of the sites.had
been filledin or drained.Of
the 29 specieswhichhad beenplentihl
Or-another
of the sites,
only one,Flanorbisleucostoma
was stilla bundantin.1964in any
site. Fourteenspecieswere.still.present
but scarceandfourteen.
wore not foundin any of the sites. The-listof sreciesrecorded
fromCossallCanalwas reducedfromsixteento seven. Siteswhich
initiaily'had
few specieshad nonein 1964.
-Thedata may be comparedwith thoseof Boycott(1928)
who recOrdedthe molluscSpresentin 76 closedpondsin 1915'andagain
in 1925. From the 18 speciesoccurring
in the 76 ponds,lloycOtt
made168.species-pond
recordsin 1915. In_1925,he made 93 new records,
108 were the saneas in 1915 and 60 of the originals
were not found.
•
•
LA
A
se'•
t,A
AA
A A
A
•
L
,
AA
A
A
Fig. 4
• • •AA Atså
••
• • ••
å
A
6 _at.
å
•
a
of range since 1920
Distribution prior to 1920
Extension
From FOrd 1946
The White Admiral
(Limenitis comillo)
o
6
1. Lithophone leoutieri
2. Eupithecio phoeniciato
3. aupithecio
4. Hydroecio
huollerordi
5. Xonthorhoe birivioto
6. Colomio tridens
•7.Sedina bUttne,ri
S. Eupi thecia egenorio
2
If
a
7
1
Fig
33
3
3 5
53
34
Distribution
S.Fronce, Spain, N.Africo
S. Fronce,S.Spain, Italy, N.Africo
Centralr S.Europe, Asia Minor
France
.France, Central r E.Europe to Siberia
Europe, Finland, Italy,. Coucosus
N. Europe, Fronce to Poland
Breeding populotions of moths
'New to Britoin'estoblished 1939-62.
Effect of viEnt
r 16
on some freshwater
molluscsin Notts & Lincs.
_0..
.11
.,3. w0,
c
0-
c
1
o
SPECIES
am
A
8
g'
c =
am
al
0
.0
,... t.
4 ,...
. ,...
2 S
0
MORDED AS PLENTIFUL
>
re
g
1949-1963
5
i
'i 8
m
a
el
0
0
T3
c
0
a.
-..-.
.1
.0
r-I
'5
''''
0
00
c
5
8
›.
A
i
'&1 'RI
DS
—1 8
g ic
0
c
;
0
0
z
STLTUS IN 1964
Planorbls leucostaan
SphaerIum lacustre
Lynnaea stagnalls
Planorbls vortex
Anodonta anima
Anodonta anatlna
Segmentina complanata
SphaerIum corneum
Physa fontinalis
Planorb!s contortus
i
---.
Abundantrr
S
S
S
S
S
S
S
S
S
S
Lymnaea
peregra
Planorbls
carinatus
V1v1parus
viviparus
Planorbarius
aorneus
Sythynia
tentaculata
Vliaparus
fasclatus
Unto pictorum
Uniotumidus
Sphaerium
rivicola
.PisIdium
amnicum
Sythynia
leach!
LYmnaea
aurleularla
Planorbls
planorbis
Planorbis
albus
LYmnaea
truncatula
Plancrias
laevis
Plancrbis
crista
Physaheterostropha
Drelssena
polymorpha
S
S
S
S
S
1
15
: Species
S
1
C
c0
-6
o
S = Scarce
- = Not fcnnd
Fig
6
c
-0
14
0
-6
Species
-4
Ir.
á
Distribution
Boy.
of lowlond peot mosses north of .Akorecombe
of Plebejus orgus var mosseyi
— The hobitor
.
...
. .
tOri
Mo s
Helsin
BRIGSTEER.
• Nichors
M055
Holker
11.4M0.5
:
•ifrAeothop
Moss
fitorecombe
Bay
Relic Feat moss in 1954
and F.ostglacial alluviumand peat - probable
former distribution of peat moss.
Recent
*
Sites from which vor mosseyi was recorded.
Fig. 7
/*
Foulshow
M055
.
_
-2 3-.
Boycottconcluded
thattherewasa considerable
amountof coming
andgoingbetweenpondsbut anydeficiency
in thecollecting
technique
wouldtendto givean exaggerated'impression
of Suchmovement.In
theNotts.andLincs,data,onlyone species
was foundin 1964
whichhad'noi-been'taken
fromthesamesitebefore:,
whereasBoycott
mademorethanhalf.asmanyrecords::again'on-his
Second.survey.
The
impression
is gainedthatthewinterof 1963 certainly
reducedand
perhapsexterminated
themollusc
populations
of manyfreshwater.
bodiesand thatbeéauseof theirgreaterisolation,
theirrecolon.isation
ia likelyto be considerably'slower
and morefortuitous
than
in intertidal
habitats.
Interaction
of habitatdisturbance
and exc tionalweather
(3)
Although
exceptional
weathermayalterthebalanceof invertebrate
populations
formanyyearsandperhapsformanydecades,
it seems'
unlikely
thatexceptional
weather,
outsidetherangeof whathas
beenexperienced
by the faunasincethepostglacial,
willnowcaUse
theextinction
of any nativespecies.Many species
however,
mustnow
be at riskbecausethe extentof theirhabitathasbecometoo small
to Providerefugesfromwhich'they
couldbe recolonised.
The extinction1
or nearextinction,
of theracemasseyiof theSilver,Studded
Blue
(Plebejus
argus)illustrates
theprocess.
(a) The.disappearance
of var.masseyiof theSilverStudded
Blue. The racemasseyiof thisbutterfly
is distinguished
mainlyby
largepatchesof blueon thewingsof-thefemalewhichin the typical
formalebrown. Its recorded
distribution
is.restricted
to the socalledmossesor lowlandmiresof Morecambe
Bay,particularly
Meathop
andHolkerMosseswhereit was muchsoughtby collectors.
It was
lastrecorded
on thesemossesin 1921—When-,
in thethirdweekof May
whenthelarvaewouldbe nearlyfullyfed,therewas an exceptionally
severelatefrost.An eye-witness
account(Wright
1942) describes
how all thevegetation
on themosseswas blackened
up to a heightof
aboutninefeetwhileon thecragsaboutthirtyfeetabove,the
vegetation
was hardlyaffected.Unlessit is supposed
thatthisevent
wasunprecedented,
the question
ariseshow var.masseyisurvived
previous
lateMay frosts.Altho4ft
thereis-_no.published
recordof
the forgerextentof thesemosslands,
theaVailable-eiridence
suggests
thattheyoriginally
occupied
mostof the valleyand coastalareas
shownon theCeological
Surveymapas alluvium
(Fig.7) . In the
LythValleYfor- exaMPle,:Seattered-frågMehta.of-MOSSee;
nevi
tree
covered,
remain-as-far-inland.as
thevillageof Brigsteer-Wherel---..-.
several
families
madea livelihood
frompeatcutting
yithinthelast
halfcentury.At thispoint,themossl'yyuld
havebeenwellabove
thelevelup to which,nearthecoast,the Vegetation
was killedby
thelatefrostof 1921. It maybe supposed
thatinTrevioue
centuries,
thesevalleyheadsprovided
therefugesfromwhich,after
similar
frosts,the coastalmosseswererecolonised.
The butterfly
has in factbeenrecorded
twicesince1921, although
notwithinthe
lasttwentyyears- on bothoccasions
frompointson theinlandside
of theremaining
mosses.
24
The casehas practicalinterestin thatMeathopMoss has
now been made a localNatureReserve,largelyfor entomological
reasons.Althoughat one timethebest knownlocalityfor var.
as a reservewouldnot haveprevented
masseyi,its earlierdeclaration
extinction.
thebutterfly's
(4)
Survivalin relationto range
From the examplesquotedaboveof the antFormicalugubris
on the coastsof Southin Ireland,thebarnacleBalanusbnlanoides
West Englandnnd fromthe similarcaseof the ScotchLrgusbutterfly
it is
Durham,NorthYorkshireandWestmorland,
in Northuftberland,
at the southern
evidentthatnorthernspeciesare liable to extinction
end of theirrangewhenhumanpressureis slightor evenabsent.
end of their
The sameis trueof southernspeciesat the northern
range.
of the molluscEno_montane.
The slowerosionof populations
limit
Englandwhereit reachestheNorth-'21est
Oig.11) in southern
to the gradual
part attributable
,
of.itsEuropeandistributionis-in
e old woodlandand in rart to its failureto colonise
disappearancof
centurywhichnow
new woods,even thoseplantedin the eighteenth
1939). Many otherspecies
habitats(Boycott.
suitable
form apparently
well knownto be at risk in Britaine.g. theLargeCopperbutterfly,
the sniderEreseusniger,the snailMenacha
the LargeBluebutterfly,
are likewiseat the extremenorthernedgeof theirrange
cartusiana
to humanpressure.
althoughsomeof theseare also slibject
had recently
A numberof southernspeciesof moths.which
earlier.As
cited
.
in Britainwere
breedingpopulations
established
if they.
it woUld-b-esurprising
their-range
colonistsat the_edger.of
bioticpressures
of climatie.and
surViVödfor long.-The resistance
by the historyin Britainof
is illustrated
againstsuchinvaeions
rhododendri.Firstrecordedin
theAmericanTingidbug Stenhanitis
1939) was as
in 1939 For:Jilson
Britainin 1901, its distribution
sitesin Southfew
a
reducedto
Shownin Fig.8but is now apparently
East England.
of speciesat the edge
to extinction
The susceptibility
speciesseemcapable
many
with the pressure
of theirrangecontrasts
The recenthistory
range.
near the centreof their
of withstanding
in
of its rangeend
northern
of the wood ant Formicarufa near the
example.
an
Englandprovides
the LakeDistrictand.in South-East
on the wood ant
(a) The effectsof habitatdisturbance
range.
its
of
centre
the
rufa on the edge and near
Formica.
s Keswick,F. rufareacheSthe Northffest
Apart fromseme'colenienear
in Britainin the areaaroundGrange-overlimitof-itsdistribution
in woodlands
Sandsand Windermere.It occurstherein abundance
on the
in
colonies
and
scattered
s
4 few
on Carboniferoulimestone
Distribution
of
Stephonitis rhododendri in 1939
introduced from Euroye
in 1901[Fox-Wilson 1939]
•
•
0-
•
•
•
•
•
Fig. 8
•
•
• •
•
• 5• • •
.
ono"
•
•••
c
•
•
•
• •
• S 41
•
• •
•
0
L
Formica rufo in the South Loke District
• Coloniesextinct
since1949
since 1912
•••••
`SO
since 1934
since 1934
since 1924
eft
/
1920 1940
I
I
I
I
I
I
1920-1940
since
;\
I
1
I
I
I
la
/111P
1929
s6,
*
Isolated
45
colonies
Atoreccunbe
gir Colon es abundant
B oy
M Limestone
40-
—— Annual rainfall
or'
Fig. 9
á
Formico rufa in South East England
•
0
Southend
LONDON
Morgote
Co
Gravesend
Gillingham; ,
•••
,
/, •
„ .
•,./•, •
vnolasto
omsoo
-..s_
'1,;
Canterbury
IedhiII
`t.`
•
•
•C‘ •
2,7/.
'Tunbrld
"../0
eWells
'4:;;
41
/
%
Oyer
Folkestone
%;;%,
5,;2;
Hosting s
Hove
Worthing
Eostbourne
K EY
Extinc
Urbon development
10 Km
Ssuores
6
Unconfirmed
Probably
extinct
Confirmed
1962-4
31
7
Fig. 10
New Records
1962-4
10
á
Distribution of Mollusca as a factor in
selecting areas for their conservation.
(Aetapsfrom Kerney unyublisked)
("Aiajs a -»
Achonthinula lamellate
0
•
•
•
••
00
Fig.it a
Appropriate for conservation in Britain
á
Lourio anglico
17
•
dical
4.°
cl
Appropriate for conservation in Britain
Fig. it b
á
Heticodonto obvoluto
1
a
0
a
0
o
0
o
4
•
*a
o
0
ci
Conservation in Central Europe more appropriate
Fig. ll c
á
Eno montano
0
I
0
oel •
o
of l
4
•
IV
• •
•
•
•
4
0
Conservation
Fig It d
in
4
0
c3
Centrot Europe more appropriate
á
Catenella orenorio
0
•
do
4ci
Conservation desirable wherever possible.
Figm e
á
Helicello strioto
o
r
o
0
g
o
0
dsl
O.
4°
•
8 ezi
Conservotion desirable wherever possible
Fig,11f
á
- 25 near
Silurianslatesto the north(Fig.9).In thewoodlands
it was howeverat one timemoreabundantand a nuMberof
Windermere
celdnieshavebecomeextinctduringthe present
formerly.flourishing
stillexistbut the canOpyhas been
century:The woodsthemSelves
of'thelargertiMberduringthe two aorldWars.
extraction
broken..fdr
littlehas been dehe
With-thedeclinein demandfor nativehardwoods,
to restockthesewoodsand as many are grazedthereis little.0nthe limestonethishas leftlargewell-isolated
regeneration.
area,in thewoodson acid soils,the
but in theWindermere
clearings
havebeen invadedby densegrowthof bracken. The extincclearings
and
(Satchell
tionof the colonieson thesesiteshas been attributed
of
1955)to the spreadof brackenthroughthe reduction
Collingwood
for the
of the groundlayerwhichis essential
directinsolation
adtivity.
ants•forazing
in Fig.10,
illustrated
In the areaof South-East'England,
validrecordsof rufa had been madebefore1960 from44 sites.
. Of the olderreby Barrett(1965)
In 1960-64thesewere examined
cordedsitessix,fivein Londonand one in Hove,had beenbuilt
over. F. rufawas net foundin a furthoxsevensiteSon the dut.,
Reigate,Loatherhead
Gillingham,
Wells,Maidstone,
Skirtsof Tunbridge
and Woodfordand thesecoloniestoo are probablyextinctfromurban
presenceof the
In 31 ten km. squares,the continued
development..
squares,ii was recorded
ten additional
ant was confirmed'and.in
In thispart
flourishing
for the firsttiMe. F. rufa is apparently
of Britain,otherthanon siteswhichhavebeen totallydestroyed',
withstandthe intensehubanpressurefor whbh the area
and can.evidently
is notorious.Its abilityto survivesuChpressurenear the centre
by the record(Yarrow1934)of F. rufa.
of its rangeis supported
contrasts
Gardensas late as 1831. -TheSituation
fromKensington
minor
with thatin theLakeDistrictwhererelatively
markedly
of the ants'habitathas resulted'in'its
over
disturbance
eictinctien
area.
a large.
of Hamrstead
(b) Survivalof the AculeateH non-tern
of the effeCtof urban
thebest documentedanalysis
Heath. 'Perhaps
is thatof the aculdate
of invertebrates
pressureon an association
Heathby GuichardandYerroW(1948)and
of Hampstead
Hymenoptera
whichhadbeen
Yarrow(1954). Theseauthorslisted49:species
informcenturybut not since. Sufficient
recordedin the nineteenth
on 27 of themto assesstheirprobablesignificance
ationis available
wereprobablyvagrants
12 Were eitherdoubtfulrecords,
-andof-these,
Of the other15, 3 appear
or havebeon rediscovered.
to havegone
of its nesting
with theirhostplants,one with the disapaearance
hosts. If the
With theirinsect'
'andpredators
siteand 11 parasites
betweenprobably
apportioned
POr are similarly
22 speciesunaccounted
doubtfuland prdbablyreliablerecords,the nuMberof specieswhich
in-thisperiodseemsto be about30. Added
has reallydisappeared
would
of the atesentcentury,.this
to the'150authenicated.reCords
suggesta.lossof not mOre thanaboutone sixthof the aculeate
of theHeath: The Heathof a centuryNib was surrounded
Hymendpiera
to
by suburbs,nowhereconnected
by fields. It is now sUrrounded
this-insect
of
sixths
five
opencountry.The survivalof
Sala:ate.'
- 26 -
underthe intensity
of urbrnpressurecharacterising
Hanpstead
Heath,
indicatesa degreeof resilience
which,thoughnevordoubtedby
agricultural
entomologists,
is sometimes
under-estimated.b
cony
servationists.
It suggeststhatin the conservattin
of invertebrates
in naturereserves,
protection
againsthumaninterference
may be
of less consequence
thanthepripersitingof the reservein relation
to the climaticdemandsof the speciesit is desiredto protect.
III CONCLUSIONS
In,the.present
stateof invertebrate
conservntion
research,
the kind of case-histories
citedformone of the few'available
sources
of guidancein.theformulation
of an invertebrate
conserVation
policy.
They 'suggest
the following
provisional
conclusionS:(1)
The invertebrate
faunaof Britainis not a staticentity
but containselementsfromdiversesourceswhichare subjectto
changesin densitYand distribution.
(2):
.0necauseOf suchchangesis variation
in climateand
weather:.
SpeciesWhch ate on the edgeof .their
distribution
in
Britainare pnrticularly
susce;-)tible
to suchvarition. Theremay.
be scientific
groundsfor conserving
themif theyform:genetically
distinct'populations
CT non-scientific
groundSin the caseof:rarities'
valuedby naturalists.
But effortsto.conserve
themcannotin the
longrun be expectedto suceeedunlesstheirhabitatiS sufficiently
largeand variedto providerefUgesfromunusual-weather
and climatic
trends. Naturerescrvesfor such specieswiuldgenerally
need to belarge,of the orderof several
.squaremiles,to provideboth the
refUgesand the'Continuity
of habitatnecessaryto permit.recdispersel
from them.Anothercauseof suchchangesi habitatdisturbance
by
(3)man. Sone of the'Species
whoSeexistence
in Britainis threatened
.by'habitat
destruction
are also speciesat the northernend of their
range. It followsthat*
preservation
of samplesof theirhabitats
as nnturereserveseannot.be
expectedto securetheirprotection
unless-the
reserVesera very lnrgeas indicated
above..A numberof
fenland,heathland
and chalkrgrassland'species
fallinto thiscategory.
(4)
Ls a converseof (3), populatiDns
of speciesnear the
centreof theirrangegenerally
appearremarkably
resistant
to human
pressureshortof totaldestruction
of the habitat.
(5)
The logicalplaceto eonserve'any-spedis
ieswhereconservationmeasuresare mostlikelyto succeedi.e.towardsthecentre
of the species'
range. Thisinpliesan international
approach'aiMed
'atconserving
sreciesWith,for example,a mainlyContinental
European
'distribution
in Continental
Europeand Mediterranean-species
in.
•Mediterranean
countries.
- 27 policyono of the
In the contextof suchan international
of
of Britainwouldbe the conservation
spedialresponsibilities
thatthese
advantage
species.It wouldbe'a practical
Arctic-Boreal
occurin the northernand highlandareasleastsUbjectto land
value
scientific
pressure.Nbre couldbe achievedof international
thesespeciesthanby 'attempting
on conserving
by concentrating
to preservesouthernspeciesagainstthe oddsof climateand land
demand.
(6)
for
on the invertebrates
information,
Thereis a soUrce-of
(7)•
and
in the taxonomists
measuresare desirable
whichconservation
listsof
groups. Provisibrol
invertebrate
in particular
specialists
st, specieswith special
scientific'interee.g.
speciesof particuler
with special
with relictdistributions;
significance;
evolutionary
- could.bedrawn
withpeculiarlifehistories
habitatrequirements;
on an international
by &Mallmeetingsof s2ebizilists
up, groupby grout),
basis.
MERENCES
K. J. 1965 (at press).
BARRETT,
Nolluscaof the
A resurveyof the.freshwater
.L.-E..1928:
BOYCOTT,'
parishof Aldenhamafterten yearswith.specialreference
..Trans.Herts.Nat.Hist.Soc.,12,1.
to the effectof drought.
and habitatsof Ena montanein
BOYCOTT,A. E. 1939.Distribution
England. J.Conch.,Lond.121,153:
•COTTTNGAOD,C. A. 1958. A surveyof IrishFormicidae.Proc.Roy.
IrishAcad.,51,213.
CRISP,D. J. ET AL. 1964. The effectsof the severewinterof 1962
63 on marinelife in Britain. J.Lnim.;Ecol.,33,165.
on the effectsof climateand
CRISP,D. J. 1965. Observations
'The
weatheron marinecommunities.Inst.Biol.Symp.
of ClimaticChangesin Britaint.'
Significance
Biological
FORD,E. B. 1946. Butterflies.Collins,London.
der CastenDetermination
K. and BIER,K. 1954. Untersuchen
GOSWALD,
in der GattungFormica. InsectesSociaux,1, 305.
Aculeata
K. M. andYARROW,I. H. H. 1948. The Hymenoptera
GUICHARD,
-1947.
832
1
district,
Heath and the surrounding
of Hampstead
LondonNaturalistII,81.
T. J. 1964. The Late-glacial
KERNEY,NL P., BROWN,E. H., & CHANDLER,
nearBrook,
historyof the chalkescarpment
and Post-glacial
Kent. Phil.Transaloy.Soc.Lond.,248,135.
- 28 -
LAMB,H. H., andJOHNSON,
A.I.1959. Climatic
variation-and
observed
changesin thegeneralcirculation.
-Geogr,Lnn.,41,94.
LAMB,H. H. 1963. Whatcanwe findout aboutthe trendof our
climate?Weather,
18,194.
LAMB,H. H. 1955. Britain's
Changing
Climate.Inst.Biol.Symp.
'TheBiological
Significance
of Climatic
Changesin Britain'.
G. 1965. Possible
climatic
agencies
in thedevelopment
of
post-glacial,habitats.
Froc.Roy.Soc.Lond.,161,363.
SATCHELL,
J. E., & COLLINGJOOD,
C. A. 1955. TheWoodAntsof the
EnglishLakeDistrict.NorthWesternNaturalist,
2, 23.
SOUTHWARD,
A. J. 1958. Noteon the temperature
tolerancesof
some
intertidal
animals
in relation
to environmental
temperaturesandgeographical
distribution.
J.Ear.Biol.Ass.U.K.
2,49.
717.s0N,
G. FOX 1939. Insectpestsof rhododendrons:
their_distributionin Britain Proc.R.Ent.Soc.,Lond.,Ser.A,14,1.
WRIGHT,
A. E. 1942. Plebejus
argusLinnaeus
racemasseyi
(Tutt)
in. NorthLancashire
and.SouthWestmerland.
Entomologist,75,7.
YARRaT,I. H. H. 1954..Recordsof Aculeate
Hymenoptera
at Hampstead.
EntoMoilogiSt
87,168..
- 29 -
DIVEZTERRATE
CONSERVATION
AND
MANLG•ti
PUNS
E. Duffey
,Atthepresenttime,69 management
planshavebeen
completed
by theConservancy
and severalothersare in the courseof
preparation.A recentanalysisof the 550 research
projectslisted
in theseplans as "inprogress"or "proposed".
Showthat-47are in
the formergroupand are concerned
with invertebrate
investigations:
Of thissmalltotal,56% are by scientists
not in theConservancy,
31% by Conservancy
scientific
staffand 12% by researchstation
staff.'Ohly'a-Sg
iali- Proportion
WereplannedsPecifically
in relation
to management
studies.A furtheranalysisof the47 projectsin
relationto (a) habitatsand (b) invertebrate
groups,is as follows:(a)
Lowland
Coastal Uplands Mountain GrassHeath WetlandSWoodland
4%
(b)
2,Z
25%
21,4
.31
. Insects Freshwater
anivnls Non-inSect
groups Unspecified
20%
The onlyconclusion
whichone can draw fromthisis that
theproduction
of ecological
knowledgeon inVertebrates'on
Nature
Reservesis poor and apartfrom soreof the work at MonksWood and
FUrzebrook
iS not orientated
towardsManageMent
studies.
The presentation
of Invertebrate
materialin the'earlier
management
plansis verypoor,bUt improvedwith the introduction
of the use of the-Chart
of'Animal
Habitats.The Chartis slowly
beingacceptedand is provingof valueas an assessment
of the
potential
richnessof the Reservefaunaand-as a-basisfor inters,
pretingexisting
listsof gpeciesrecorded.This has been done in
severalcasesby-the-Conservation
ResearchSectionusinEold:records
forwhichecological
datawas not recorded'at
the time. We-feelthat
a habitat-analysis
is a valuablepreliminarY
for any zoological
projecton a NatureReservewhetherthis is a general'survey
or a
plannedexperimental
studyof a management
problem. The designation
of habitatcategories
can be supplemented
withbotanicalinformation
and in somecases,thismay be very detailed.Where the information
on plantsand animalsis cemprehensive,'it
is possibleto makea
usefUlcross-reference
betweenthe Sections:In the faunaand on
the vegetation.The management-of
a NatureReserve'for
invertebrates
is essentially
the maintenance
of'hahitats'although
not all.areef
equalimportance.For thisreason,it is important
to findout as
much as possibleaboutthe faunain each habitatcategory.
30
Comprehensive
faunalstudiesof ecologically
complexNature
Reservesare a formidable
problemand requirethe servicesof many
specialists
including
staffto maintain
recordsand collections.Of
the 21,000 ( approximately
) speciesof Bidtishinsects,onlyabout
5,000 are "accessible"
in the sensethat therearo modernor older
reliablekeys available.Llthougha few NationalNatureReserves
are comparatively
wellknown,nonehas the detailedknowledge
available forWythamWoods,Oxford. As an oxampleof the valueof relating
habitatrichnessto animalcommunities
the following
may be taken.
Woodwalton
Fen and WickenFen are both well knownentomologically
and includea habitatrangefromwoodland,scrub,reedbed,herbaceous
fen to varioustypesof waterbody. ScoltHead is an extensive
dune
systeMwith.some shingleformation.L habitatchartwas completed
for eachReserve,
and the species'
totalsfor severalimportant
orders
of invertebrates
are listedbelow.
Woodwalton
Fen flckenFen -ScoltHead Island
m Totalsof speciesrecordedin Britain
The figuresfor ScoltHead are probablythe most accurate
because
it is relatively
simpleecologically
and has changedlittlesince
biologistsfirstworkedthere. The two fensare much mere.complex
ecosystems,
moredifficultto wOrk thoroughly
and have charged
considerably
sincerecordsof animalswere firstmade.
Invertebrate
studiesdesignedin relationto management
problems
should.be
carefUllyselectedto avoidthe dangerof findingthatone
has to sort throughan immenseambunt
of materialwhichcannotbe
identified
withoutspecialist
helpand a greatdeal of timeusing
keys. Two usefUlmethodswhichproducevaluableresultsare:(a) the selection
of a few,or perhapsonly onc, speciesof special
interestwhichcan be studiedintensively
in relationto habitatand
the environmental
faCtorsinfluencing
nuMbersand behaviour.
(b) the selection
for spocial.study
of the animalcommunity
of a
habitatwhichis of importance
in management
workbecauseit is
extensive,
vulnerable
in someway to outsideinfluences
or has been
artificially
maintained
for a longperiodby someformof exploitation.
- 31 -
by
Singlespeciesstudieswhichcan be illustrated
value
special
of
are
Copper
ge
on
the:Lar
work...
the
to
ce
referen
of certain
significance
the ecological
becausetheydemonstrate
in the habitatin whichthe specieslives. This opens
components
the co-operation
Studiesinvolving
theway to othertypesof management
intensive
deserve
which
species
of
s
Example
ines.
discipl
other
of
whichfeeds
butterfly
studiesof this typeare theBlackBair-streak
to tell
is
Hunt
Mr.
which
ly
butterf
Blue
Large
the
orn;
blackth
on
numerous
and
studied
has
Moore
Dr.
which
lies
dragonf
certain
us about;
specieswhichcanbe selectedfromlesswell knownOrders.
valuablefor
is particularly
The LargeCopperbutterfly
Fen and.hasa single
studybecauseit occursonly on Woodwalton
of.thisinsect
ion
populat
The
Dock.
Water
Great
the
nt,
foodpla
has yaried
1953
since
)
emerged
adults
of
number
by
d
(as measure
100-120.
about
is
it
years
most
in
h
althoug
250,
and
60
between
edgesof
the
along
is
nt
foodpla
the
of
ution
distrib
natural
The
cted
constru
ially
artifie
are
Fen,
ton
Woodwal
at
which
yS
waterwa
dykes. In suchplaces,the dockgrowsintoa large,vigorousplant.
The insectdoes not like suchplantsfor egg-laying;the-optimum
as the
for thispart of the lifecycleare not the-same
conditions
are
eggs
for the growthof the foodplant.The
optimumconditions
usuallylaid on smallerplantsawayfromwaterand in surrounding
whichdoesnot smotherthe waterdock. The presenceof
vegetation
on its Own is hot enoughto maintainthe insect; it
the foodplant
of this
habitat.The identification
mustbe growingin a particular
re
therefo
is
it
or recreating
habitatand the methodsof preserving
leads
This
of the insect.
the firststepin ensuringthe survival
and theircontrol,germination
to a studyof watertablefluctuations
the.
by.thedock seedand the factorsaffecting
required
conditions
growthof the youngplants.
Anotherasoectof the studyof thisspeciesconcerns
and after
dynamics.The eggshatchin late.summer,
population
feedingfor a littlewhile,the firstand secondinztarlarvaemove
until
to the deadleavesaroundthebase of thelant and hibernate
after
and
high
is
ty
spring. DuringthisperiOdmortali
the following
heavily
be
may
larvae
in the summerthe surviving
feedingrecommences
by a tachinidflyPhryxevulgaris.The conservation
parasitised
problemarisingfrom thisis to decidehow largethe populntion
mustbe to avoidextinction.The nuMberof eggslaid eachyear is
veryvariable:1960
1961
1962
1963
NuMberof eggs laid 1,500 1,700 3,800 10,900
femalescan lay an averageoflabout350 eggs although
In captivity,
as manyas 600 havebeen recorded.Since1960,the nuMberof eggs
lnidper femaleon the Fen has averaged72 (20.55{.of averagetotal
in captivity).
- 32 -
The figuresare as follows:1960 1961 1962 1963
AVeragenuMberof eggsper
femaleon theFen
24.
69
126
The percentage
survivalof larvae-after
wintermortality
is generally
very low:1960/61
Wild colony 6.4%
Cagedcolony 4.3.
1961/62
1962/63
10.0A 3.5%
17.1%
1963/64
1901/65
1.7%
3.1%
3.1%
The traditional
methodof maintaining
Ahis butterflysince
its introduction
in 1928 is to collectthe younglarvaeemergingin
the springand.Place
themon foodplants
in muslincageswherethey
are protected
frompredatorsand.parasites.
Since1961 however,a
population
of LargeCoppershas been maintained
in anotherpart
of theFen whereno.protection
is provided.The comparative
figures
for survival
are as follows:% survivalof larvaeaftersPringemergence
to pupation
We nOw have,
someevidenceto Shdwthat,with the population
at the levelit has been maintained for manyyears,
thereis a strong
risk of extinction if the larvaeare not protected
duringthe stage.
up to pupation;
but we are alSobeginningto demenstrate
that suitable
habitatscan be re-created
in the Fen for the successful
regeneration
of waterdockswhichdevelopintoplantsattradtive
to the egglaying.female.
Studieson the faunaof-Selected
habitatscan nlsobe
illustrated
by reference
to ',7oodwalton
Fen. The Fen dykeswere dug
about100 yearsafro for the transportation
of peat. Thesewaterways:haveto be cleanedout from timeto timeor else they soon
becoMechokedwith vegetation.They forma typeof waterbody not
usuallystudiedby freshwater
biologists
andbecauseof their
uniformwidth,depthand methodof maintenance are very suitable'for
experimental
work. Dr. Morrishas been studyingthe development
of thefaunain.anewlyconstructed
dyko and the influence'on
the'
aquaticbugs andbeetlesof.the
annual
cuttingof submerged
and
marginalvegetation
in olderdykes. The presenceand quantityof dead
plantmaterialin the waterhas a significant
effecton the composition
and nuMbersof aquaticanimals.
- 33 -
Research
The mainhabitatstudiesof theConservation
of a type
fauna
the
on
been
Sectionduringthe last two yearshave
"Openas
Chart
Habitat
on theAnimal.
of Breckheathclassified
6"
than
less
ly
beinggeneral
the vegetation
groundformation",
this
in
d
inclUde
be
can
ion
in height. Many othertypesof vegetat
some
and
s
pasture
upland
nes,
habitatcategory;chalkdown,sand-du
a
of
studies
cal
ecologi
for
heathlands.Theyhave the advantage
and a faunawith a limited
simpleand oftenuniformstructure
numberof species.
The main objectof Brecklandstudyis to assessthe
almostcomplete
faunain relationto the.:
changesin the invertebrate
(a) a
chosen:
were
areas
of rabbitgrazing. Two study
cessation
myxosince
Heath
Weeting
at
ed
thickuniformturfwhichhas develop
small
moSses,
,
lichens
mostly
matosis;(b) very Shortvegetation,
annuals,a littlegrassand with SOM bare groundand stones.Very
in thesetwo sitesandthe
distinctfaunascan be demonstrated
growthin recentyears-pp-arsto haveresulted
surgeof vegetation
in the loss of a nuMberof speciesof animalswhichare more
The material
onditions.
s
Breck-c
of the pre-myxomatosi
characteristic
d.
examine
being
fromthe many samplestakenis still
The secondpart of thiswork is to try to identifythdse
for
of theBreckhabitatwhichhave special'significance
components
the
of
parts
Those
.
species
ting
the survivalof the More interes
rabbit
are isolatedgrasstussocks,
habitatWhichhavebeen.studied
.-ce'of
'presen
stones,
nce.of
er-abse
e
r
presenc
holesand scrapes,
ge
knowled
The
Plants.
heather
and
d'gorse
mdbilesand,dung,isOlate
d
obtaine
at
with.th
d
coMpare
be
then
will
derivedfromthismosaic
work on.the two studyareas. A few
from the more.comprehensive
Arctosa
thework on Breekland.sdiders,
from
examplescan be quoted
on vary
but
nes
sand-du
cOastal
on
found
y
is.a lycosidcommonl
perita:
there.
is
llhere'
occurs
it
heaths,
Brock
few inlandsitcs. On thc.
such
as
soon
As
ion.
vegetat
short
ed
exposedsandwith scatter
it disappears.Two salticids,
vegetated,
areasbecomecompletely
occurin similarhabitat.s
s'also
aequipe
s
r Euophry
Attulussaltatoand
areasmay be
coastal
.
of
typical
althdugAthe latterwhichis not
is
ion
vegetat
the
as
long
as
foundwherethereis no bare ground
ted
ivegeta
sparsel
s
6peri
require
ulatus
short. Lith hantesalbomac
plant8
taller
ated
Or.isol
s
tussock
stones,
are
there
habitats.where
0 tila seabricula
aroundwhichthe webcan be spun. The crab-spider
countryandthis
duneain
or
heaths
dry
is rarelytakenaway from
.•
scrapes
or
holes
rabbit
with
is oftenassociated
SUIDJARY
Thereis stillverylittleknownaboutinvertebrate
(i)
faunason NatureReserves.Yuchprimarysurveyinrelationto a
is required The use of the Chartof Animal
habitatclassification
studieson Nature
to invertebrate
nary
prelimi
e
Habitatsis a valuabl
al.
essenti
be
may
areas,
complex
cally
Reservesand in ecologi
speciesand'theshortage
The largenumberof invertebrate
(ii)
projectsmuStbe
management.research
means.that
of specialists
studieson a
detailed
arc
methods
usefUl
'Two,
selected.
carefUlly
to ecological
with specialreference
few,or a singlespecies,
on
in relationto the Reserveas a wholeand.studies
requirements
becauseit is of specialsignificance
the faunaof a habitatselected
purposes.
for conserVation
of thistypefor
knowledge
of ecological
.Theaccumulation
(iii)
Formation,
a rangeof habitatswithinfor example,the Onen-ground
suchas grazing.
treatments
the effectsof management
will illustrate
It will alsorevealthosespecieswhichare of valueas indicators
and the pattern
in the environment,
conditions
ecological
of particular
of habitats
in relationto the distribution
of animalcommunities
Shouldemerge.
Summar of Discussion
from the
A memberaskedwhetherthe LargeCoppersuffered
country.
this
in
does
it
as
Holland
in
samedegreeof parasitism
fly seemedto be
Dr. Duffeyrepliedthattwo speciesof ichneumon
but the parasiteof the Large.Copper
in Holland,
the mainparasites
Fen had nearlyalwaysbeen the tachinidfly,Phryxe
on Woodwalton
was extremelyDempstersaidthatPhryxevulgaris
.Dr.
vulgaris.
by his work at MonksWood
prOlificand thishad been demonstrated
by thisfly on the SmellCabbage
wherethe degreeof narasitism
to the
is of the orderof 15-2074. Referring
bUtterfly
White,
in Holland,Dr. Duffeysaid thatthe LargeCopperhabitat
situation
for
exploited
in theDutchfensis in an areawhichis extensively
on whichthe eggsare laidoccur
hay and reed. The dock.plants
and
distances,
by considerable
separated
in smallgroups;sometimes
that-the'
to theDutchchtegolOgists
was expressed
when surprise
theyreplied
insectcouldmaintainitselfundersuch.conditions,
of the..
smallgroupsseemedto be characteristic
that scattered
may redUde7the
the LargeCopper. Such dispersal
distribution.of
.of parasitism.Dr. Satchellpaidthatif thu LargeCopper
incidence
parasitein Holland,thenIt
sametachinid
is not subjectitd-the
attemptto conserveit in this
couldbe arguedthatwe,should.not
whichmay be particularly
environment
countrywhereit is in a different
Dr. Duffeysaid that theLargeCopperhad existed
unfavourable.
on othersitesih thiscountryand gave as an
withoutprotection
exrmpleWickenFen, whereit thrivedfor Peveralyearswithout
Fen was drained.
until.Adventurers
protection
- 35 -
STLTUS AND CONSERVATION OF THE LLRGE BLUE B
MACULINEA ARION L.
O. D. Hunt
INTRODUCTION
The status in Britain of the Large Blue butterfly has
declined to a point where it must be asked:1.
Will it.surviva without conservation?.
2.
Can-it be conserved?
in mind the Devon Trust Tor Nature-Conservation
With this
decided in 1962 that a survey of the status of the species in Devon
and Cornwall ought to bu made, especipaly as its breeding grounds
in these two counties were now-generally thought to be the-only ones.
of consequence remaining.
The Nature Conservancy was approached-and steps taken to
arrange for the organisation of any survey that might eventuate.
Proposals far a too-year fact-finding survey wore put forward to
the Conservancy,by a Joint:Committee on which the NaturaliSts'
Trusts.for Devon and Cornwall, the Royal Entomological Society, the
SocietY for the'Promotion-of Nature Reserves and the NatUre ConserVancy were all represented. A grant was obtained and the survey
has now been under Way during the years 1963 and 1964; while further
work, now in-train, will be continued-during 1965 if support from
the ConserVancy,is fortheoming.
This paper is presented as a brief narrative of what has
been accomplished, summarising the position in the light of the .d.ata
so far available.•
LIFE HISTORY
The briefest remihde::.of the life-history of the Large
Blue will serve to Show that-its survival depends entirely on the
maintenance of a special habitat.
For part of its larval existehce, the Large Blue is
Myrmica sabuleti and M.
parasitic on ants of the genus It/Tr:Ilea,.
and Cornwall—The
species'inDevon
host
main
the
being
scabrinodis
flowers of wild thymo-,-Thymus drucei, on which
eggs are lald'on the:the caterpillar at first feeds exclusively. -Later it is adopted by
the ants and taken underground into their nests for the sake of
secretions from its honey gland which the ants are able to excfle.
It stays in the ants' nest from autumn to spring, feeding on the
ant larvae, and pupates there, the imago emerging from underground
in early Summer.
- 36 -
THE SURVEY
BASIS LND PROCEDURE
At the start.of the enquiry, very little up-it6-date inform-,
ation was available abdut:(a:
(b)
in Devonand
the extentof the LargeBlue population
Cornwall;
the confinesof itsbreedinggrounds.
fileon the
l
However,theRoyalEntomologicaSociety's
in whichthe most valuable
LargeBlue was placedat our disposal,
itemswere an accountof a surveyof the coastalarea of a part of
NorthDevon and NorthCornwallcarriedout in 1948 by CaptainR. A.
intothe1950's relating
extending
Jacksdnand somecorrespondence
to protectthe LargeBlue on
to +he effortsof a Devonlandowner
his property.
we were indebtedto Nr..Alan
For laterinformation,
in the fieldduringthe seasons
Kennardwho had madeobservations
1961 and 1962.
researchinto the
was an intensive
Of basicimportance
relatingto theLargeBlueby Mr. G. M. Spooner,Secretary
literature
enquiries
who not onlycarriedout exhaustive
of the JointCommittee,
with authorbut corresponded
references,
in tracingand evaluating
and collateddata frommanypartsof
ities,examinedcollections
to the reasonsfor
himselfspecifically
the country.Addressing
the declineof theLargeBlue in Britain,Nr. Spoonerhas edbodied
in The
a valuablepaperon this subjectpublished
his findings'in
r
for•Septedbe1963.
Entomologist
of the surveywas to locateand
The firstrequirement
definethe presentbreedinggroundsof the LargeBlue and,if
remainingtherein.From earlyin
possible,estimatethe population
were directedto an obsen,
operations
the 1963 seasontherefore,
was knownto
vationalsurveyof regionswithinwhichthe butterfly
data,of all
with appr3priate
have occurredand to the listing,
for the
appearedto be favourable
siteswhere the conditions
speciesin respectto:1.
2.
3.
wild thyme;
the presenceof the foodplant,
species;
the presenceof Myrmica,of determined
the presenceof Shelterin the form of groundconfiguration
vegetation.
and suitable
- 37All available
helpwas thendeployedto watchthese
sitesfor the appearance
of thebutterfly
and coapilea recordof
the nuMbersseen,,
whileone sitewas selectedfor an attempted
censusby markingall,specimens
capturedthroughout
the.scason.
RESULTS
The summerof 1963was Wet,windyand coldand the nuMbers
of LargeBlue seenwereprobablyinfluenced
by this as.wellas by
the fact that
.the previousseason,1962,had alsobeen a verypoor
year in whichvery fewlaarge
Blueshadbeen reported.Out of the
34. locations
wherethe surveyhad judgedconditions
to be suitable
for the LargeBlue,"only
13 producedrecordsin 1963and the total
numberrecordedthroughout
the flyingperiodwas only85, two of
the sitesaccounting
.for more thanhalf of this total. The best,
siteyieldedonly 27 so thateven therethe speciesappearedto be'
at a low ebb,whileat most of the'sites
the impression
was of neat
ness.to vanishing
point.
Colourmarkingwith quick-drying
cellulose
paintsapplied
as smallspotson the wingswas carriedoutin four sites,especially
in one whereeasy accessenabledfrequentobservation
and whereit
was hopedto estiMatethe population
by the capture-recapture
technique.Over a periodof ten days,thissite yielded.23
captures
and,recaptures,
representing
a totalof 12 separate
individuals,
figures,which
scarcely
providemuchfor analysisbut serveto confirm
a verylow.status.
Oneinteresting
outcomaof thisexperiment
was to show .
thatwhile theLargeBlue has,in general,a sedentary
habit,tending
to remainthroughout
its lifetimewithin smallarca,yet movemcnts
of individuals,do
takeplaceovera sufficient
distanceto suggeat:
that,givena good seasonwith enoughbutterflies
involved,
travel,•
especially
if wind-assisted,
coulddistribute
the speciesfromone
breedingcentreinto othersuitable
sitesif suchexist.within
range. Whatthat rangemay be cannotbe stated.with
any certainty:.
tho longesttravelebseryed.in
tbis,experiment
wasuabbuthalfa '
milebut thismas coveredso quicklymithwind helpingthatconsidlniply
greaterdistancesmightbe.expected.
The breedingsitessurveyed
are in a series.of
cooMbes
and cliffslopesalongthe,ceast,
separated
onlyby stretches
where
the requisite
conditions
for foodand shelterare lacking. /larking
experiments
of the kind hereattempted,
evenif inSufficient
for
censuspurposes,
can at leasthelpin understanding
the relation
betweenthesebreedingsitesas linksin a relatedchain,and the
relevanceto survivalof everylink:-
- 38 -
The surveyin 1964 followedmuch the sameplanbut with
n moreto censuswork
and
y
less timegivento preliminarexploratio
thesite of
On a selectedsite. Unfortunately,
and concentration
as
whichhad been earmarked
the previousyear'smarkingexperiment
workwas ruinedby agricultural
suitablefor'somsfurtherintensive
marking
but a successful
thewinterof 1963/64,
n
reclamatioduring
most
the
been
had
was carriedout at the sitewhich
experiment
populousin 1963and whichprovedso againin 1964. Here,though
the dataare not high enoughto requirecomnuter
the figures.in
s be drawn.
can
t
sorm significanconclusion
analysis,
at thissite
s
The totalof separateindividualmarked
and somemore
once
at least
was 119. Of these,44 were recovered
five times. Analysisof the figures
thanonce,onebeingrecaptured
thewholeof the flying
throughout
a totalpopulation
indicates
not exceeda maximumdailypopulation
seasonof not more thanar,
ing 70 and an average"life"of the butterflyof 3 to 4 days. The
longest"life"recordedwas 9 days and manymusthavebeen very
rate can be equatedwith the
d the disappearance
short-liveif
deathrate. Thishowever,may notbe the case.
in the 1964 season,with
The weatherwas favourable
but althoughthe totalnuMberof LargeBlues,
abundantsunshine,
recordedon all siteswas deublethat for 1963,the nuMberof proat
ductivesiteshad droppedfrom13 to 10 and the concentration
80%
over
yielding
was far.greater,
the two mostpopuloussite's
of the habitathas a fore.
of the total. This apparentshrinkage
this,
but offsetting
bodingaspectfor the futureof the species,
from
resulting
recovery,
of
thereare groundsto hope for a measure
balanced,
thus
situation
the
With
of 1964.
the goodweatherconditions
duringthe
shallbe continued
it is mostdesirablethatobservations
surveyed.
been
far
so
1965 seasonon all the sitesthathave
Duringboth yearsof thissurvey,not a singlerecordof
the LargeBlue has been obtainedfrom thatpart of the surveyed
of the speciesin
areawhichwas the classichauntand stronghold
or more.
thousand
a
the earlyyearsof thiscentury,yielding
partwhich
that
in
ner seasonto collectors.Moreover,
specimens
to a
points
status
low
its
grounds,
providesits presentbreeding
from
deduced
as
ago,
decade
a
Obscrved
markeddeclinefrom thelt
of individual.s
recordsand from notesand.recollections
documented
in the fieldduringthe mid-1950s.
CAUSESOF DECLINE
it wouldseemthatthe answerto the
From the preceding,
is quite
withoutconservation?"
survive
species
question"Will.the.
on
conservati
"Should
as
be.posed
question
probablyno, whilc if the
yes.
ly
emphatical
be
must
answer
the
ed?",
measuresbe contemplat
- 39 -
Althoughthereis evidenceof the abilityof the-spedies
tho marked
'
areas,
to survivefor yearsin smallnumbersin 'small
a history
with
s
specie
a
of
trendof declineshYwnby this•survey,
abundance,
former
of
region
after
e region
of totaldisappearancfrom
7 ."What
2.
Ne..
n-of
ion
quest
eratio
to,a consid
bringsus undoubtedly
of conservation?"
are the requirements
the causeof decline,the evidenceaccruing
In considering
surveygivesgeneralsupportto thepicturepresented
fromthe-present
by Spoonerin the studyalreadyreferredto. Brieflythe factors
are:responsible
1.
2.
3.
4.
5.
Heavycollecting;
of breedinggroundsby agriculture;
Destruction
on breedinggrounds;
Scrubencroachment
AdverSeweatherconditionsi
Fire.
,
practices,
ltural
suchas collectingagricu
Humanagencies
necessarily
means!
no
by
are
,
theory
in
and fire,thoughcontrollable
is.
chment
encroa
scrub'
es,
agenci
l
so in actuality:Of the natura
con
ly
direct
ot
n
er,
weath
the
l:
ameroble to a measureof contro
be circuamented.
may not impossibly
trollable,
- Spoonerhas adducedthat theLargeBlue has
Collecting
wheretherehas been
disappeared
ting.and
collec
heay
both survived
inglyyulnerable•
exceed
is
s
apecie
,
the
heless
no collecting.Nevert
other
ra and when
extrem
been
-as
whiehl
of
nce
to thisfactor,the incide
that-.
le
probab
highly
seeMs
ng
it
as•well,
adversefactorsareoperati
for the coupde grace. Of
may easilybo.responsible
collectors
aerioUsthre-Ato surviva.
most
the
not
s,evenif
e
all-singlfactor
be a •
wouldcertairly
which
of
ion
ppress
today,it is one, the.su
tion.
potentmeasureof.conservri
ground
of the reughYuntilled
- ReclaMation
Agriculture
tand2.
habita
its
d
redube
has
ng
breedi
for
needs
that theLargeBlue
ially
potent
tor
collec
the
ng
escapi
of
s
therebylessenedits chance
for
Except
e.
declin
of
ration
accele
rapid
for
s
a proces
therefore,
of. f;,-.
thereis no-part
onwallsand hedgebanks,
verylimitedareas
grow.
can
,
thymu
food
wild
,
the
plant
where
ground
cultivated
may be
and herbicides
ed of insecticides
-The.increasuse
.
ce
eviden
te
defini
no
but
Blue
Large
the
Of
e
declin
involvedin the
of the survey.
the.course
of thishas-sofar beendiscovered'in
likely-to
winds
high
and
kwayed
ly
common
re
the
area•a
Cropsin
that
ible
imposs
is
riot
It
ence.
Sccurr
normel
of
causedriftare
means.
thia
from
ed
suffer
has
plant
the food
-40 of rabbitsaffordsanotherinstanceof a clash
Suppression
and thoseof theLargeBlue,
of agriculture
betweenthe interests
helpedby rabbitgrazing.
for the survivalof wild thymeis,greatly
- Wild thymedemandsthe absenceof
-Scrubencroachment
on factorsthat suppress.
and dependstherefore
vegetation
smothering
orsare many,
tsllergrowthand preservea Shortsward. Suchfact
ts;
agen shallowness,
being erosionby wind and wind-borne
examples
walls
and Sharpdrainageof soiloverrocksand ridges,on
elevation
farm
and
als
anim
ant-mounds;grazingby wild
and mounds,especially
of
th
grow
rich
stock;-burningoff in earlyspring(swaling).The
enis doubtless
on ant-mounds
characteristic
thymeso frequently
with
ed
ciat
with soilasso
by theperiodictopping-up
couraged
for thyme
The idealconditions
ds.
moun
d
bite
activelYgrowinginha
ciesacting
agen
of
ion
froma combinat
growthwillbe thoseresulting
y le,
stab
ivel
arat
s
balance,sometimecomp
to producea favourable
example
one
e
phas
a passing
as representing
oftenmore transitory
een
betw
od
of whiehis probablyto be seenin the interimperi
by
swalingsat suitableintervalsefyears,accompanied
successive
ing.
e
the effectsof selectivgraz
without
The effectsof grazingwill notbe understood
scope.
le
arry
and
for whichthereis obviousneed
intensive'study,
tor,
'fac
thiS
of
ct
one aspe
Thereis hOwever,no.doubtconcerning
ual
virt
,
time
a
for
in nuMbersand,
namelythat the greatreduction
proa
had
has
sie,
mato
myxo
e the rabbitfollowing
disappearancof
e.
thym
wild
on
ct
effe
rse
foundlyadve
of
Weather- Spoonerhas adducedevidencethatdecline
by
nt
exte
some
to
theLargeBlue in spmoof its formerhauntsmayincreases
ed
ciat
asso
ing
,
with climaticfluctuationstrac
correlated
s,
year
"
"bad
s
with
with goodweatherseasonsas well as decrease inemay wellbe
.
theseoccurinsuccessionDecl
especially-where
e
to a pointwher it only needsthe interbroughtby suchinfluences
to
colleCting
otherharmfulfactorlikeexcessive
ventionof sbnc
bad
of
on
essi
succ
a
be
could
achieveextinction.Equallyeffective
s
area
ding
bree
of
ers
in nuMb
reductions
following
weather-seasons
broughtaboutby othercauses.
firedamn,»to the interests
Fire - Two distinctseurces-of
: d.
cite
Blue maytie
of the Large.
planned
, deliberately
the
The firstof these'is-swaling
uragethe
enco
to
r
orde
earlyspringin
burningof coarsegrowth.in
hincludes
whic
and
urel
finerswardgrasseson the rough,untilledPastis allowedup to the
ing
thebreeding.areasof the butterfly.Swal
lypractised
end:ofMarch,afterwhichit is illegal.It is wide
hit is
whic
for
for thepurpose
beneficial
and is undoubtedly
groundis
same
the
and
intendedprovidedit is properlycontrolled betweensuccessive
od
not &waledtoo frequently.The optimumperi
but in the
to locationand climate,
swalingsmay vary according
eyearsis
thre
that
able
coastalareashereunderstudy,it is prob
Ll
rectin the economyof
abouttheTight interval.If thisis cor
theeintervals,rves
t -knnin-whether.sam
yet
pasturingflt-is-no
balance
ona nice.
ending
dep
e,
Blu
of the Large
best the requirements
Sheltering
swardand the-taller,
betweena ghortthyme-favouring
of this
further'study
heatherand gorseetc. Considerable
growth.of
canbe made,but one
pronouncements
is neededbeforeany confident
geBlue haslived and thrived
thingthatcan be saidis thattheLar
is knownand can be expected
with swalingfor as long as its historyeaboutin the practiceor
com
to continuoto do se unless-changes
butterfly.Deleterious
al
effectof swalingthatare inimic to them too greatan areabeing
resultfro
resultswouldalmostcertainly
swaling,the groundis likely
swaledin one year,for in the yearof
.To haveenoughground
to be too denudedto suitthebutterfly
it is essentialthat
son,
sea
every
in the rightcondition
available
llhaveremained
part of any one breedinggroundShast
a substantial
t iod probablyat lea threeyea,rs.
unburntfor a sufficienper
the
, smallerthe areaof a breedingground ed,
Clearlythereforethe
mfuland,as alreadynot
greaterthe riskof swalingbeinghar
for theplough.
ation,
of breedingareasby advanceof reelam
reductiOn
ng insttheLargeBlue. Too freauent
is one of the factors.operatiaga
likely.
be veryharmful,but sincethisis.
swolingwill alsocertainly
bad
n pasturequalityand ariseonly from k.
to causedeterioratioof
d lesslikelyris
, mustbe considerea
and ignoranthusbandryit
causingundesirably
mby
har
do
burningcan
Too fierceor prJlonged
g rfherbage.
dwa
damage.affectin
deep-sented
a muchmore
The otherfirehazardto theLargeBlue ig Sed
by accident
caU
e
fir
mer
swaling:it is the sum
serious.one'than
is
. ingthe swalingseason,thebutterfly
or carelessnessDur
immUnefrom
sumably
in the ants'nestswhereit is premer
underground
m
however,,froegg
by fireaboveground. In the sum nd
destruction
,
umn
aut
ergrou in the
layinEin Junu and July tillit goesund
overa
and fireextending
es
fir
e
to surfac
it is whollyvulnerable
n-on
tio
ina
pleteexterm
breedingareaeJuldreadilycauseits comatextent,notbeing
site. Summerfiiesare oftenof gre g,an it may be held that
that.
lin
undercontrolfrom the outsetlikeesWa
m someformersiteshas
fro
Blu
ge
e theLar
the disappearancof
of summerfires,causedeither
hnppenedfrom thiscause. The menace
ussedthrough
or malice- oftenby the sun'srays foc
by carolessneas
h expansion
wit
y
rapidl
bottles- is one whiehis worsening
discarded
orcar
mot
by
influx
and its aCcelerating
of the humanpopulation
.
dplaces
intocoastalareasand hithertoremotewil
CONSERVATION
the causesof decline,someattemptcan
Havingreviewed
ef conservatimand assessthe
) cussthe requirements
be made't-dis
of aimsand methods.
practicability
42
measurescouldbe aimedat:Conservation
wanion
of collectig and other.
or reduction
Elimination
1.
destructive encies.thatthepresentlow status
'Thereis littledisagreement
renderthe act of collecting
of theLargeBlue and its vulnerability
whileeven the taking
one of wantondestruction,
cabinetspecimens
•
purposesaimed'at
experimental
to be keptalivefor
of specimens
strictest
and the
consideration
needs.careful
aiding.conservation
is thoughtto be impossible
of collecting
control.Elimination
mootedare:mightbe,effectedand-mmA4a-methods
but reduction
Enclosure
In the knownbreedingareas,thisis not at presentviewed
of the landwith a
as practicable.Thereis multipleownership
bodies.
extentof it as yet in the handsof conserving
negligible
So far as is known,onlyone areaexistswherea limiteddegreeof
of
is alsoa possibility
ana -there
mightbe effective
enclosure
A publiccoastalfootpath,runs
if fundswere availal-le.
acquisition
groundsand the onlybar to
throughor adjacentto the'breeding
whichis..steadily
accesslies in a varyingdegreeof remoteness
becominglesSened..
(a)
'Policing
breedingcentres
Patrolby wardensof the mostimportant
.
be foundto serve
can
helpers
voluntary
if
be:practicnble
couldifonly
flying
the
during
expenses
their
to defray
made available
and funds:be
seasonof thebutterfly.In the absenceof any law prohibiting
but
onlyuse persuasion,
,
wardens'could
the takingof,butterflies
of
question
the
invoking
and
landlords
of
with_theco-operation
is likelythatsuchTatrolscould.beusefUllyeffective.
trespass,,it
(b)
Appeal
to refrainfromtakingtheLarge
Appealto collectors
societies,
Blue,made to thewidestpossiblerangeof entomological
not withoutconsiderable
and put intoaction',
has been suggested
It has been oppoSedon the scorethat
doubtas to its advisability.
givento the rarityof a species,the graver
the widerthepublicity
on the attitudeof_
a sad reflection
the menacefromcollectors,
in general:"
eollectors
(o)
of breedi- areas
to reelnmation
Resistance
of reclamation
As with the case of enclosure,_mithholding
or
purchase
by
certainty
with
accomplished
be
only
projectscan
leasingof landithoughmuchcan and has been doneby cultivating
goodwilland intereSt
contactwith farmersandlandownersto .secure
of-landsterilisation
for
payment
by
Compensation
conservation.
in
altitude
the
on
depend
will
Mach
practicable.
be
cases
may in sone
Trusts.
and financial
statusof CountyNaturalists'
2.
-
Controlof ScrubEncroachment
swalingand grazingcouldbe effecPlannedand controlled
with- tiveif the goodwilloffarmerscan be securedto co-operate
bodies. Mechanised
Trustsor Otherconserving
CountyNaturalists'
to carefUlplan,avoidcontrolled
cuttingof gorse,bracken,e'tc.,
is
of shelterto the extentriskedby &waling,
ing the destruction
hopeof success.
.triedout with considerable
being
Transolantation
The surveyhas shownthatthereare many fornerbreeding
be verygood for re-introduction
appear.to
groundswhereconditions
and
of theLargeBlue. A projectto achievethisis desirable
whenevera strpngcolonybecomesavailable
ahouldbe undertaken
canbe spared.
fromwhichtransplants
or assistedreari
Artificial
the view that
so far have not encouraged
Experiments
rearingawayfrom the naturalbreedinggroundsis likely
artificial
however,
to resultin any largenumbersbeingraised. It is desirable
in assistedrearingshouldbe carriedout,
thatfurtherexperiments
of conditions
if thesecouldbe directedto improvement
especially
in the naturalbreedingareaby boostingthe quantityof thymeand
of antsof therequiredspecies,togetherperhaps,with somemeasure
to preventescapeand protectfrompossibledestructive
of enclosure
weather.
agenciesand inclement
5.
of certainaspectsof the life
Furtherinvestigations
the
of cannibalism,
the incidence
historyare needed,especially
or assistedconditions,
of whichunderartificial
circumvention
production.In general,
and lead to increased
rightbe.possible
of the life historyand ecologymightwell lead
greaterknowledge
rearingin the field- in effect,to farmingthe
to ways of intensive
species.
Summor of Discussion
was whetherthe decline
The mainpointraisedin discussion
by a
of theLargeBlue in Britainwas paralleled
in the population
habitat.
or changein the nature,of themost favoured
disappearance,
Hr. Huntpointedout thatthereexistat the presenttime,sites
whichappearidealfor the insectbut whichare not populated.He
coloniesof the LargeBlue had
also thoughtthatin someinstances,
changesin the habitat,
by bad seasonsor temporary
been destroyed
again,therewere
but when thehabitatand weatherbecamafavourable
nearbyfor recolonisation.
no insectsavailable
44
es
Dr. Mooresaidthattwo notableIand:USe-changoccurred
effects
in the yearsafterthe lastwar whichhad very significant
of the LargeBlue. The rangeof
on the nuMbersand distribution
.alongthe coastwhere therewere extensive
the insectwas mainly
end a
with manyant:hills
grasslands
areasof roughcattle-grazed
to plough
greatdeal of thyme. Afterthewar, itbecame profitable
grants
with the aid of Ministryof Agriculture
up thesegrasslands
grasslands
and manyLargeBlue sites4sappeared. Surviving.coastal
were fencedoff in manycases,so thatscrubgrowthwas able to
alsoby the disappear,
aided
spreadin the absenceof cattle-grazing
ance of the rabbitaftermyxomatosis.He believedthatin suchareas,
if monaycouldbe foundto managethem
the changeswere reversible
effeCtively.
-45IN BRITAIN
OF DRAGONFLIES
THE CONSERVATION
N. W. Moore
Station
MonksWoodExperimental
SUIS1RY
are at the edge-oftheirrangein Britain.
Dragonflies
1.
and do not have
arc highlyadaptable
specie's
Most of the forty-three
of extinction
danger
in
are
two
preferences.Only
exactingecological
threatened.
become
could
and
but severalothersare verylocal
pollution,
of habitatand increased
Owingto destruction
2.
Britain
lowland
in
rarer
much
becoming
are probably
dragonflies
is becoming'
abundant.spedies
formerly
of
conservation
and so;active
necessary.
3.
-to
of dragonflies
. Enoughis knoWnaboUtthe requireMents
needis for
easy. The immediate
makeactiveconservationmeasures
research.
further
for
than
rather
knowledge
existing
of
application
the
Should'
Reserves
Nature
National
in
ponds
derelict
that
It is suggested
and otheraquaticanimalsby clearing
be made sUitablefor dragenflies
y southsideof the ponds,
on the
woodygrowth..especiall
surrounding
whichdo not possesstheM.
reserves
in
dug
be
should
and new ponda
that such
shows,
Reserveaand elsewhere
on National:Nature
Experience
canbe highlysuccessful.
measures
in National
a vide rangeof.habitats
The methodof selecting
4.
of
protection
NatureReserveshas resultedih the "accidental"
75%of 'theBritishdragonflyfaunain them. Rare species
at least
procedure.The degreeof
by the
are givetsomeprotection
by carryingout the measuressuggested
shouldbe increased
protection
and somespecialmeasuresShouldbe.takenin the caseof
in 3 above,'
veryrare species.
two or three.
INTRODUCTION
The Odonataare a smallorder- Of the 5,000or so living
occurin Britainand of thesethreeare
species,only fortY-three
t
populations...
migrantsand do not maintain.permanen
onlyoccasional
in 1946.butwas
was discevered
(Coen rion scitulum)
One.species
The fortyby the greatsea floodsof 1953..
exterminated
apparently
and the Britishfaunais
threespeciesbelongto ninefamilieS,
the worldfauna.
fairlyrepresentative-of
-4.6As with butterflies
the distribution
of British
dragonflies
is tolerably
well known and interest
in the group extond-s-beyond
that
of specialists
- though not to the extent
that reserves
are set
up expressly
to protect
them.
Dragonflies
are essentially
tropical;
the order is at the
edge of its range in Britain.
This is well illustrated
by the fact
that while thirty-two
species
breed
in Hampshire,
only one occurs
in the Shetlands.
Nest British
species
are tough and adaptable,
and
are not tied
to very exacting
ecological
conditions,
therefore
they
ere potentially
easy to conserve.
Their conservatiun
has been briefly
discussed
by Moore in Corbet
et al 1960.
THE NEED POR CONSERVATION
None of our species
is endemic and so there
is no special
responsiblity
to conserve
British
species.
Yspossible
exception
is 0
astra
curtisii
which is a rare
species
from the world point
of view.
By virtue
of their
size,
dragonflies
provide
special
opportunities
for ecological
and ethological
study and are of educational
and aesthetic
value,
and so are worthy of special
conservation.
measures.
Until
recently,
abundant
and conservation
developments
in farming
scale
throughout
lowland
folloWs:-
1.
suitable
habitats
for most species
were
measures
were not urgent.
But post-war
now threaten
breeding
areas
on a large
Britain.
The principal
threats
are as
Greatly
increased
efficiency'in
draining
due to widespread
use of mechanical
equipment.
This. means that many ditches
•and.ponds.vfrich
used to contain
Permanent
water arc now
only temporary
and so are useless
for dragonflies.
.flaily
.other
ditches
still
contain
permanent
water,
but their:rate
of flow, varies
much irlôrc than in the past - from nil to a
.fast
torrent.
These 6trearre are unsuitable
for all British
species,
which are either
adapted
to live
in fast-floming
water or to slow-moving
and str:tic
water,
but never to both.
Modern mechanical
equipment,
improved
methods of making
poor land productive,
and Government
grants
have increased
land reclamation,
particula”ly
of lowland
heaths
(Moore
1962),
and.of
small areas
of marshy land.
'Me loss of farm ponds
purely
arable
farming,
due
and
to changes
from mixed to
the provision
of piped water.
.."
In recent.years,
pollution
by pesticides
has been added
to that-of
industrial
wastes,
detergents
and sewage..
Particularly
hazardouS
are the organochlorine
insecticides
which are now universally
distributed
in British
river
systems
(Moore and Walker,
1964 and unpublished).
The
—
— 14-7
like
is increasing.This,.
use of aquaticherbicides
many
altersthe habitatby'killing
treatment,
'mechanical.
of
the development
and by encouraging
aquaticpl'ants
Theseoftenhave to be treatedlater
nesistant.species.
In generalaquaticherbicides
herbicides.
-bi-more-foxic
to insects;
non-toxic
are
in-use:-today-relatively
weed
destruction.of.aquatic
large-scale
nevertheless,
of thewaterto an extent
may resultin de-oxygenation
whichis.likelyto be harmfUlto aquaticinsectslike
whiehcannottolerateverylow oxygentensions.
drag,:nflies
work has been done to evaluatethe effects
No systematic
that
above,but the generalimpression
of the factorsmentioned
lowlandBritain(e.g.the Fens and Romney
in cultivated
dragenflies
yearsago,is
Marsh)are muchrarerthantheyto twenty-fivc
is unlikelyto improve
sharedby'many dbservers.Sincethe situation
thereis a good casefor takingactivemeasurestowardsconserving
Britishdragonflies.
policyby Whichits National-Nature
The NatureConservancy's
has led to
Rescryesare obtainedto covera wide rangeof habitats,
bothby design.
pondsand streams,
of manyfreshwater
the protection
and accident.And whileno reserveboundaryhas.beendrawnspecibreedingarea,.atleast
Specialdragonfly
ficallyto,protect-a
.occurin NationalNature
breedingspecies
thirtyofLthe-forty
d
to
havebeen_designate
S.S.S.Is.
.
Reserves(see-appendix)Five
protectrare speciesof dragonflies;and NationalNature-Reserves
to practically
give sore sertof protection
together
and S.S.S.Is.
mnnyof the
all theBritishspeciesof Odonata.NeVertheless,
and mightwell not surviveif
are very-small
nrotectedpopulations
destroyed.Therefore,
werebreedingplaceSoutaide.them
neighbouring
breedingplacedof dragonfly
to increasethe'number
it is desirable
roscrVes..
withinexisting
IN RESKRVES
POPULATIONS
DRAGOTTLY
1EASURESTO INCREASE
speciesaretairly catholicin their
Sincemostdragonfly
easily. The followthesecan bc providedrelatively
requirements,
be bornein mind:must
requirements
their
s
about
ing generalisation
1.
2.
water.
e
permanent
All species-requir
are foUndin ponds•
of theBritish.speoies
Thirty-five
riversorboth. Only'five(Agrionvirgo,
or-alow,moving
Cordulegaster.
tennis, Go hus vul atissimus,
Platycnemis
Ox astracurtisii are confinedto flowingwater.
boltonli,
Thirty-three
to thirty-four
speciesare foundin neutral
or alkaline'
waters; mostof themwill tolerate
acid
conditions.
Only six or sevenspecies(Coenagrion
mercuriale,
Cori rion tenellumAeshnacaerulea,
Orthetrum
coerulescens,
Leucorrhinia
dubia,Sympetrum
danaeand
perhapsIschnurapumilio are confined to acidwater.
All speciesare absentfrom shadedpondsin woods. The
extentto whichthisis due to de-oxygenation of the
waterresulting
frembacterialactivityon deadleaves
or to the lack of plantsand associated
animalsdependent
on themis not known.
5.
All speciesare territorial;
as a resultpopulatindensity
of adultmalesis low and dispersal
is generally
efficient.
On the otherhand,localpopulations
of rare speciesare
vulnerable
to overcollecting,
sinceit is quitepossible'to
collectout the wholemalepopulation
at one siteafterthe
maindispersal
has occurred.
6.
A few speciesare known to hnve specialrequirements:Coen rion armatumis apparently
dependent on frogbit
H drocharis
morsus-ranae
L.); Erythromma
najas isdependent
on waterliliesand or Potamogeten
spp.; Sympetrum
sanguineum
is probablydependent
on Typhalatifolia
and/or
Equisetum
spp.
For most speciesa smallunshadedpond surrounded
by aquatic
vegetation
willprovidea suitablehabitat.Recentstudies(bbore
1964 and unpublished)
show thatthe creationof permanent ponds
lesdSrapidlyto colonisation.
For example,bomb holesmadein
the 1939-45 war on heathland
of theArne S.S.S.I.supported
fifteen
specieSof dragonfly
by 1954. Twelvespecies'of
dragonfly
were
seen at twentypondsdug for experiments
on aquaticherbicides
at
W)odwalton
Fun N.N.R.only threeyoarsafterdigging.Felling
treesarounda shadedpond at Ham StreetWoodsN.N.R.,whichhitherto
supported
no dragonflies,
resultedin it beingcolonised
by seven
speciesthe following
summer.
Most reservescontain ponds,but many,especially
thosein
woodlandreserves,
havebecomeso overgrown
thattheysupportfew
or no'dragonflies.
By fellingtreesroundsuchpondsand thereby
lettingin the lightand reducingthe leaffallinto them,woodland
pondscan be made suitablefor dragonflies
and otheraquaticanimals.
It is particularly
importantto fell treeson the southern
edge of
the pond:since
Ttheseprovide..most
of the shade.
.
thnt,unlessthereare specific.reazons
It is suggested
cleared
be
Should
Reserves
Nature
National
on
ponds
againstit, all
whichdo not
in thisway,and pondsshouldbe dug on thosereserves
dragonflies
common
of
nuMbers
the
means,
this
By
them.
possess
yet
increased-on
greatly
be
would
and numerousotheraquaticspecies
speciesof dragonfly
NationalNatureReserves.Abouttwenty-two
way.
this
in
helped
be
would
RARE SPECIES
SPECIALMASURES FOR CONSERVING
measuresaro onlyrequired
At presentspecialconservation
for a few species:-
2.
Ox astracurtisii- thisspecies is confinedto
Spain,the Southof Franceand one or two riversin
England. Its.most famousBritishsiteis ari
southern
in which
of the riv..2pollution
but accidental
5.5.5.1.
siteis'
the
Since
it.
exterminate
easily
could
it breeds
should
eOnsideration
interest,
zoological
general
of great
be givento makingit a Nationalor LocalNatureReserve.
Coe -rionarmatum- in Britainthisspeciesis confined
to one site,whichis'an5.5.5.1.The habitatony have to
if the sr:eciesis to.Survive.
artificially
be maintained
confinedto a few
Coen rion hastulatum thisspecies-is
At Presentthese
smalllochsin theHighlandsof SCotland.but a watchshouldbe kept
do not appearto be threatened
on the species.
Aeshnacaerulea;.
Ischnurapumilio,keshnaisoceles,
Leucorrhinia
motallica,
ochlora
Coen ribn mercuriale,'Somat
the.best
spme.of
but
sites
several
at
dubia- all occur
protection.may
active
and
the.War
since
lost
oneshavebeen
are scheduledas
of the-sites
becomenecessary.Sevc.tral
S.S.S.Is.
REFERMICES
C. and Moore,N. W.
MOORE,N. W. 1960 in Corbet,P. S., Longfield,
Dragonflies.London.
MOCRE,N. W. 1962. The Heathsof Dorsetand theirConservation.
J.Ecol.,50,.369-391.
'among.
Competition
MOORE,N. W. 1964. Intra-and Interspecific
49.;71.
J.Anim.Ecol.,
(Odonata).
Dragonflies
W. and WALKER,C. H. 1964. OrganicchlorineInsecticide
MOORE,'N.
Residuesin WildBirds. Nature,201,(4924),1072-1073.
- 50 APPENDIX
Species(43)
SpeciesprotectedConfined
Present specifically
on to flowing
Statusin N.N.Rs
S.S.S.Is.
water
Gomphusvulgatissimus
2
Cordulegaster
boltonii
1
Brachytron
pratense
1
Aeshnacyanea
1
Aeshnajuncea
1
Aeshnagrandis
1
Aeshnacaerulea
2
Aeshnamixta
1
Aeshnaisosceles
3
Anax imporator
Cordulialinaenea
1
Somatochlora
2
metallica
Somatochlorn
-u-ctica
2
curtisii
Oxygastra
3
Libellulaquadrimaculata 1
depressa
1
Libellulafulva
2
Orthetrum
coerulescens
1
Orthetrum
1
cancellatum
Sympetrumstriolatum
1
1
nigrescens
Sympetrum
Sympetrumvulgatum
SympetrumfonscoloMbei IA
IA
Sympetrum
flaveolum
Sympetrumsarguineum
Sympetrum
1
danae(=scoticum)
Leucorrhinia
2
dubia
1
Agrionvirgo
Agrionsplendens
1
Lestessponsa
1
Lestes.dryas
2
Platycnemis
1
pennipes
Pyrrhosoma
1
nymphula
Ischnuraclegans
1
Ischnurapumilio
3
Enallagmacyathigerum
1
Coenagrion
1
puella
Coenagrion
pulchellum
1
Coenagrion
mercuriale
2
Coenagrion
hastulatum
3
Coenagrion
armatum
3
ErythreMma
najas-Ceriagrion
tenellum
Totals:
4.3 30t 6
1. Wide distribution
in
suitablehabitats
27
2. Confinedto localareas 8
3. Very local
5
m. Migrants
3
G. Generallyfoundin-flowing
waters.
-51Summar of discussion
Dr. Mellanbysaidthattherehad been a greatincrease
in recentyearsin the areaof waterin thiscountry.He wondered
how important
gravelpits and reservoirs
were for the conservation
of dragonflies,
now that thesehabitatshad becomemorenumerous.
Dr. Mooresaid that the interestof reservoirs
for dragenflieswas slightin the firstfew years,but theycan becoMecolohised
very quickly.If therewas a greatdeal of fluctuation
in the r
watertable,thenonlycertainspeciesof dragonflies
can adåpt 4themselves
to theseconditions.Gravelpits tendto get polluted
by refuse,but sometimes
theycanbe of very considerable
interest.'
Mr. Eltonenquiredwhat sort.ofinteraction
therewas
betweendragonflies.
Do theyfor instance,
establish
territorieS?
Is thereinterference
betweenspecies,and havedragonflies
attempted
to pairwith thewrong species?
In reply,Dr. Mooresaidthattherewas a greatdeal of
interspecific
fighting.In the caseswheretwo dragonflies
of a'
diffetent
:typefought,the largerusuallywon The femalelays her eggswhen the male is not presentso thattheyare not generally
affected
by interspecific
strife. Mr. Skellamreferredto Dr.
Moore'sstatement
thatof fortyspeciesof dragonflies
in Britain,
twenty-eight.occur
in NationalNatureReserves,and thatfive
Sitesof SpecialScientific
Interesthad been established
partly .
becausetherewas a dragonfly
interest.Mr. Skellamaskedhow many'
extrasPecieshavebeen addedto the twenty-eight
by establishing"S.S.S.Is. In reply,Dr. Nboresaidaboutthree.
- 52 -
BkSIC ECOLOGICLL KNO,LECGE ItrUIRED
FOR INVERTEMLTE
CONSERVATION
E. Broadhead
Department of Zoology, Leeds University
All ecologists would doubtless agree that the more v.e
know about the mechanisms of survival and'maintenance of populations
of animals in the field, the more intelligently and more surely shall
we be able to modify the environment in favour of species we wish
to encourage and also to modify it to the detriment of species we
be
wish to discourage.Both theseaspectsshould,in my opinion,
and
procedures of
of conservation
kept in mindwhen we arc thinking
with regard
- particularly.so
basisof conservation
the theoretical
therearc so many
s
because
of invertebrate
to the conservation
amongthem.
and kindsof interaction
speciesand so many interactions
of inverteto thinkof the conservation
important
It is therefore
of varietyof speciesin any
bratesin termsof the maintenance
one placeor area. In sayingthis,I am not therebydenying
rare specieson aesthetic
someoutstanding
of conserving
the importaxce
interest
groundsor becauseof somespecialscientific
or educational
whichexistsas a small
the LargeCopperbutterfly
as for instance,
in
Fen. As Dr. Duffeyhas indicated
in Woodwalton
localpopulation
this
of
the eggs and larvae
to thisSymposium,
his contribution
markedfor census
butterflymay easilybe countedand the adult's
uniqueopporalmost
estimates.Here is an
movementand longevity
dynamics
tunityfor studyingin verygreatdetailthe population
a studywould
Such
range.
its
of a veryrare speciesnearthe edgeof
procedure.
haveundoubtedvaluein decidingconservation
of varietyof species,
regardnow to the conservation
Yiith
with the
is concerned
required
basicinformation
the most imr.ortant
I
area.
given
a
in
exist
can
that
species
nuMberof herbivorous
greater
much
a
has
second
The
approach.
of
lines
two
to
wish to point
important
unduubtedly
have
Both
first.
the
than
content
theoretical
practicalapplications.
briefly,althoughit is the kind
vTith
The firstI shalldetal
majorityof conservation
great
the
probably
which
on
knowledge
of
in practice.If we know
based
be
s
will
invertebrate
for
programmes
of speciesof
occurrences
between
correlation
the
about
enough
of associations
or
species
plant
of
occurrences
thc
and
herbivores
of the habitatto
of plant species,thenwe can plan management
of the maximumnumberof animalspecies.
encouragethe maintenance
to TableI whichindicates
reference
by
illustrated
be
may
This
species
of the three.psocid
each
of
density
between
the relation
—53
with
namedtogether
e.g.the 6 treespecies
andplantasSOciation,
but
theycarry; thesepsocidsfeedupontheepiphytes
theepiphytes
food
by the amount,of
entirely
determined
arehtot
theirdensities
is Eli socUsMclachlani
We seehow,restricted
of tree.'
on each.kind
is of special
othertwo species.E. mclachlani
with'the
compared
psocidsin thiscountry
as one of the fewlichenophilous
interest
as beingonlyimperfectly
interest
and amongthese,it is of special
on thedeadlarch
adaptedto a lichendiet. It is muchmorenumerous
on the treethanon thelivingones. TheHa Hireplantation
branches
on theeastShoreof MaihamTarnis an idealhSbitatfor thisspecies,
elsein the
herethananywhere
being_much.denser
thepopulations
Pennines.
Table 1
in
on sixtreespecies
species
Densityindicesof threeElipsocus
1952
thewoodedareasaroundMalhamTarn,SepteMber
Total-
No. of
1) includes
26 fromone sample;31 fromanother..
2)
38 fromtwo samples.
includes
- 54 Teble 2
of PsocidSpecieson Larchin lateSummer
Abundance
TheRelative
fromtwenty-four
The figuresehowtotalcaptures
1954. Equal
Septetber
24th
August
8th
localities,
and from
living
from
taken
were
nuMbersof samples
deadbranches.
151; Metylophorus
immaculatus
269;Stenopsocus
guestifslica
gibbosa29; Trichadenotecnum
35; Psococerastis
nebulosus
12; Cuneopalpus
pedicularia
15; Lachesilla
sexpunctatum
3;
variegata
Loensia
7;
cyanope8; Caeciliusburmeisteri
unident14
and
1;
cruciatus
Graphopsocus
3;
L. fasciata
ifiednymphs.
treeswouldclecrly reducethepopulation
Brashing_the
suchas these,one couldeasilyset out a
facts
With
enormously.
population
to allowthe densest
designed
nrocedure
management
woodln.pd
graprie.
management.pro
A
course,
Of
EirentUally
exist
to
mclachlani
of
nutber
large
a
to
reference
with
designed
be
would
area
for any one
and some
be necessary
wouldinevitably
of.species.Compromises
than
attention
more
merit
to
considered
be
doubt
no
specieswould
others.
of
therecording
- involving
This kind of.information
or
food
different
to
eference
r
with
sites
different
densitiesat
all
of
first
would-collect
one
kind
the
is
resources
substrate
point. It is not
. is a naturalstarting
for mostinvertebratesIt
usefUlfer con'servation
bu.tit is extremely
verYdeepknowledge
•-purposes._
Elipsocus
hyalinus
Amphigerontiabifasciata
Elipsocus
westwoodi
1961-
Philotarsus
picicornis
Mesopsocus
immunis
Mesopsocus
unipunctatus
••
1.:•;;;;..•
E
200
160
Mesopsocus
immunis
120
80
40
160
Mesopsocus unipunctatus
120
80
40
AAR mAy JUNE
JULY
AUG
SEPT
tAAR APR mAY JUNE
JULY
AUG
SEPT
ApR
mAY
JUNE
JULY
AUG
SEPT
/AAR
ApR
mAY
JUNE
JULY
AUG
SEPT
á
ov. 30
a
25
3
20
1958
1959
15
1960
1961
Pleurococcus
Nymphs
Pleurococcus
Nymphs
,1 Adults
/ 1
Adults
Nymphs
I
Nymphs
Adults
I 1 I Adults
600
/Total
Total Mesopsocus
Mesopsocus
All other psoc ds
All other
•
psocids
—• —
00
- 300
Mesopsocus immunis
Mesopsocus immunis
Mesopsocus unipunctatus
Mesopsocus unipunctatus
:Ea)300
2-0
Pleurococcus
Pleurococcus
per unit psocid
biomass
per unit psocid
biomass
1•0
0
MAY
JUNE JULY
AUG.
SEPT.
APR
AY
JUNE
JULY AUG.
APR. MAY
JUNE
JULY AUG. SEPT.
APR. MAY
JUNE
JULY AUG.
SEPT.
á
-55-
The secondline of approachfollowsfrom the first.
_given_now
fundamental-problem:.
It is.the-tonsideration-of-a-more
by
allowfor its exploitation
what conditions
only one foodresource,
muMberof speciesof herbivores?Lot me developthis
the greatest
to sometypicalwoodlandinsects,thePsocoptera.
with reference
.arevery commonin uplandareasinBritain. On the
Theseanimals
bark of larchtrees,theycanbe as abundantas are the big herbivorousmammalson the plainsin Africa.2Kendeigh(1961) gives
,--for-the-biemass_of_grassland
rom
figures-ranging-f0.18-to-8.34.g./m
the
in Africaand NorthAmericaand we haveestimated
herbivorms
specieson a comp1ete2
of twoMesopsocus
weightof the population
larchtreeat Harrogateat the time'ofpeakbiomassas 1.18g./m .
of bark surface.
on
Table2 givesa listof all psocidspeciesoccurring
in late summer.
theirrelativeabundance
larchtreesin Yorkshire.with
speciesand sevenabundant
lichenophilous
Therearc two Omandant
al mixturepresenton
thePleurococcus-fUngspore
speciesexploiting
of Closelyrelatedspecieswhich
the z;itups
thebark. Table3 indicates
meMbers
occuron larch. Exceptfor the two Mesopsocus.species,Ahe
on a singlelarch
in.habitat
difference
of eachpair dhowa pronounced
of alllthe
individuals
in nuMbers'of
tree. Figure1 ahowsthe.change
together
of psocidoccurring
ng
six commonPleurOcoccus-eatispecies
tile-feet -of-theliving-larch-branches-in.a
on the-V.:Si-Halat Harrogateover fouryears. Elisecus h alinusand Amphigerontia
on:thedeadbranchesthanon these
occurmore abundantly
bifasciata
on the
eodursmoreabundantly
westwoodi
livingbranches.Eli so'cus
and the two
livingthanon the deadbranches.Philotarsusicicornis
speOiesare asabundanton the deadbranchesas on the
Mesopsocus
the closest.
speciesdhowtherefore
livingones. Thosethreecommonest
the samefood..
and all are exploiting
in habitatrequirement
similarity
intobiomass,and
resource.In Figure2, the nuMbersare.converted
Thp.miXedspecies.
of food.isalso indicated.'
the changein.abundance
foodlevel
effect'on
exertan appreciable
of Mesoesocus
populations
on,thesotwigs. They take the foodearlyin the season,andthe
Onlya
fourlatersummerspeciedtogetheraccoUntlfor
remaining
theMesoosocus?
do
th:tri
food
smallbiomassand'takemUchloss
relatively
OntheSelarchtreesthen,we havea greatdiversityof
the samefood.
and-eating
together
specieslivingherbivorous
a closed.homogenous
in
that
dhow
to
evidence
of
plenty
iS-now
There
requirethe same
they
if
co-exist
species.cannot
two
environment,
the
expressed.in
was
this
way,
resource.In a.purelytheoretical
non-predators.
two
between
interaction
cf
equations
LotkaArolterra
work by much laboratory
it has been demonstrated
Empirically,
and
confusum
mixed*Tribolium
on
(1956)
work
Park'S
particularlY
populations.
castaneum
- 56 -
T&blo 3
.Foodresources
and habitatsof fivegroupsof Closely-related
psocid
speciesonLarch in Yorkshire
FOOD
HABITLT
Proportions
of
populationon
above600feetabovesea level;
xx.- below 600 feetabovesea level.
The possibilities
.of reducingor obviating
the results
of competition,between
two suchspeciesby introducing-environmental
heterogeneity
are several:•Co-existence
of potentially
competingspecies.Two species
requiring
the samefoodresources
may co-existin a stable
and permanent
way providedthatthe two nopulations
are
Separately
controlled,
eachby its own'density
dependent
factor,'at
a leYelbelowthatwhichwouldresult.in
food
shortage.
Co-existence
of comneting
species.Speci:esi
competing
fer
food(Le...the
exploitation
type
.ofcompetition).
may
neyerthelessteo-exist
provided.that.there
existSa balance
of advantages:(a) in time. As Park has Shown,the competitive
prowesS
of two sneciesmay be reversed
by changeof climatic
environ&mt. An cnirironmont
whoseclimatechangesin
a periodic/
- 57 -
way with the appropriate frequency about an appropriate
mean value may then allow prolonged co-existence of two
competing.species Whose fluetuations would be out of phase.
On the basis of what is now known-oboUt mixed_species_populations of Tribolium, this system could eqsily be engineered
in the laboratory. In the field, seasonal changes in frequency of the chromosome types "Standard" and "Chiricahua"
in Drosophila at Pirien Flats in California (Dobzhansky; 1943)
approaches this system most closely.
(b) in space. Co-existence is possible provided that one
species.is favoured (or has a refUge) in one part of the
habitat and the other species in another part. Movement
from these refUges would produce a zone.of coaeXi-steneewhich would bcestable so long as the refuges remained invidable.
_
(c) Skellamrs model cf.two hypothetical plant species (1951)
ineorporates a balanee of advantages.- The environment is
conceived as carrying a certainnuMber of sites suitable for
the growth of.seeds of species A and B. If seeds of the
two species fall at the same plabe,..A.minsbecauSe it grews
more quickly. Both.plants are mortal however, so releasing
vacant places and species B disperses its seeds-More vddely
than does.species A so.that there will always be some
sites reached first by B in which.it can then maintain its
position.
(d) A common enemy (parasite or predator) could act as a
stabilizer to the IpopulationsiftWo competing species
provided that dt.is habit.rforming with respect-to preference
Tor prey• species, i.e. it comes to prefer whichever species
one.
happens to be the...commoner
(a) We can conceive an extension to Park's Tribolium model.
Park found that alternative outcomes were possible from the
struggle for existence in-replicate mixed species populations
of Tribolium under identical cliMatic conditions. By allowing a limited migration between replicate vials, co-existence
would be greatly prolonged, although it would not be stable.
-(b) This is not far removed from tho transient co-existence
of two competing species - one of which is moving towards
extinction.
These ideas are-hypotheSes one priori models which are
avellable to the field ecologist trying to interpret what exactly
is happening in any given situation in the field. In order to decide
which model or coMbination of models is. appropriate, the field data
required'must relate to changes in time. I.wili now illustrate
thiSby brief reference to a study of the population dynamics of two
co-existing Hesopsocus species on larch trees - a study conceived-
- 58 -
in part as an attemptto utilizethe conceptual
schemesarrivedat
fromlaboratory
studiesof mixedspeciespopulations.The final
interpretation
is summarized
by model1 above.
In a plantation
at Harrogate,
two Mesopsocus
snecies,
M.
immunisand M. unipunctatus,
feedon-thesamefoodand live together
on the sametreesat the sanetime. The eggsand each of the six
nymphalinstarscan be identified
to species.Weeklysampling
from
the terminaltwo-foot
regionof livingbranchesin 8 blocksof trees
providedthe observational
data and the remarksmadebelowall apply
to the population
in thissampledzoneof the tree.
Figure3 givesa quantitative
pictureof the lifehistory
of the two speciesin 1958 and the flightperiodof the mymarid
Alaptusfusculus,
whichparasitizes
the eggs of both theseMesopsocus
species.Anoth..x
parasite,
a braconid,attacksthe 5th - 6th instar
nymphsof both thesespecies.From thisinformation,
lifetablesof
M. immunis,H. unipunctatus
and of the mymaridcan.be.
constructed.
Table4 showsthe life tablefor M. immunis.From theselifetables
and a furtheranalYsisof the resultson whichthe life tableswere
based,we can construct
an empirical
mathematical
model. Figure4
indicatesthe modelqualitatively
and Table5 givestheparameters
of thismodel. The Mesopsocus
life cycleiS dividedinto8 stages
and the quantitative
expressions
describing
thesestagesfallinto
threetypes:mean survivnrship
valuesfor stages1,.3, 4, 5. This is
(a)
the equivalent
of simplifying
our modelby regarding
mortalitiesin thesestagesas constant. It is alsothe
expression
of inadequacy
of the model,whichcouldbe
improved
by incorporating,
in placeof these.survivorships,
equations
describing
the relationship
betweensurvivorship
and eitherclimateor animaland plantenemiesor both.
the use of thu Nicholsonian
host-narasite
equationfor
stages2 and 6-,
an equation
whichis itselfana priori
-Model.
for stages7 and 8, empirically
deteiiittndd
relationships
betweennuMbersof femalesat thebeginning(N ) and at
the end (N ) of the preoviposition
periodand Eetween
N and the1nuMberof eggslaid. Both theseequations
for
each speciesdescribeequilibrating,
i.e.densitygoverning
processes.
When thismodelis workedout (Fig.5) fromarbitrary
initial
densities,
the numbersof the fourspeciesreachequilibrium
values
closeto the densitylevelsrecordedin the field.
Thiskind of'emPirical
modelcannotimmedi
ntelybe translatedinto a conservation
Programme,
but it can pointthe way with
referenceto theseparticular
speciesand,itcan.do_this.in
two ways:-
2000
1500
1000
500
fig 3
s
Nymphs
In
egg
21
APRIL
eggs
a
26
2
9
s_
l
sr
30
adtults
No of
adult
mymarld
on
00 twig
23
1ST
adults
16
RJNE
in host eggs
hs
in_hosts e
M. unipunctatus
hatching
co mplete
n
M. irnmunis
19
mAY
12
Mymarids
free nymphs
hatching
complete
5
f ree
\._ Mymarids
28
in\
Nymphs
l
7
Adeus
/
/
21
JULY
IA
\\
28
Eggs
Eggs
AtjuGusT
18
25
Flhght ° f
adult .n1)11-flan
'd
4
8
SEPT.
Empty
eggs
Dead
eggs
in
Larval
p orasite
eggs
Nymph
in
eggS
Em
Dad
a fasste
NymOs
30
I
255
6.3
58.6
9.6
128
15.
199
522
á
LifeHistoryof two Mesopsocus
speciesand theirparasites
EACH MESCPSCCUS
SPECIES
AUG.
MYMARID
BRACONID
Egg Mortality
fungalattack
mymaridattack
adults
u'e -uP
loss of larvaeby
suckingpredator
suckingpredatorand
winterloss of egg
by winterloss of
batchesfromtwigs
host eggs
=
NV'
OCT.APRIL
MAY
juNE
NymphalLosses
birdpredation
braconidattackA
U.
Adults
mortality
and migration
duringpreoviposition N -N
o 1
period
JULY
Egg-laying
- u'e-aP
-(pupae)
N1-Eggs
Figure4
adults
(la:Lrvae
pupate in
soil)
á
60
10
20
30
40
50
Fi3.5
(Scale X 10
for mympricl)
of
larch twig
128
10"lengths
per
Number
70
80
90
5th
Generations
loth
15th
20th
M. immunis
Braconid
Mymarid
M. unipunctatus
25th
21
19
425
46
á
at Harrogate.
lamas1958-61
BudgetforEasopsocue
andNatality
Uortality
1957- 58
Stage
Period(x)
Approx.
duration
in weeks
Factors
Nos.
surviv'g Nos.
et begin dying
duringx
of x
1x
1183
9
EGG
attach
ibmarid
8:deadSap
attaob
Fungal
producing
Predators
of
Promcompletion
to onegg-lsying
zet at winter
Total
rain
26
loseof egghatches
of
Prombeginning
to
hatching
attackon
braconid
6tb instar
11
Fredatort
of
Prombeginning
attackto
braconid
peakadultincidence
2
attacking
7:caconid
5thand6thinstars
4
of adulta
Predation
andtheiroviposition
ADVIL: Yrompeakincidence
of adultsto oompl
tionof egglaying
253
35
61
wipty
egga
Winter
di
-
834
1958 -
1960- 61
1959- 63
59
Percent
ageIon
1004x 1x
21
3
175
dx 100
15
29
48
92 - 17
dX 1
i34
113 17 378
g
55 9
45
3:
20
61 -13
-_
248
1X
647
94
72
5
-
dx 1C0
1,
..f;
12
51
29 421
45 186
42
74 2
701
90
i
25 11
18
I
eggs
"-otal
70
7
4
-642
475
1961- 62
6
9 -125
-160
647
376
81
I
- 61 Table 5
THEPARAMETERS
OF THE MODEL(NUMBERSPER 16 TWIGSAMPLEUNIT)
Period
1.
Factor
Eggslaid
Eggsavailable
(u')
-=
Fungalattack
Eggsavailable
to
mymarid
Eggs surviving
(u)
rymphswithineggsat
beginningof winter
Nymphswithineggsat
Nymphsreadyto hatch
onsetof winter
in spring
Nymphsreadyto hatch
5th-6thinstarnymphs
from eggsin spring
available
to braconid
6th instarsavailable
(u')-> 6th instarssurviving
(u)
6th instarssurviving
No
8.
N
1
Mortality
+ Migration
Oviposition
Regressions
below
Attack-bysucking
predator
Winterlass of egg
batchesfromtwigs
Predation
by birds
and lacewings
Braconidattack
6thinstarnymphs(No)
Femalesat end of preoviposition
period(N1)
Eggs laid
PARASITEs
Mymaridsproduced
by
oviposition
in hosteggs
Braconids
produced
f.
PariolN
N
o
1
8. PsriodN 3 Eggs
1
2+6 Parasite
attack
Mymaridsattacking
the
following
year (p)
M.
immunis
Survivorship
• 0.879
(n = 5)
Area of discovery 0.003096
(a)(n= 4)
0.879
Survivorship
(n = 4)
0.612
Survivorship
(n = 3)
0.185
Survivorship
(n = 4)
Area of discovery 0.1807
(a)(n= 3)
Sex ratio(2:total)0.597
Regressions
belcw
Mymaridattack
Eggs surviving
mymarid
attack
6.
Parameter
M. unipunctatus
0.955
0.01656
0.776
0.974
0.267
0.0653
0.506
:-
Mortalityof hosteggs:
from suckingpredator
and by winterlossof
hostegg batches
Survivorship
0.879
0.776
Survivorship
Survivorship
0.612
1.0
0.974
1.0
Braconids
attacking
the
follcwing
year (p)
E U A.T.I 0..N..
S 13S.E.D
M. immunis
logeN1= 0.809+ 0.364logeNo
M. unipunctatus
logN1=
e
1.323+ 0.305loge ()
eggs = 16.16+ 28.38logeN1
logeeggs = 3.564
+ 0.694logeN1
u = u'e-aP
á
- 63 -
it helpsus toexplorespeciesinteration
in.a speculative
way in so far as tho modelis greatlysimplified
and random
variables
are excluded;
itean.be.elaborated_by
substituting,
for the'constant
survivorship
terms,equations
relatingmortality
(or
migration)
with otheranimalsor with criMatic.factors,
e.g.
nymphalmortality
canbe relatedto lacewingand bird
nuMbersand for the adultperiodthe regressions
citedcan
incorporate
temperature
as a significant
additional
term.
This wouldgiveus somepredictions
of the consequences
of
changesin climateor of.otheranimals.
'Finally
I shouldlike to.cndmy talk with a synthesis.of
the mainpointsI have justdiscussed
and a commentupon the relation
of basicecological
ideasto conservation,
(seeFig. 6). I have
mentioned
someof the a priorimodelsconceived
in a purelytheoretical
vay independently
of considerations
of the properties
of narticular
speciesof animalsor plants. These,historically
specking,
are
gradually
testedagainstour ecological
experience
at the variouslevels
of complexity
- initially:with
the simplestaniffials
in the simplest
and mosthighlycontrolled
laboratory
situations
(e.g.Park'sTribolium
cultures).Theselaboratory
investigations
themsolveS
resultin
models,simpleempirical
models,whichthe fieldecologist
can use,
and test,with referenceto naturalpopulations
of animalsin uncontrolledor largelyuncontrolled
environments.
Thesefieldstudies,as
for instance,
in the case of our own work on Mesopsocus,
can also
resultin simpleempirical
models,whichgiveus a littlefurther
insightintothe mode of workingof thatmuchmore complexentitythe wholebioticCOMMunity
in any one region. This summarizes
the
structure
of researchin population
ecology. In long termview,
it shouldimpingeto an important
degree,upon the ways in which
conservation
is viewedand the way conservation
programmes
are
develOped.In shorttermview,the kind of information
I indicated
in my firstlineof approachwillprovidethe mostvaluableinformation
for developing
managemnit
plansto conservespecified
invertebrate
animals.
REFERENCES
BROADHEAD,
E. 1958. The psocidfaunaof larchtreesin northern
England - an ecological
studyof mixedspeciespopulations
exploiting
a commonresource.J.Anim.Ecol.,27,
217-263.
BRa.DIT&O,
E. and DATTA,B. 1960. The taxonomyand ecologyof British
speciesof Peripsocus
Hagen(Corrodentia,
Pseudocaeciliidae).
Trans.Soc.Brit.Ent.,
14: 131-146.
BROADHEAD,
E. and THORNTON,I. W. B. 1955.An ecological
studyof
threecloselyrelatedpsocidspecies.Oikos,6(1),1-50.
on
populations
A. J. The Mesopsocus
E. andWAPSHERE,
BROADHEAD,
closely
two
in
regulation
larch- a studyof population
a commonfoodresource.
exploiting
species
coexisting
._related
(Inpress).
IX. Temporal
populations.
of natural
T. 1943. Genetics
DOBZHANSKY,
of Drosophila
of populations
changesin thecomposition
28,162-186.
Genetics,
pseudo-obscura.
, pp.
468
Prentice-Hall
S. C. 1961. AnimalEcology.London,
KEVDEIGH,
J., PLRK,T. andSCOTT,E. L. 1956. Struggle
PARK,T, in HEYMAN,
and statmodel:biological
forexistence.The Tribolium
on Math.
Symposium
Berkeley
isticalaspects.Proc.3rd
41-79.
andProbability,
Statistics
populations.
in theoretical
J. G. 1951. Randomdispersal
SKELLLM,
196-218.
18,
Biometrika,
AND FIGURES
SOURCEOF TI-LBLE,S
1955;Table2 from
& Thornton,
Table1 fromBroadhead
1958andBroadhead
1958; Table3 in partfromBroadhead,
Broadhead,
& Datta,1960.;Tables4 and5 andFigures1 - 3 and5 fromBroadhead
(unpublished).
andWapshere
FIGURE6
IDEAS
IN RELATIONTO BASICECOLOGICAL
CONSERVATION
VIEN
A LONG-TERM
towardshighergradesof abstraction
of our understanding
the direction
The structure
all
Its longterm
on
influence
conservation
theoryand
practice
at
experience
testedagainstour ecological
are successively
levels
the ideasexpressed
these
these
are
our
practical
loreetc.
the ideasand wisdomimplicitin traditional
i.e.the
madeexplicitand furtherelaborate,
theory
population-ecological
of developing
utilisation
practice.
for conservation
starting
points
The startingpoint
of The
historically
i.e.
NatureConservancy
of
the declaration
NatureReservesand ther
achieve
by conservation
and political
executive
action
the scientific
development
selection-basis.of.
lore
1 traditional
procedure
';intuitive
á
- 65 SURVEYS
WHY WE PIED INVERTEBRATE
M. G. Morris
is takingat its startinEpoint,the idea
ThisSymposium
althoughexactly
shouldbe conserved,
s invertebrates
thatpopulationof
people.
different
to
things
different
what thisimpliesmay hean
on
conservati
the
in
surveys
of
place
the
with
ThemeIII is concerned
knowledge.
of
sort
one
with
deals
It
populations.
of invertebrate
by Dr. Broadhead.I shouldlike to take this
suchas was mentioned
G. E.
the helpI have.hadfrom.Mr.
y acknowledging
opportunitof
g
formulatin
in
today,
here
be
cannot
ely
unfortunat
who
Woodroffe,
forward.
put
Shall
I
s
someof the ideason the needfor_surveywhich
in what I have tO say,
willbe somerepetition
,
Inevitablythere
but perhapsthiswillbe no bad thing.
A gooddeal is knownaboutwhat speciesof invertebrates
occur-inthiscountry,althoughon many asoects
at leaSt).
(mesofauna
we knownext to nothing. In terms-ofspecieS
ecology,
of invertebrate
we have someideaof what we want to conserve.But it is obviously
of
t9 approachthe conservation
, well as undesirable,
impossibleas
species.
indiVidual
of
standpoint
the
from
generally
invertebrates
cases. As a
and atypical,
to thisare special,
The exceptions
briefly
must-look
we
surveys,
need
we
why
reasons
the
to
background
s
Population
te
invertebra
of
on
conservati
to,the
at someattitudes
whichare current.
e placeswhere
maintenancof
Moat of us wouldagreethatthe,
of terresanimalslive is the mosturgentneed in the conservation
It is customaryto call different
invertebrates.
trialand freshwater
thisis
,
sortsof placeswhereanima'sare found!habitats'though
of the
loosely.HoWever,"conservation
usingthe term 'habitat'
habitat"is a usefUlbasicideaeven if it is alsorathermuch of a
of
on
g
of approachinconservati
clichd. One of the advantages
by way of habitatsratherthanspediesis-thatone can
invertebrates
when
particularly
readilyintroducethe conceptof communities,
very littlework
usinghabitatin its broadsense. Unfortunately,
theyneed
s
of communitiebecause
has been doneon the conservation
cut
and alsobecausetheirgroupings
studyin themselves
intensive
We mustbe Clearthanan association
c
groupings.
across'systemati
y
not necessarilconstitute
ene sortof placedoes
of animalsfound-in
we can prea communitY.HoweVerlittlewe know abOuta community,
sortof countryside,
of a particular
supposethat it is characteristic
In
are thingswe ean all-recognise.
and 'habitats'
or 'habitat'
we can do most to protector conserveare
the-'habitats'
practice,
Odourtogetherin specialareaswhiehare,of course,
thoseWhi:ch
as
NatureReservesare important
s
-of'variousorts.
NatUreReserves
for two good reasons:-animai.habitats
areaScontaining
- 66 -
and
be expectedto containinteresting
Theymay reasonably
1.
of animals.
populations
important
but theyare
physically,
They are not onlyprotected
2.
eithermaintains
Maragement
managedto a greateror lesserextent.
themto this
converts
or
state,
desirable
themin an ecologically
of habitat
diversity
for
management
Reserves,
condition.In many
management.
object'of
important
an
typesis
MostReserveshavebeen selectedon-thebasisof theircover
Reserves,
thoughthisis not the casewith geological
of vegetation;
a familiarattitude
with which we are not concerned.This illustrates
thatif onelooksafterthe
of invertebrates,
to the conaervation
willlookafterthemselves.
invertebrates
the
flora,
and
vegetation
Cne of the dangersof thisas a principleis the senseof false
of ignorance
the maintaining
securityit gives. It encourages
theCenser,existence,
of
years
fifteen
After
aboutinvertebrates.
conserinvertebrate
to
attitude
shouldhavea morepositive
valley
if thereis nothing
vationthan this. However,the principle,
one.
better-togo on, is a reasonable
Throughthe willinghelp of RegionalOfficersand their
from
been able•o analysethe recordsof butterflies
have
I
staff,
F.N.Rs.and L.N.Rs.).It
NatureReserves(N.N.Rs.,
125 statUtory
recordsto analyse
thatwe have sufficient
is onlywithbutterflies
of specieson Reserves; thispointin itselfis
the representation
for this
are of course,atypicalinvertebrates
of note. Butterflies
partlybecause
very reason. Also, someReserveshavebeen selected
figurevery
interestin whichbutterflies
of theirentomological
fluctuations
highly. Buterflyspecieshaveundergonecountrywide
in the lastfew yearsand many of the
in numbersof greatmagnitude
recordsI havemay be out of date; thoseof certainspeciesfrom
about
the information
are. Nevertheless,
MonksWood certainly
from theseReservesmay be of interest.
butterflies
LargeBlue
Fritillary,
Only threespecies(Glanville
and LulworthSkipper)have not been foundon one or otherof these
you
of the speciesis concerned,
Reserves.As far as conservatiOn
havebeen
measures
will rememberfromMr. Hunt'stalk thatspecial
takento conservetheLargeBlue. Some otherlocalspeciesof
on our Reserves,suchas the
butterflyare verywellrepresented
Skipper. Speciesrecorded
and Chequered
BlackHalrstreak
Swallowtail,
by a few old records.
frOm onlyoneReserve,or whichare represented
AdonisMile
includethe ScotchArgus,PUrpleEmperor,HeathFritillary,
a period
7spottedSkipper. The AdonisBlue is undergoing
and Silver
is scarCe,to say the
whilethe HeathFritillary
of declinegenerally
least,in BleanWoodsin whichit was formerlyabundant(incidentally
the returnfromBleanWoodswas a 'nil'one).
- 67 -
At the otherend of the scale,the MeadowBroenhasbeen
and othercommonspeciesfrom
recordedfrom63 of theseReserves,
nearlyas many. 43 Reserveshaveno recordsat all, thoughthese
SkullSite and Wren'sNest.
includesuchplabesas Noss,SwanscoMbe
A further17 Reserveshavea knownbutterflyfaunaof 5 specieseach
poor
may havea butterflyfaunaactually
or fewer. A few of thesein speciesbut mostare just.under-recorded.
we are about
will showhow ignorant
I hope thisdigression
althoughI mustadmitthat
occuron ourReserves,
whatbutterflies
we were muchmore
beforethissurveyof the records,I-thought
ignorantthatI findthatwe arc. But if thisis the casewith
you can imaginewhat it is like for lesspopulargroups
butterflies,
Reserves.
and on lessimportant
of invertebrates,
Nature 'Reserves
Thereare two clichdswhichare applied'to
laborto describetheirfunction- "livingmuseums"and."open-air
thatalmuseumis a pladewhereone goes to see
atories".I.presume
andthatis)that the museumexistsfor the edificatiOn
Something,
amusementof peopleand not for the sakeof the exhibits.I alSo
thingsthata curatorof'a museum
presumethatone of the important
must do is to knowwhat is in his museumand to issuecatalogues.
In the base 'ofa livingmuseum,we are of course,dealihgwith an
entitieS
complexand aboveall,dynamic)collection,thc
immensely
no excuse
is
that
But
at severallevels.
of whichcan be considered
(Reserves).
is in our musetims
what,
for not beginningto catalogue
but onfY
laboratories',
NatureReservesare also 'open-air
invertebrates.
on
research
for
a few of.themare gettingused as such
tao kinds.
The researchdone in Reservescan be usefUllydividedjLnto
to find
designed
reseiu-ch
is,
that
research,
There.isconservation
conserthe
for
Reserve's
in
canbe used
out what sortsof management
populations'or
species
as
dered
vationorplantsand animals,.consi
iS basic'ecologiCal
as'theneedmay be.. And tht;re
communities
.
processes
ecbIogibal
about
researchwhichis designedto.findoui
some
be
may
There
and the flowof energywithinthe ecosystem.
rightpeopleto do these.different,
aboutwho are the.
disagreement
1 wouldregardthe
but related,kindsof research.Personally,
the.
y,
while
the-Conservanc
of
the specialconcern
formeras.being:
Universities.
by
lattermightbe.done.mainly
whiehhas
Eltonand MillerPublisheda paper,
In 1954,
ecOlogists,
animal
of
.
thinking
the
on
influence
had considerable
Quiteearlyin•theifpaper,
both withinand outsidetheConservancy.theydissurVey
the authorsdiscuss.thelecOlogicaI of animals.and
tinguishthreelevelsof survey:
limits;
habits,tolerance
lists,lifehistories,
..Species
1.
It is'thissortof
and'simple-AUantitatiVe'studies.
qualitatiVe
later:
most frequ'ently
surveyto whichI willbe referring'
- 68 -
balanceand
processes,
population
studies,
Quantitative
2.
of populations.
inter-action
Energyflowwithinthe ecosystem.
3.
It can readilybe seen thatEltonand Miller'sthree
or inventory,
roughlyto the catalogue
levelsof surveycorrespond
respecprocesses
and researchon ecological
i-esearch
conservation
between
tively.At the highestlevel,thereis littledifference
surveyand reSearch(afterall,at all levels,surveyis a typeof
connotation,
has an extensive
At thelowerlevele,'survey'
research).
one.
an intensive
'research'
Eltonand Hiller'stype 3 surveyis one of the most
kindsof research.It is
and long-drawn-out
comprehensive
difficult,
i.e.it is done overa longperiodof time. Thereare
continuing,
will do a surveyof thistype,suchas
no signsthattheConservancy
the Wythamsurveyin the near future,nor is thereany likelihood
thatone willbe doneby any otherbody. This typeof surveywill
alwaysbe veryrare.
can,
Type 2 surveysare of the sortwhich thcConservancy
been
not
have
done
been
have
that
althoughthose
and does,undertake,
survey,
3
type
the
and
type
This
as is desirable.
as comprehensive
on
are of the sortthatis best doneby staffworkingfull-time
a
only
play
to
expected
be
will
amateurs
as
working
them. Experts
minorrolein thesesurveys.
done
Type 1 surveys(specieslistsetc.)are appropriately
and .
by expertsworkingas amateurs(thisincludesprofessional
whichis not cart of their
who are doingsomething
amateurzoologists
of this sortof surveyis thc
work). The prototype
professional
J. S. (Edit.),1923 seg,The Natural
surveyof WickenFen (Gardiner,
Fen). Ar. Steelwill tellus how someof the surveys
History.ofWicken
now being donegrewout of the surveyof Spurn.Hesd.
has givenlittlethoughtto
In thepast,the Conservancy
and controlof peopletakingpart in surveysof
the organisation
is willing
the surveyor
thissort. Unlikethe generalcollector,
to conformto a definedsystemof recording.Thisbeingso, it is
positivehelp
should.give
the more importantthatthe Conservancy
to its surveyors.Somepointshere are:and plant surveysshouldprecedezoological
Geological
1.
surveysand the resultsshouldbe availableto surveyors.Both
to the
surveysand listsof plantspeciesare important
vegetation
surveyor.
of habitat
systemof recording
simpleand generalised
2.A
for the
types,basedon theplantsurvey,shouldbe made available
to conformto.
surveyors
- 69 -
of termsof distribution
An agreedsystemof definitions
3.
e.g.
shouldbe used,with the emphasison objectivity
and abundance
to "rare".
is to be'preferred
"twospecimens.only"
collecvolicher
ThereShouldbe a clearpolicyregarding
4.
tions.
I have referredto type1 surveysas beingof the catalogue
aboutthe
presentstateof.knowledge
type. In the'
or invento.ry
theyare more
thztt
fluggest
however,
of invertebrates
conservation
about
valuablethanthat,as theyprovidethe essentialinformation
vcry
of
absence
the
In
conserve.
to
like
should
we
what species
many type2 surveys,thebest thatcan be done aboutconservation
species
in manyReservesis to relateintelligently
of invertebrates
is.
thiq
am-surethat
I
species.
the
data-about
liststo biological
Conservancy.
the
of
staff
by
one of the tasksthat Shouldbe-done
realisedthatthis sort
At the sametime,it shouldbc thoroughly
inferior'alternative
very
a
is
lists
species
from
of extrapolation
surveysof communities
which
in
2,
surveyat level
to a comprehensive
wouldbe included.
aspectof surveyswhichwill'bcof increasing
An important
withinbroad
particularly
of Reserves',
is the comparison
importance
of this.
Comparisons
heaths.
systems,
sand-dune
typese.g.woods,
apparent
not
are
that
ReserVeS
between
sortmay highlight.differences
in
may.beof importance
.differences
at firstsight..Andthese
typeS.'
'ofhabitat
for.diversity
managingReserves.
but equally
in Space,is 'important
of Reserves,
CompariSon
or more so is •he comparisOn
at differenttimes at
of'Surveysdone
truewhereactive the specioslistlevel. This is particularly
the very•few
is takingplacein a Reserve.An exatple.of
management
Fen.
doneat Woodwalton
surveysof thistype is thatof Coleoptera
by Buck (1962), whichincludedthe waterbeetles. Balfour-Browne's
of thisReserve
earliersurvey(1951) of the aquaticColeoptera
he
includedrecordsdatingfrom1910-1930 and,not surprisingly,
absence
recordedmore speciesthanBuck. However,a remarkable
abundant.
a speciesrecorded.as
fromhis listwas Aciliussulcatus,
by Buck and whichnow occursin nuMbersin the Fen. The fact that
did not recordsuchan obViousspeciesis surely
Balfour-Browne
evidenceof a markedchangein the natureof the waterbodiesat
Fen.
Woodwalton
on
so far surveysof invertebrates
I haveconsidered
surveysof
NatureReserves.I am not goingto considerlarge-scale
animalsoverthe countryas a whole,althoughI shouldmention
RecordsCehtreat thisStationwhereDr. Perringis
theBiological
data for somegroupsof invertebrates
in distributional
interested
the surveyof
over the wholeof the BritishIsles. Nevertheless,
areaswhichare not NatureReservesis of someimportance.I believe
thatit is of littleuse knowingwhatwe have in Reservesif we do
- 70 -
not know the contextof theserecordsi.e.we do not knowwhat
animalsare to be foundoUtsideReserves.Fortunately,
we have,
a
good dealof:information
of thissortat the specie's
list level
in the formof countyand 'area'listsof invertebrates.
In the
future,theremay be a strongcase for surveysof areasoutside
Reservesand.other
protected
areasto see.whathas happenedto
theseareaswhichhave notbeen conserved.I imaginethislinks
in withwhatDr. Satchellhas said.
SU/VARY
Thereare threelevelsof ecological
surveyof invertebrates,eachof whichis of importance
to theConservancy.
Surveysof energyflowwithinthe ecosystem
are unlikely
to be undertaken
exceptin a very few cases.
Surveysof population
processes,
balance,interaction
of populations
are likelyto be doneby Conservancy
staffto answer
oroblemsof conservation
research.
Surveysat the specieslist and lifehistorylevelare
beingundertaken
by expertsworkingas amateurs.Not only do suéh
surveysgive theConservancy
catalogues
of thc invertebrates
in
NationalNatureReserves,
but theygive the onlyinformation,
in
most cases,fromwhichmanagement
can be plannedand done in the
Reserves.Such surveyscan alsogive information
on the relative
richnessof Reservesof similartypeand on thepossiblediversificationof habitats.
71
ORGANIZATION
ANDFRACTICEOF
iSIVESURVEYS
W. O. Steel
As is now perhapswell known,theShellCompanyhavemade
available
to theNatureConservancY,.an
annualsum of moneyto be
used for entomological
surveysand my talkdealswithproblems
associated
with the planningof these'.Some of thepointsdealtwith
are applicable
to entomological
surveysin generalbut the main
concernis with intensive
surveysof the typedone on Rhumbetween
1960and 1963. Thesesurveysare undertaken
by entomologists
who
are not on the Conservancy's
staffworkingfor a fortnight
or so
annuallyovera nuMberof yearsin a Reserveout of easyreachof
theirhomes'orlaboratories.
The objectof suchsurveysis simply
to findout what is thereand whereit occursi.e. to providean
inventory
of the insectspresenttogether
with as much information
on theirbiology,distribution
and abundance
as can be colleeted
in the timeavailable..
For Obviousreasons,no quantitative
data
can be givenand the surveysdannotbe considered
in anyway ecological..Theymay well howeverserveas bases for futureecological
work in the areas:concurned.
.Havingdefined.the
aims of suchsurveys,it mustbe
decidedhow theymay.best
be done and'heretwo mainmethodsmust
be considered.The first,whichmay conveniently
be referredto
as the 'bulk'methodConsistsof collecting
aS manyinsectsas
possibleusingall available
means,i.e.beating,sweeping,
trappingt,
soilextractions
etc.'The materialcollected
is killedand simply
bulkedin tubesor papersWithoutany examination
or selection,
exceptperhapsfor size to make-packing
easier. ThiSimethod
is
widelyusedby expeditions
to the.Tropics
and efsewhere.
It is in
fact,the'onlymethod
.whichcan be used in SuehplaCeswherethe fauna
is verypoorlyknOwnand it would'betime-consuming and pointlessto
try and'identify
the insectsOh thespot. Also in suchexpeditions,
as much.aspossiblemustbe collected
in.a Shorttimeand the nuMber
of collectors
is limited.The advantages
of:themethod are thatmore
or lessanyonewho can recognise
insectscan be employed.for
collect-.
ing; all ordersof inSectsare covered'and
no greatdemandsare made
on the Surveyheadquarters,
'a tentbeing sufficient...No
literature
or equipment,
etherthan.collecting
equipment,
iS necessary.The
disadvantages
.arethatmuchunnecessary
materialis collected,
some
speciescanbc overlooked
becausetheyare not specially
searched
for, thereis a dangerOf destroying
habitatsby the collecting
methods,and a greatdealof timehas to be spentaftcrthe'survey
in sortingand mountingthe materialand gettingit identified.
- 72 whichherbcan wellWealled
Usingthe secondmethod,
by specialist'sYeach
are collected
insects
method,
the 'selective'
are sortedand
group. The specimens
workingon his ownparticular
present
and thevariousspecies
at theSurveyheadquarters
identified
becomeknown. Thismeansthatas timegoesby, moreand
gradually
thenuMber
therefore
in thefield.and
morespeciescanbe identified
smaller.The
becomesprogressively
collected
of speciesactually
few
relatively
methodare that
of this'selective'
advantages
beingoveraretaken andthe chancescf.anyspecies
specimens
beenshownthat
lookedare veryMuchless- it hasrepeatedly
morespecies
groupproduces
Of a particular
colleeting
specialized
which
Also,any species
collecting.
of thisgroupthandoesgeneral
pointof view
or distributional
fromthebiological
are of interest
foror studied.The
searched
andcanbe specially
are recognized
mustbe employed
of themethodare thatspecialists
disadvantages
for the surveyso thatonly-apartof the insectfaunawillnormally
mustbe madeat theSurVeYheadquarters
be dealtwith,andprovision
workusingmicroscopes.
for identifiCation
GreatBritain
the insectfauna-of
Withcertainexceptions,
typeto be
of the:selective'
is wellenoughknownfor surveys
(of theNatureConservancy's
sub-committee
and the-survey
undertaken
believesthatthismethodshould
lIdaiSon.Committee)
Entomological
Thiswill,it is hoped,helpto guard
pOssible,
be usedwhenever
ia probably
andaboveall,against-what
againstovercollecting
or
thedestrUction
surveys,
dangerin entomological
thegreatest
as uscdheremeanstheparticular
of habitats.'Habitat'
alteration
or groupsof insects.Not
of variousinsects
environments
breeding
but some
activitieS,
by collecting
all of theseare easilydamaged
or mosson
patchesof vegetation
suchas deadtrees,isolated
in character
altered
or considerably
topscanbe destroyed
mountain
for some
makeit difficult
This.may
collecting.
throughthoughtless
if it is a smallone.
particularly
in theReserve,
speciesto survive
carriedout
beensatisfactorily
havealready
surveys
'Selective'
to Rhum,
in addition
at SpurnHead,MalhamTarnand Inverpolly,
use to be madeof the time
possible
and theyenablethegreatest
canbe doneduringtheday and identification
- collecting
available
or in bad weather.
and otherindoorworkin theevenings
outof
to thecarrying
themainobstacle
At thepresent,
personnel
in finding:
but the difficulty
surveysis not lackof.money
NatureConservancy
workof the typerequired Because'the
to undertake
it doesnot follow
areasurveyed,
in havinga.certain
is interested
entomologically,
thattheareais of anygreatinterest
necessarily
Maybe yery small.
species
i.e.the numberof 'rare'or 'local'
cellectors
primarily
ar.;
manyentomologists
On theother.hand,
as possible.
of as manyspecies
is the acquisition
whosemaininterest
whichare new to them
onlyspecies
andperhapsrecord,
Theycollect,
or of whichtheyhav6not a fullseriesand ignorethe rest. This
information
to obtainaCcurate
why it is oftenso difficult
explains
- 73 -
s obviously
of commoninsects.Such entomologistare
on the distribution
for surveywork,whichrequirespeoplewho,besidesbeing
unsuitable
mustbe preparedto spend
reliablein theiridentifications)
seeingany speciesof
withoutneeessarily
severalweekscollecting
is desirable,
n
interestto them. Anotherqualificatiowhich
particular
stages
larval
the
of
is a knowledge
thoughperhapsnot essential,
arc
areas
survey
the
As
of the insectson whichtheyare working.
the
of
case
the
in
weeks,
normellyvisitedfor someeightto ten
thereis no
Rhum sUrveybetweenearlyJune and earlySepteMber,
adult's-Laze
tho
in
be
will
present
thatall the insccts
guarantee
therefore.
may
larvae
identify
ability'to
The
duringthe survey.
of the
idea
an
gives
also
It
of thelist.
add to the Completeness
The
breeding.
are
species
the'varieus
which
typeof habitat.in
s make somepeoplefeel
of suchentoMologistmay
pparentshortage
people
less experienced
ng
for'recruiti
case
a.good
that thereis
however,
done
is
this
If
collecting.
in
and beinglessselective
anotherproblemwill have to be faced,thatof getting.thematerial
as obtaining
identified.Thisin itself,can be justas difficult
survey.
the
to undertake
specialists
in the area
available
n facilities
The accommodatioand
can be
survey
a
before
considered
be
course,
of
must
concerned
reffioter
the
of
some
near
or
n
in
accommodatio
arranged.Sleeping
is verylimitedand unlessit can be
Reserves,suchas Inverpolly,
peoplethatcan take
the nu.Mber-Of
,
augmentedconsiderablylimits
theConservancy could
perhaps
that
here
is.made
n
Thesuggestio
part.
Reservesas
n
in.such
accommodatio
the
increase
to
caravan
a
dbtain
wpm
n,
accommodatio
sleeping
to
addition
In
and when required.
can be used. This
wheremicroscopes
is necessary
formof 'workreom'
supplyis essential.
but-anelectricity
neednotbe veryelaborate
caravans
Althougha surveywas carriedout in the Ainsdaleareaiusing
a greatdeal of timewas wastedbecauseof
withoutelectricity,
couldbe providedby the Conserlighting.Nicroscopes
inadequate
for the surveypartyto
vancybut it is perhapsmorepracticable
providetheirown. Thereis no dangerof losshere as insurance
can
policieswhichcoverall the riskslikelyto be encountered,
be obtainedverycheaply.
shouldnot have
It is desirablethattheporty themselves
of
on Rhum Showedthatthe preparation
to cookmeals. Experience
be used
mealstakesup a greatdeal of timewhichcould_otherwise
'Thesaving
more in fact,thanmightbe e3COected.
for collecting,
Wereprovided
of timeon the lastvisitto _theislandwhen all.meal's
staff,was verynoticeable.
by the Conservancy
to save timein
is also desirable
Some formof transport
In
Reserves.
in largo
gettingfromplaceto place,particularly
has
Reserve
providedfor the staffof the
the past,the transport
has provedto be
and thisarrangement
been usedwhenavailable
satiSfactory.
- 74-
Much has been saidhereregardingthe need for specialists
'for collecting
and identification
and it maYbe of interestto non- .
entomologists
to-havea.ronghlidea
of the nuPberof speciesinvolved
and the takonoMic
stateof the varionsInsectOrders. Thisis
shown.inthe accompanying.tablea
and givesan.ideaof the identifi,c'atiOn
probieMs
involved.It mustbe eMphasized
thatalthoughinmany casesit is statedthatadequate
keys are available,a'certain
amountof experiencesometime
,
å large
s
amount,
isnecessary
before
thesecan be uaed satisfaCtorily.
Even then,namedspecimens
may
be neededfor comparison
beforea correctidentification
can be
obtained.Onlyrarelycan a personwith littleor no knowledge
of
the subjectsipplyread througha key and identifyspecies.
Table I
ApproximattotaI:
e
20,000
- 75 Table II
Taxonomic Status pf the Er itiO?
of ites
1.1s.ect--FaTI,J9.259:1414ing..paras
soil living species requiring specialized
Apterygote Orders
-Collecting methods: Not . many workers; literature more
or-less adeqUate, largely published abroad but. in English.
Ephemeroptera, Odonata,.Plecoptera - adee4ate keys available to
adults and larvae.
Orthoptera, DerMaptera, Dictyoptera - adequate keys to 'adults but
not to larvae.
Psocoptera - very few.workers. Prench work available on adults
and R.Ent.Soc.Handbook in preparation; little published
on larvae.
Hemiptera:.
era
good keysavailableto adults; no comprehenReteroptsivekey to larvaebut at leastone personworking
on these.
- very few workers;
(leafhoppers)
Auchenorhyncha
Homoptera
thirdand final
available,
two R.Ent.Soc.Handbooks
one in preparation.Littleknownaboutlarvae.
whiteflies)
(aphids,scaleinsects,
Sterorhyncha
Homoptera
to some
available
keys
but
keys,
ive
- no comprehens
larvae.
about
known
Little
groups.
- very few workers,partialkey available.R.Ent.Soc.
Thysanoptera
Handbookin preparation.
to larvaeand adults.
- keys available
,
NeuropteraMecoptera
on larvae. Britishkeys
Trichopter-a severalworkers,particularly
l
to adultsout of datebut morerecentcontinentakeys
paperson larvaewith
available.A nuMberof scattered
somekeys.
on micros.
Lepidopter-a adultmacroswell known; work necessary
Larvaenot well knownand muchwork needed,some(continental)keysavailable.
- adequatekeys availableto adultsof mostfamilies
Coleoptera
and to a few larvae. Interestin larvaeon the increase.
Hymenoptera:
at least
Symphyta- adultscoveredby R.Ent.Soc.Handbooks,
larvae.
to
key
ive
corprehens
one fairly
- a numberof peoplenow workingon thesebut
Parasitica
a greatdeal stillto be done. At presentonlya
of some familiescanbe identified
smallpercentage
work is necessary.
and majorrevisional
well knownand keysavailable.
Aculeata- reasonably
othersmuch
Diptera- severalworkers,somefamilieswell knovrnbut
less so and requiringfurtherwork.One family,Cecidomyiidae
verypoorlyworked.
- 76 -
Summaryof Discussion
followi a ersb Dr. MorrisandMr.Steel
priorto surveywas
analysis
of habitat
The question
thevalueof thiswhile
emphasised
meMbers
raised.SomeSymposium
whichthe surveywas
information
thatit assumed
othersthoUght
of
to gather.It waspointedout thattherecording
designed
duringa surveYcouldresultin a poorer
on habitats
information
thatthe
It was alsoemphasised
listbeing'obtained.
species
habitats
are foundarenot theircomplete
placeswhereanimals
in this
was mentioned
of larvalexamination
and theimportance
of one species
how manyspecimens
context.The valueof knowing
was alsomentioned.
itshabitat
to determine
mustbe dbserved
to be givento
importance
It was agreedthattherelative
on the time
typesof surveydepended
and ecological
theinventory
werebeingdone.
whilethe surveys
available
and specialists
- 77 -
SOMEFEA
• D.
INTEREST
OF CONSERVATION
ARISINGFROM SURVEYS
A. M. Massee
Introduction
Englandconsistof a
in South-East
Many of the woodlands
sweetchestnutia.the
of trees,of whichthe introduced
variety,
predominant species. Othertreescommonlygrowingin these
maple,hornbeam
hawthorn,
are oak,hazel,birch,.aspen,
woodlands
elderand crab apple.dFormerlY,.
rn,
,
and lesscommonlywhitethd
but the adventof the
oak formedthe canopyin thesewoodlands,
theihortageof tirber;resultedin
SecondWorldWar combined.with
most of the oaksbeing-felled.
are grownfor periodic
The treesin-thesewoodlands
dividedintocants,and once
The woods:are
cuttingor.coppicing..
Thereis a
in everytwelveyearsor so; eachCant.iS alear-felled.
chestnut
sweet
steadydemandfor the 'batt'and spilesmadefrom
s
k
trees. The largechestnut'bats'form the main framewor'support
The
today.
in hop gardens
e
erectedin wireworkstructurused
smallerchestnut'bats'are splitand made intospilesand are UsCd-.
and housingestates.The
properties
for fencing agricultural
floraof the newlycoppicedpart of the woodlandmay be very rich;
wood anemone and dogs mercury
primrose,
,
s
suchplants_abluebell
the cuckooflower,wild
may carpetthe floorand in addition,
7wheat,goldenrod and fOxgloveoftengrow freely
straWberry,.cow
thevigorousnew growthsof the
for two or_three_years-until
trees'housein' the groundflora,which
and other.
sweetchestnut
out.
gradually'dies
Two exarplesof woodlandsthatredeivethe coppice
t theNationalNatureReservesof Ham StreetWoods
treatmenare
and BlearWoods,Kent.
aISo supporta largenuMberof
Thesecoppicedwoodlands
nowbe discussed
of-which.will
examples
orderi,
insectsof various
in somedetail.
The Conservation of ColeopteraHerd tera-Hetertera Looidotera
underco ice manaement.
and otherinsectsin woodlands
A veryrich insectfaunacan be maintained in coppiced
acreageis cut downeach
s
woodland everyyearprovideda reasonable
will ensurethat treesof all ages
winter. This typeof management
- 78
-
to providethe
fromone to twelveyears,will alwaysbe available
kind of
this
with
associated
-insects
needsof a wide range.of
aercage
small
a
leave
to
it is essential
habitat.In addition,
associated'
species
the
that
so
cutting.,
without
of treesto mature
for.
catered
with the oldertreeswillbe
AspenCoppice
Large areas of young aspen can readily be established
when the tree is under coppice management. The metallic, green
weevil Byctiscus populi (L.) prefers the young saplings of from
one to four years old, and these should always be available to
supply the needs of this weevil. The female rolls the edge of the
leaf inwards into the shape of a cigarette, and in this dwelling,
the eggs are laid. The various stages of the larvae are completed
within the rolled leaves, and later on, the adults emerge and feed
on the foliage. In rare cases, two or more leaves may be rolled
together by the weevil, but this is exceptional. Rutidosoma
globulus (Herbst.) is another very local weevil that feeds on aspen
leaves of this age group, althoUgh it occasionally occurs on willow
and poplar also. Aspen from five to ten years old meets the requirements of the three species of the genus Zeugophora (Chrysomelidae)
and the larva of the Sallow Kitten Harpyia furcula Clerck.
Chrvsomela tregulae T. and its larvaare morepartialto the young
leavestowardsthe tipsof the shoots. The Miridplantbugs,
(Fallen)
Fieberand Neomecommabilineatus
Bracharthrumlimitatum
aspensprovide
inhabitaspenof all ages. The oldercatkin-bearing
and the gallhabitatfor weevilsof the genusDorytomus
a suitable
forminglonghornbeetle, Saperdapopulnea(L.). Luperusflavipes
s oldertrees.
also-inhabitthe
(L.) (Chrysomelidae)
HazelCoppice
Hazel is the hostplantof a wide rangeof insects,
to the age groupof the trees,in
and the faunavariesaccording
The youngbushesand thoseup
birch.
and
aspen
to
a mannersimilar
greenweevil,Byctiscus
metallic
the
by
selected
are
years
five
to
betulae(L.)and thebrightblue variety,c.1prini,IS (Sy.). The
of this speciesfollowsthe samepatternas B. populi
life-cycle
beetles,Cryptocepholus
on aspen. Threevery localChrysomelid
(L.) are also
(L.)and C. bipunctatus
coryli(L.),C. sexpunctatus
and mate.
feed
adults
the
which
upon
bushes,
young
with
associated
hazel
older
much
to
off
move
females
the
oviposit,
to
ready
When
plants,mossesand the unicellular
bushesthatsupportepiphytic
larvae
sp. upon which the casebeoring
greenalga,Plcurococcus
and tracesof
feed. The case is formedof cast skins,excrement
hazelbark. The bushesin the age group fiveto ten yearsattract
rod weevil,Apoderuscoryli(L.)and severalspecies
the conspicuous
Phyluscoryli(L.) and Psallusperrisi
of Hiridbugs including
—79 —
••
with oak.
(MllsantandRey),the latterbeingnormallyassociated
Palomena
bug,
id
Pentatom
the
attract
bushes
earing
catkin-b
older
The
y
partiall
The
bushes.
fruiting
to
confined
is
which
(L.)
prasina
ly
especial
is
(Panzer)
ns
chloriza
ris
Malacoco
bug,
Mirid
us
predacio
its
of
search
in
plants
other
to
move
will
but
hazel,
to
attached
speciesthatlaysits eggs on the shoots
prey. This is an interesting
theyare neverinsertedinto the tissue
and
buds
the
of
axil
near the
of the Shoots,as is the habitof most speciesof laridbugs. The
occurson hazelof all ages,
e
Eriophyidmite,Ph to tus avellanaNal.
theBritishIsles. The microscopic
and it is abundantthroughout
and
swellconsiderably
mitesfeedwithinthebuds,whicheventually
are commonlyreferredto ds 'BigBuds'. The mitesalsocausea
of the femaleflowersand preventthe fruitssetting.
malformation
BirchCoppice
birch,and
Many speciesofinsectsare associated.with
bushes
provided
s
woodland
coppice
in
ained
be.maint
most of them.can
insects.
ous
herbivor
the
for
food
as
e
availabl
are
ages
of various
unctier Payk.is one of the
Cr toce-Dhalus
be'atle,
The Chrysomelid
of England.
the south,east
in
noted
insects
birch
ing
most interest
on a few
d
persiste
has
that
species
uncommon
and
local
It is a very
last'
the
for
Kent
Wood,
Darenth
of
corner
one
in
trees
birch
young
the
ed•over
sca.tter
trees
t
commones
the
of
one
is
Birch
fiftyyears.
itself
confines
beetle
this
yet
and
wood,
the
of
acres
of
hundreds
its range;.Theadult
to'extend
seems
to one smallarea;andnever
has not been foundon
and
bushes
young
very
of
age
the'foli
.
feedson
olderbushesin the vicinity;In orderto havea nuMberof bushesin
for thebeetleto feedUpon,a numberof bushes
a suitablecondition
seasonstoensure a constantSupplyof young
downmost
cut
been
have
growths. Is it too much to hope that•thiS implemethodof conser
in the same
for thisbeetleremaining
vationhas partlyaccounted
localityfor-somanyyears?: As with otherspeciesof the genus,
c
ng
to veryold birches'supportiepipheti
C. punctigerdisperses
larvafeeds
etc.,on thebark wherethiscase-bearing
plants,-moSseS,
When fUllyfed,the larvamoveson to grassesunderthe treesand
pupationtakesplaceon the grassstems. 'Theblackweevil,Magdalis
in a
its larva.feeds
is fcundon-oldertrees,jand
(Is.)
carbonaria
underthe bark. Two otherspeciesof weevil,
smallcell-juat
Curculiobetulae(Steph.)and C. rubidus(Gyll.)are specifie
the.tops
birch. The fermerusuallyfeedson the foliagegrowing'at
more catholicand may be
of the treee,and the latterspecies--is
.(Chrysomelidae)
tridentata
foundanywhereon the tree. Labidostomus
beetlewithblue head,
buff-coloured
is a verylocaland uncommon
thoraxand logs. Ii occursonly on largetrees,and thc hairycaseplantsand other.foreign
bearinglarvaefeedon the epiphytic
with old.trees.At East Malling,the species'
growthsaSsociated
yearsor more,and it is neverfoundon,
of twenty-five
selectstrees
betuleti(Fallen).
g
youngbushes. SeveralMiridbugs includinPsallus
up
s
build
sometime
and
and P. falleniReuterare specificto birch
- 80 bug, Elasmostethus
largepopulations.Two speciesof Pentatomid
grisea(L.) are confinedto fruiting
(L.)and Elasmucha
interstinctus
bushes. They deposittheireggs on the foliageand the gregarious
immaturestagesfeedon the femalecatkins.Both specieshibernate
in the adultstate,but are rarelynotedduringthu dormantseason.
ChestnutCoppice
devoidof
sweetchestnutis strikingly
The introduced
insectsbut the verycommonweevil,Stro hosomusmelanogrammus
abundanton the foliage,uponwhichit feeds.
(Forst.)is sometimes
in
plentifUl
The woedLeopardMoth,ZeuzerapyrinaL. is sometimes
and the larvais familiar
the stemsof the-purestandsof chestnut,
who comeacrossit in the courseof theirwork. The
to woodcutters
chiefinterestof chestnutcoppicecentresaroundthe two or three
the fellingof the wood. Duringthis'
following
yearsimmediately
period,the insectfaunais usuallyvery rich. For example,three
with newlycoppicedareasof
verylocalplantbugs are associated
manyKentishwoodlands.The red and blackShieldbug, Eurydema
to Kent,
restricted
,
largely
(Scop.)(Pentatomidae)
dominulus
is confinedto the cut-downareasof chestnutin whichthe cuckoo
flowergrows. The adultfeedson the leavesof the cuckooflower
in April and May and lays its eggson the stemsof the plant. The
progencyalso feedon the leavesand on reachingthe adultstage,they
in thickmoss. The speciesis
and laterhibernate
firstaestivate
gregarious;as many as fortyor morehuddletogetherin a small
area of moss. In April,theyseekothernewlycut-downareasof
AO
the cuckooflowergrows. In one smallexperiment,
woodlandwhereadultswere markedwith yellowpaintand replacedin
overwintering
on Cuckoo
were recovered
May, 8 specimens
the moss. The following
flowerssomeeightyyardsaway.
Fieberprovidesanother
The flatbug,Aradusaterrimus
part of a
from.ono
very goodexampleof an insectthatdisperses
to enableitto survive.
conditions
wood to anotherto find.suitable
The Aradidwas firstrecordedin Englandin 1865when two specimens
were found,one in the centreof Norwichand the otherat Darenth
Wood. In 1930,the specieswas recordedagain,this timein old
wood chips. Since1930,the specieshas
sackingwhichcontained
at East Mailingand elsewhereand alwaysin
been foundmany-times
by
of chestnutchippingwhichare infested
with•piles
association
making
When
are madeby woodcutters
fungi. The chestnutchippings
are left to rot.when the
spiles. The chippings
up and sharpening
are foundin the funguseggs
and
larvae
wood is cleared. Adults,
s, bugs
the year..After-two-yearthe
infestedchippingsthroughout
feed
they
which
upon
fungus
the
IroM the chippings
beCauCe
disperse
most
a
is
It
fungi.
and
is crowdedout by otherspeciesof mould
.
of thebug swarming
to Observeaggregations
fascinating:experience
behind
goes
sun
the
When
August.
in
day
warm
a
on
on the chippings
- 81 -
a-cloUdor at-d9isk3.the
bugs dispersein thepile of chippings
and
breeduntilthe pilebeconesunsuitable.for
theirsurvival.
'TheStenocephalid
bug, Dicranoce
halusmediusMasant.
and Rey) is a verylocalbug thatoccursin coppiced
wodd wherewood
spurge-grows.
This veryagilebug-IaYsitS eggsin the stemsof the
plant,whilethe adultsfeedon the fruits. This speciesis also'
partlypredacious
andruns abait-rapidlYin
searchof its prey. The
bug moveson as the wood growsup and wood spurgedies down. -The
lacebug, Oncochilasimplex(H.-S.)is alsoassociated
with this
plant,but the bug onlyoccursin the cut-downareasof the Wood.
Unusualinsecthåbitats
It is the customof woodcutters
to burn all surpluswood
and cheshutbranchesnot suitablefor makingsoiles,sinceit is part
of theircontractnot to leaveany wastewood aboutwhen the coppicing
has been completed.Followingthe firesin thewoods,the areasof
burntgroundprovideidealsituations
for certaininsectsto breed
in. th fact,two minute.and
veryuncommonbeetlesrequirethiskind
of habitat.The,Histerid
beetle,AcritushomoeathicusWoll.and
the Staphylinid,
Microe lus tesserula
Curt,occuramongsttheburnt
embers,and are attachedto the reddishfungusPvronemaconflueris
(Discomycetes)
whichoftenoccursin suchconditions.Thesetwo
spediesbreedamongstthe eMbersof burntgroundfor two seasons
and thendisperseto othernewlycut downpartsof the wood.
Stool-oak
. Althoughmany of the oakS KentishWoodlands
havebeen
clearfelled,.
a few stumpsusually
YreMainthatare one or two feet
high. Thesestool-oaks
nroduceveryvigorousgrOwthswhichare
oftensubjectto severeattacksof mildew. The mildewedleaves
and Shootsprovidethe foodneededby the larVaeof the localmoth,
theLunarDoubleStripe,MinucialunarisSchiff.,a soeciosmuch sought
afterby Lepidopterists.
It is fortunate
the mildewedshoots,which
seem to be essentialfor the rearingof the larva,also'occur
on the
pollarded
oaksgrowingin the hedgerows,
becausethe.stool-oaks,
onceverynumerousin theForestry
Comndssion's
propertyat Ham Street,
are rapidlydyingbecauseof continual
cutting;
The stool-oak
is also the habitatof the red weevilwith
blacklegs,Attelabusnitens(Scop.),a soeciesrarelyfoundon 'mature
oak althoughit will surviveon 'scrub'
oak.
The i, ortanceof i - rowi woodlands
A veryrich faunais oftenassociated
with ivy growing.on
trees,walls,fencesetc.,and thisplant shouldalwaysbe encouraged
whenpossible.For example,very old ivy withblack stemsprovides
the habitatfor the verylocalPlatystoudd
beetle,ChoragusSheppardi.
Kirby,a shinyblack speciesthatpossessesthe powerof leaping.
- 82 The thickstemsof old ivy providethe habitatsuitablefor the
Scolytid
beetle,Kissohagushederae(Schgitt),
a speciesresident
in BleanWoodsNatureReserve. The Anobiidbeetles,Hebodiais erialis
(L.) and Ochinaptinoides(Marsh.)are alsoassociated
withwellestablished
ivy; The minute.ladybird
beetleClithostethus
arctuatus
(Rossi)may occasionally
be foundon ivy leavesin Octoberwhereit
feedson the aphidsassociated
with thisplant. The weevil,
Lio hloeustessulatus
(NEU.) and the varietymaurus(Rhrsh.)
both
occurwhere the ivy growsthicklyon treesandwalls. The beautiful
and delicatelacebug, Dere h sia foliacea(Fallen)is another
inhabitant
of thishost,and thebug feedson the newlyformedleaves
whilethe Homopteron,
Issuscoleotratus(Geoff.)selectsthe leaves
of a moreleatherynature.
Summaryof Discussion
It was askedhow insectsat the topsof treeswere dhserved.
ObservatiOn
aftergales,ropingdown of treesand cuttingthemdown
were mentioned.
The detailedinformation
givenby Dr. Masseefor named
specieswas particularly
valuedand it was suggested
that specific
obServational
evidenceof this sort.mas
important
in conservation
to bridgethegapSUntildetailedexperiMental
workwas available
for
a betterbasis for_management.
-The-difficulties
of identification
of specimens
and retaining and storagc.of
materialwere discussed
in detail,particularly
in relationto standardmuseumpractice.The importance
of having
materialreliablydetermined
was stressed
and it was the view of
severalSymposium
membersthatmore taxonomists
shouldbe recruited
into theConservancy.The suitability
of taxonomic
work as 'parttime'researchfor conservation
branchstaffwas mentioned.
- 83 -
COLLECTIONS
THE-ROLFOF INVERTEBRATE
I TEE CONSERVANCY
DN
Editedsummaryof talkby
T. Huxley
y formedsixteen
From timeto timosincetheConservancwas
problemsof invertebrate
yearsago, staffmeetingshaveconsidered
to
been.limit
s
discussionhave
Generally,
conservation.
. , ed
fauna
invertebrate
aspectse.g.biological-recordS'i
particular
plans,a chartof animalhabitatsfor Reserves,
in management
sections
s to
s surveys.Onlyrarelyhave suchdiscussionled
collectionand
nd
problems.a
te
y copingwith invertebra
an aoceptedmethodologfor
y
policy
Conservanc
Nature
conclude.that.no
it wouldnot be unjust.to
has y6t emerged.
conservation
for invettebrate
LiaisonCommittee
of theEntomological
The deliberations
1958,it
reflectthissituation.Sinceits firstmeetingin :Larch
the
notice
y's
has donegood servicein bringingto the Conservanc
,
nservation
require.co
invertebrates
namesof placeswhereparticular
problems.
hoc
ad
recurringalsoin clarifying
and it has been invaluable
howeverhave seldomfounda permanent.answer.
l
Fundamentaquestions
In'thisrespect,it seemsthat theCommitteehas failed,partly
of the Conservancy's
becauseof the limitedexperience
perhaps,
, tendto avoid
It may be thatas scientistswe
representatives.
is not our individual
matter
Whose
givingPositiveanswersto qUestions
- 9th.meeting
the
that-until
observe
concern,if so, it is pertinentto•
on.
full-time
employed
of-staff
no member
n
of the'LiaisoCommittee,
work hadbeen present.
invertebrate
is that thereare !.
aboutinvertebrates
The main-difficulty
sizesuggesting.
average
small
so many of themand that,deSpitetheir
thatof
'as
comlicated
as
is often
autecology
,
simplicitytheir
part-timer
a
for
difficult
very
is
it
ly,
vertebrates.Consequent
to makemuch of a
informedaboutinvertebrates
to bo sufficiently
Conservancy.The
the
e
for
they.creat
to theproblems
contribution
so immersedin
often
is
fUll-timer
the
reversemay alsobe true;
with wider
himself
concern
to
time
spare
hiS work thathe has no
issuet.
thatwo employmorepeoplewhose
therefore,
It it essential
theywouldneedadditional
but
gap,
this
bridge
to
is
sk
special:ta
employment;for instancethojob of
Werk to justifyfull-time
collections.Thesehavo
te
y
loekingafterConservancinvertebra
t extra
a requiremenfor
and
curating
in
a.problem
eated
already:cr
curaterswillof necessitybehandlingall kinds
staff.—Because-our
84
of invertebrates and getting to know the appropriate specialists
and literature, they will be in a specially advantageous position to
assist in dealing with many of the currently vexing questions which
arise from the need for e.g. inventory lists and voucher specimens.
The creation of an invertebrate service, based on collections is a
natural development of this idea, giving a service sirdlar to the
other basic services •n the Conservancy. Such a service would not
of course, remove all obligations on Conservation Branch staff to
think or take action about invertebrates, any more than the
existence
of thobiometrics,
geology,climateor libraryservicesfor example,.
have done in theirrespective
fields.
Becausethisideaof an invertebrate
servicehas recurred
to me in variousformsduringthe last tao years,I used ThemeIV
as an opportunity
for advocating
it in public. ThemeIV was designed
to coverthe role of thoConservancy's
invertebrate
collections,
the
problemof invertebrate
recordsgenerally
(andespecially
whether
theReserverecordis a suitablemethodfor storingthisinformation)
and tho needfor voucherspecimens.In this summaryof my contribution,I have not repeatedall the information
presented
by me at
the SymposiuM.and
the following
factsconcerning
theConservancy's
collections
must suffice.
Thereare fourmainConservancy
collections
and foursmall
ones (seeTable1)9 containing
aboutas many specimens
- at an estimate
of four specimens
per species- as the 26,000volumesholdin the
Conservancy's
libraries.SOMQ of the collections
pre adequately
lookedafter,othersare not; onlya few are catalogued;noneis
housedin permanentspecially
built quartersand only one station
(Edinburgh).
is doingsomething
aboutthis. The collections
havebeen
acquiredin severalways; by bequests,
requestsand collecting
by
NatureConservancy
staff. hest havebeen formedas aids to identificationand althoughvouchercollections
alsodo this,onlyone of
the largercollections
has the holdingof vouchermaterialas'a
main aim. Storageequipmont
has cost the Conservancy
about£900.
'Theexistence
of thesecollections
establishes
that
collections
of somekind are necessary
for the properfunctioning
of
the Conservancy.This conclusion
oill stillbe trueeven if some
partsof collections
are holdonly temporarily
by theConservancy
becausetemporary
collections
will alwaysremaina smallproportion
of the totalstockheld at any one time. Furthermore,
justas
permanentConservancy
reference
collections
areboundto enlarge,
theConservancy
Cannot'avoid
the additional
taskof maintaining
some
sortof minimuMvouchercollections.
Alternative
procedures
to holding actualvoucherspecimens
for ensuringthe validityof National
NatureReserverecordshavebeen suggestedand foundwanting; some
kind of vouchersystemis inevitable
and althoughsomeConservancy
stationsmay be able to make arrangements
with a localmuseumwhich
- 85 -
will guarantee to hold their voucher material in perpetuity, others
will not be so fortunate and will have to accept a heavy curating
burden.
At this point in my talk, I enlarged on the subject of
Conservancy servicing sections in-general. Although the initial
substance of servicing operations is often things e.g. books, maps
or files,, they also involve people, place, time and cxpertise. In
addition to these five essential elements, there is another factor
- extremely imnortant in a democratic orgarization - a 'comrunal
acceptance that servicing operations are essential and should be
made to work wall. The Conservancy has sometimes prided itself that
it has not been sUbject to Fnrkinson's Law but there is a danger in
taking this Law too seriously; without basic intcrnal services,
the pcople.who are-directly Servicing
the operational staff, i.e.
the public,_cannot cope. Fortunately - or unfortunately - according .
to one's point of view, the outwardly servicing Conservancy staff
have become adept at making superficial knowledge go a.long way,
In respect of invertebrates, organized public pressure has not yet
become great enough to show up our basic ignorance.
An important point about a good internal service is that
it is generally able to deal with more than its original task.
Libraries, for example, not only house books but also deal with printing,
publications and press.cuttings. Az an example of another basic .
service which works well, I described at the SymposiuM the Edinburgh
Office Registry, and as a measure-of, the-communal importance attached
to it, I pointed out that scientists as well as •administrators had
collaborated in its design. A similsr collaboration, with scientists .
taking the lead, is needed-to overcome the Conservancy's organizational
difficulties concerning invertebrates.
With regard to an invertebrate collections service, I
suggested that eventually there might be five reference collections
at Furzebrook, Monks Wood, Merleweod, Bangor and Edinburgh - with a
curator in charge of each and clerical assistance guaranteed„--Each
collection should be housed in a special room with working space for
two persons, the additional space being required forrvisiting
taxonomists (for no curator could be expected to identify everything).
Thc final nuMbor of reference collectiOns is not important and in
to imply that the collections
recommending five, I meqnt simply
service must be deployed in several centres in Great Britain as it
is essential that curators not only look after speciMens collected
from Nature Reserves, but also actually visit and themselvcs.collect
in some of thc Nature Reserves in the group of regions which they.
service.
Scientific
Officersas well as Experimental
Officers
shouldbe involved
in the running, of the service.The situation
Shouldbe avoidedof an Experimental
Officerstuckawayin a corner
with Someinsects,responsible
to a Principal
Scientific
Officer
or a SeniorPrincipal
Scientific
Officerwho-never
gets enoughtime
to help in and guidethe work. Indeed,as sciuntists,
the curators
Shouldhavecloselinks,or perhapsevenbe directly
responsible
to the head of theConservation.Research Section,ratherthanto
theDirectorof the Stationin whichtheyare located.
The dutiesof the c011ections
servicewouldrelateto the
wholefieldof invertebrate
surveyon Reserves,
adviceon permits
-tocolleet,invertebrate
data for management
plansand invertebrate
nomenclature. Eventually,
all listsof speciesnow scattered
through
filesandReserverecords-houldbe collated
by the appropriate
coliebtions
serviceand in thispart of theirtask,the curators
wouldwork in closecollaboration
with theBiological
RecordsCentre.
Thisidea of a deployedinvertebrate
(collections)
service
was the main thoughtthatI wantedto get acrossat the Symposium.
Unfortunately, both in my paperand in the discussion
which.followed,
it got ratherlost in theproblemof.National
NatureReservevoucher
speciMens.Thosecertainly
createan enormous
potential
commitment
whichwill neverbe solveduntilseveralStationshave triedto tackle
it pith the full supportof theConservancy..
Yet a decision
as to
whetherwe try to persuadeNationalAllscums.to
curateour voucher
specimens
or whetherwe try to take.on'thecommitment
ourselves
is of
lesS-imriortan
at
cethe presenttimethan thatwe agreethatpermission
to collecton NationalNatureReservesshoulabe subjectto the .
condition
that at least one voucher specimen of every species collected
be offered to the Conservarey... The specimens need not be acceptedef COUrSe
if we alreadypossessa voucherof it brknowwhereone is
lacated.
Thereare,I appreciate,
dozensof difficulties
in this
latterproposaland therewillbe lots of exceptims(i.e.
exceptional
people)involved.NUch moreimportant
is the fact that
we cannot
put thiscondition
on collecting
intoeffectuntila deployed
collections
service
is established
and thiswill takemanyyearsto
becomestaffedfully. Inevitably,
therewillbe somecentresof the
ConservanCy
- perhapsat NenksWood and Edinburgh
- whereit may
be possibleto makebetterprogressthanelsewhere.Againthis
doesnot matterso longas someprogressis made somewhere.
- 87 -
benefitsShouldbefelt. Just as the advent
Additional
at the 9th Meeting
appliedentomologist
of our firstfull-time
turningnointt in its
a,
was
Committee
Liaison
of the Entomological
be
cura-tor
first
our
of
arrival
the
so-could
deliberations,
supply_.
must
colleagues,
another.From thc start,he and laterhis
manner
such
service.Untilin some
a broad7based_invertebrate
experts
ofd invertebrate
the "eutside"-worl
111:-16t
theConservancy
It
versa.
vice
or
them
of
eut
it will not get the-best
half-way,
in
half-way
world
"outside"
the
Conservancy
the
is because.
.
. . has met
so
havoLbeen.
we
.that
s-,
respectOfr other-organismnotahIrbirds,:
in makinginroadsinto theproblemsof their-CohacrvatiOn
.successful
Branch,workingin
The timehas now comewhen the Conservation
Sectionand the
Research
with the Conservation
collaboration
a similarorganizational
that
demand
Should
Centrei
Records
BiofOgicaI
invertebrates.
on
of
the.conservati
effortbe 'put-into
. _
_
Summar of Discussion
valueof main-,
whichdoubted'the
question
ato
.--Ifi'nnswer........
any way in which-Mr. Huxleysaid-that
r
'specimens,
taining-VOUche
and he'did not
ofWork can be reducedis.agood thing.,
the'VolUMe
should
to maintainavOuChercollection
intend -thatthe-suggestion
ways would
'eventually
beHapplled too rigorously.He thought_that
Mr. Eltona sYStehr.
doingawaywith-theneedfor stich
be-foUnd'of
be-cOmbined-withvoucher r---uld
collections.co..--reference
aSked'whether
system:
celIaCtionS-in-one
can
thatalthougha VoucherCellection
HuxIey-said
-not alwayswork
this-did
collection,
as a reference
alSe-funetion
collection
A reference
-03111a.
inl-the-reverse-way.
_ . be bOught fram
Tor instance.
entOmplogical-suppliers
-
-88Table I'
NuMbersof speciesin Conservancy
Invertebrate
Collections
(plussign
= presentin unknownnuMbers;nuMbersin brackets= sub-totals).
Namc
Edin- FUrze- Merle-MonksNor-.
burgh
rook wood Wood wich Oxford Rhum
Pangor
P''rotozoa
Porifera
Coelonterata
Plat holminthes
Rotatoria
Nematoda
Annelids
Pol chaota
Oli ochaeta
Hirudinea
Aollusca
Lamellibranchia
Castrooda
ARTPLROPODA:
rustacca
A ia oda
Anachnida
45
iliones
5
Araneae
40
Acari
Psoudosco iones
Insocta.
400
A ten ota
Orthotera
Dermater-a
Pleco.tera
hemerotera
Odonata
HeIrd tera
MC ale tora
Neurotera
Trich tern
Le idoptera
Colcotera
c.2001
H no tera
c.2001
Di tera
TOTAL445
1
50
.+
430
25
7
10
30
11
15_
1011
50
1
50
8
5 •
28
5
1
425
13
4
420
11
1114
1478
2
35
1
8
1
13
3 1.
13Ë
10 :
162
11
151
30
700E
488
200
50 '
f
•
50
, 10
30
2741
12
150
110
2
1482
60
5
1
1
12
8
50
10
40
1
150
711
50
3
1
165
30
5
c .250
42
1
201 ,
201
21
4
4
152
169
50
100
350
1357 ! 1966
2
111
2
11
7'
100
1
10
5
28.5e
420
400 1
500
12
c. 611 80
1926
1388
60
0
10
165 ,1 e.250
!
51'
83'
542'
- 89 -
GENERAL
DISCUSSION
meMbersraised
Duringthissessionon the last afternoon,
of the
of valueto the discussions
peintswhichtheyconsidered
is an editedversion,%thembs.Althoughthe following
different
discussed
to all the subjects
we have triedto includea reference
the manypointsof view expressed
in orderto placeon .record
duringtheSymposium.
E.DUffey said thatone of the
Openingthe discussion,
was thatafterfifteen.
arisingout of the Symposium
basic-points
were .
years,we were stillworkingon thebasisof proposals.which
Paper(Cmd.7122)..Did.
yearsago in the 1947-White
framedseventeen
conservation_
theConScrvancy's
we not requirea re-assessment-of
all thathadbeen said:about
policy,takinginto-consideration
studiesas well as otherwildlife?We couldnot
invertebrate
isolation.
conserVation.ia
of invertebrate
discussthe-problems
of the ConserVancy's.rscientific.staff.
from the othercommitMents
.
amongthe meMbersof theSympoSium
hadbeen generalagreement
-Tlaere
of theConscrvandts
objectives
the scientific
thata re-assessment-of
waS-necessary..
programe in relationto conservation
research
A secondbasicpointhe•said;arosefrom-the-papers-and
surveys.Was theregeneralagreement'
en invertebrate
discussions
and developed-in
thatsurvoyson NatureReservesshouldbe-extended
with-similar
and-otherorganizations
orderthatthe Conservancy
what it was we were trying.to.Consere?
should'know
responsibilities
of wildlife-on '
In short,did we not needa nationalinventory
NatureReserves?
J. Eggelingsaid thatin Scotland,the firsttaskwas..te
of-majorhabitatson NatureReserves,
includerepresentatives
judgedin the firstinstancefrom the botanicalinterest.A state-.
needcd.....
policy as a wholewas certainly
mentof-theConservancy's
beCauseit was used in a
thiswerd "interest"
He had mentioned
staff,it was generally
varietyof-ways.Among thc Conservancy's
Should
but'itwas not alwaysclearto others.-"Interest"
understood,
of habitatsand
be definedso thatpolicyaboutthe preservation
hoW far we ware
speciescouldbe known. It was not knownat.present
rare speciesen Reservesin relationto
preparedto go to safeguard
ef the habitat. This alse tiedup with-therequest
preservation
thatwe Shouldhavewildlifesurveyson land outsideNatureReserves.
- 90-
whatDr. Eggelinghad saidand
J. Satchellsupported
shouldbe made of what our aims should
felt thata clearstatement
'Maintenance
.of invertebrates.
be in relationto the conservation
Shouldbe acceptedas theprimaryaim,but
of habitatdiversity
speciesof
of particular
anetheraim shouldbe thepreservation
specialinterest.He wouldlike to see writteninto the recommendof the view that theseaims
an expression
ationsof the Symposium
lonsis.He pointedout
couldbest be achievedon an international
especially
communities,
thata numberof importantinvertebrate
in the current
ones,were not well represented
marineand freshwater
snecieshad
thata particular
seriesofReserves. The "interest"
of appeal
(e.g.relictspecies)or aesthetic,
mightbe scientific
to the generalpUblic.
C. Eltonwishedto clarifyone pointwhichaffectedthe
earlyhistoryof the NatureConservancy.At thattimeCaptain
of havinga largestafftrained
the possibility
Diver considered
work,but insteadan
in taxonomyto do surveyand inventory
and'agreatdealhad
was instituted
of regionalofficers
organisation
deVelepodfromthat. Therehad neverbeen a policyor a feeling
was underan obligation
that,becauseof the Charter,the Consurvancy
in its Reservesalthoughit was assumedthat
to make inventories
thatit was moreimportant
thesewouldbe done. Mr. EltonthcAight
by
intoa fund of knowledge
to Collectand convertthisinformation
studieson Reserves.SomeReserveshad been
makihgscientific
e.g.HavergateIslandfor the
largolYfor one species,
established
to make a surveyof the mud in whichtho
Avocet. Was it necessary
in
shouldbo established
Avocetfed? SOme systemof nriorities
was
casesof thissort. He saidthat the pointhe wished.to.make
not the impossible.
thatwe Shouldattemptthepossible,,
secondpointabout
referredto Dr. Satchell's
F..Perring
interestand wondered
of speciesof narticular
the preservation
occurredcouldbe the
they
where
land
of
whetherthe acquisition
thanthe Conservancy.
rather
Trusts
ef Naturalists'
responsibility
Societyof theBritishIsleswas doingthisthrough
The Betanical
method. In many cases,small
gainedby the inventory
information
to theNature
embarrassment
an
be
would
so
or
areasof an acre
because
yet therewas a case for theirpreservation
Conservancy,
of the existenceof a rarity.
acceptedthe
E. DuffeywonderedwhethertheConservancy
to conserverareanimalspecies,and how raritywas
commitment
assessed.Dr. Satchellhad givensomeguidanceon one aspectof this,
aboutthe rarityof
how was information
Reserves,
but when selecting
speciesto be used? Were the criteriathc samefor all animals,or
forbirdscomparedwith invertebrates?,
standards
were theredifferent
- 91 -
J. Satchellsaid thatwhat he had in mind whenhe menwas thatpriorityshouldbe given
view-point
tionedan international
to the endemicspeciesof thiscountryand thosewhichhave their
should'be
Othercountric.s
in Britain.maincentresof distribution
Unionfor the
to do likewisethroughthe International
encouraged
and more
of Nature,so thata more comprehensive
Conservation
couldbe formulated.
basedpolicyOf conservation
scientifically
J—Frazer askedwhetherit was not possiblethat"an
a Reserveestablished
awayf"rom
aninalspeciesmight-mpve
interesting
fer its protection.He alsoWonderedat_whatpointin time
becomenaturalised.
Speciescouldbe regardedas-having
introduced
from
W. O. Steelgave as an examPlea speciesof beetleNew Zealand,firstrecordedin 1940,whichwas now breedingin
speciesof the
Berkshire.He alsomentionedtwO otherNew.Zealand
samegenus. One, firstrecordedin 1870,was now one of the commonest
Britishbeetles; the otherhad comein justbeforetho War and now
somespeciescould'
extendedas far as Yorkshire.On thisevidence,
withintwentyyears.
becom naturalised
C. Eltonwas of the opinionthatsuchspeciesshouldbe
thatno specialeffortShouldbe made to preservethem.
studied.but
and
between.England
J. Eggelingstressedthe difference
Waleson the one handand Scotlandon the otherin regardto their
that-thezhole-of'
Trusts. Therewas no likelihood
Naturalists'
there -Werejust-as
Scotlandwouldbe coveredby County-Trusts,-yetScotland
Of-interest.in
plantpopulations
manyhazardsto animal.and
as in Englandand Wales.
out
Of-carrying
Severalmembersthenspokeon the'ProbleMS
National
both insideandcutside
of wildlife,
inventory
a-natienal
of a monthly
publication
includedtheSuggestions
Nature-Reserves.
species
aboutintereating
journalin whichinfo/nation
conservation
staff.
Cenei- -vandy
te-cOuldbe madeaVailable
and theirdistributipn
consultedto
Should-be
specialista
thattaxonomic.
It was sUggested
meMbersthought
and'other
.were in-danger
species
-findout'which
to copewith
ahouldhaveits own taxonomists
7 -thatthe-Conservaney
the wildlifeon NationalNatureReserves.surveying
pro-blem.of
-:the
The argumentagaincame roundto the questionof rarityand how commitments.
be-definedin relationto the Conservancy's
this-Should
the proposalwhich
was referred'to,particularly
White-Paper
-The-l947
flora"
theBritishfauna-and
that "A representative-shmpie'of
stated.
Seriesof NatureReserves.
on the-National
Shouldbe preserved
were•
rare specie-a:which
C. Eltonstatedthat-there-were
that it.wasbetternot:to
specialcases,;e.g.the-Avocet,.blit
emphasizethemtoo much. On WythwaHillne6rOxford,therewere
6,000- 7,000speciesof animnls,and so one fifthcf the.British
On 120faunaoccurredon two.squaremilesof-English.cOunttyaide.'
thehe wouldhave thoughtthatnearly.all
NationalNatureReserves,.
most of the rarities.
speciesof theBritishfaunawere reservedincluding,
- 92 N. Mooreaskedwhetherit was agreedthatas many species
as possibleshouldbe conserved
so that-they-Coad-be
passedon to
futuregenerations.The rare oneSShouldbe conserved
as far as
resources
permitted.Richnessof floraand faunamustbe maintained,
endemicspeciesconserved,
and dertain
butterflies
and birdsprotected
for aesthetic
and educational
reasons.
R. Goodiersaidthatthe maintenance
of diversity
was a
main aim accordingte manyparticipants
in the Symposium,
but that
it couldnotbe appliedin all cases. It had'not-been
too difficult
to.obtaina representative
selection
of different.
majorhabitats,
but
diversity
in relationto the multitude
of the Britishfaunawas
impossible
to achieve.Diversitycouldbe established
in relation
te vegetation,
but not in the samesenseto soilsand climate.Diversity
shouldnotbe introduced
at the expenseof sonic,
Otherfactorwhichwas
worthconserving.In the selection
of characteristiclabitats
which
were represented
on NationalNatureReserves,
a certainminimol
area for eachwas implicit,
so thattheycouldsurviveand their
faunaswith them. Theymust not be endangered
in an attemptto
achievemaximumdiversity.
J. G. Skellam endorsed
Mr. Goodier's
remarksand said:that
certaintypesof uniformity.were
necessary
to enhancescientific
knowledge.
N. Mooredeniedthatit was his intentien-that
diversificationshoulddestroyan existinghabitat; if it did so, it could
not be supported.
T. Huxleyagreedthatit was impossible
to includethe
requirements
of all 20,000Britishinsectson NationalNatureReserves
but pointedout thatin practice,someapproximation
to thiswas being
attempted
when additional
habitats-desired
on NationalNatureReserves
werebeingdefined.AlthoughtheConservancy
beganby definingin
verybroad termswhat shouldbe includedon NatureReserves,they
werenew chosen- woodlandsfor instance- on thebasis of a dominant
species.The samnprinciple
mightforma similarbasisfor the
selection
of futurereservesfor invertebrates;
actionof thisSort
wouldbe neededto.safeguard
endemicand rare species.
R. Steelerepliedthatwoodlandreserveswere not
selected
purelyon thebasis of dominantspeciesbut thatbroader
principles
were takeninto accbunt.
M. G. Morriswarnedngainstsuchattitudesthat"if the
florawas conservedthe faunalookedafteritself"and askedmeMbers
of the Symposiumto keep the exceptions
in mind.
C. Eltonpointedout thatthe habitatclassification
in
management
planswentbeyondthis to includeanimalhabitatssuch
asdeadwood.
- 93 -
of MoccasDeer
to theColeoptera
A. M. Massee-referred
faunaoccurredin a very limitedarea. He
Park wherean exceptional
thatsuchplacesshouldbe left out of
any suggestion
deprecated
NatureReservesSeriesbecausethe insectswere not
the National
so rare on theContihent.
the valueof surveywork in assessing
M. Labernmentioned
in tao majorwoodlandreserves
the successor failureof management
region.
in the south-east
J. Phillipssaid thatan attemptwas beingmadeto
on thosereservesand thatit would
coppice-with-standards
perpetuate
speciesof such
be valuableto knowwhethertherewere invertebrate
to use.
whichformof management
whichwouldindicate
conditions
work was done.for
J. G. Skellampointedout that.survey
a specialpurpose. It couldbe a meansto an end for the selection
of thebest sitefor a naturereserve,or to measurethe success
practice.The aimsof a surveyshouldbe clearly
of conservation
stated.
on manngement.resear.ch
E. Duffeysaidthatthe ignorance
He statedthat
illustrated.
had been abundantly
for invertebrates
were intensivestudieson single
two usefultypesof investigation
of
and studiesof the,faunas
speciesin relationto theirhabitats,
habitatswhichwere changingor beingmodifiedin some
particular
way. It was betterto takea limitedproblemand expandthis
in relationto the mostusefUllinesworthfollowing.To do this,
that
were requiredand he suggested
manymoreappliedecologists
to theConservancy, ' thisneedshouldbe emphasized
J. Dempstersaidthatif the principlethatspecialspecies
was required
thenmoreinformation
were accepted,
mustbe conserved
necessary
was
It
species.
the
on whatwas governingthe nudbersof
of the speciesin the fieldand more
to know thepatternof behaviour
aboutits generalbiology.
thatfor many years,he had.stressed.
C. Eltonmentioned
the needfor more studieson populations.
was made
reference
In the finalpart of theDiscussion,
work.
inventory
and
survey
problemsof scientific
to thepractical
scientific
all
problem,
national
thatas thisis a
It was emphasized
in someway or other. On '
to contribute
staffshouldbe encouraged
i'twas gene:rally
data,
the questionof storing'andindexingthe.
at the present
agreedthatone or two centreswouldbe sufficient
Experimental
Wood
Monks
and
Scotland
time; the Edinburghofficefor
g
the
latter,
the
to
StationforEnglandand Wales. With regard
body while1119 ResearchSectionwouldact as a collating
Cons,rvation
Centre4
Redords
materialwouldbe storedby the Biological
processed
.z,,,
undertaken
be
not
could
It was pointedout however,that thistask
staff.
withoutadditional
t '
á
á
UJ,30
H