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JournalofEcology(1981), 69, 1017-1059

A COMPARATIVE STUDY OF GERMINATION


CHARACTERISTICS IN A LOCAL FLORA
J. P. GRIME, G. MASON, A. V. CURTIS, J. RODMAN, S. R. BAND,
M. A. G. MOWFORTH, A. M. NEAL AND S. SHAW

ResearchCouncilUnitofComparative
NaturalEnvironment PlantEcology,
ofBotany,The University,
Department S 102TN
Sheffield

SUMMARY

(1) Using a standardizedprocedure,a laboratorystudywas made of thegermination


characteristicsof seeds collectedfroma wide rangeof habitatsin the Sheffield region.
Measurements wereconductedon freshly-collected seeds and on samplessubjectedto dry
storage,chillingand scarification. Responses to temperatureand lightfluxwere also
examined.
(2) The data have been used to comparethegermination biologyofgroupsof species
classifiedwith respect to various criteriaincludinglife-form, family,geographical
distribution,ecology,and seed shape,weightand colour.
(3) Marked differences were observedin the capacity of freshly-collected seeds for
immediategermination.Of the 403 species examined, 158 failed to exceed 10%
germinationbut 128 attainedvalues greaterthan 80%. Germinationwas high in the
majorityof grasses and low in manyannual forbsand woody species. Withrespectto
initial germinability,major familiescould be arranged in the series Gramineae >
Compositae > Leguminosae= Cyperaceae > Umbelliferae. Many small-seededspecies
wereable to germinate immediately aftercollectionand seeds of thesespecieswereoften
elongatedor conical and had antrorsehairs or teethon the dispersule.High initial
germinability was conspicuousamongthe speciesof greatestabundancein theSheffield
flora.
(4) In the majorityof species,germination percentageincreasedduringdrystorage;
this effectwas most markedin small-seededspecies. Among the seventy-five species
which respondedto chilling,some germinatedat low temperature in darknesswhilst
othersweredependentupon subsequentexposureto lightor to highertemperature or to
both.Responsesto chillingwerecharacteristic of theUmbelliferae. In all of thelegumes
examined,rapidgermination to a highpercentagewas broughtaboutby scarification.
(5) Underthe experimental conditions,all of the annual grassesshowedthepotential
forrapid germination. High rates were also observedin manyof the annual forbsand
perennialgrasses.Low ratesof germination occurredin the majorityof sedges,shrubs
and trees,and were particularly commonin species of northern distribution in Britain.
Rapid germination was characteristic ofthespeciesofgreatestabundanceintheSheffield
flora.Rate of germination showeda progressive declinewithincreasingseed weight,and,
withsome exceptions,therewas a positivecorrelationbetweenrateof germination and
therelativegrowthrateoftheseedling.
(6) In sixteenspecies, germinationin the lightwas found to be dependentupon
exposureto diurnalfluctuations in temperature. Underconstanttemperature conditions,
the majorityof grasses, legumes and compositesgerminatedover a wide range of
temperature, and the same featurewas evidentin species of ubiquitousor southern
distributionin the BritishIsles. A requirementfor relativelyhigh temperaturewas
apparentin sedges, in plantsof northerndistribution and in a majorityof the marsh
plants.The rangeof constanttemperatures conduciveto germination tendedto be wider
in grasslandplantsthan in woodland species. Rapid germination over a wide rangeof
temperatureoccurredin many of the species which attaingreatestabundance in the
Sheffieldflora.
Publications
0022-0477/81/1100-1017$02.00 (?1981 BlackwellScientific
1017

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1018 Germination in a localflora
characteristics
(7) Althoughgermination in mostspecieswas promotedby light,some wereinhibited
under relativelyhigh light flux.In 104 species a marked reductionin germination
occurredifseeds werekeptinthedark,and in manyspeciesthisinhibitory effectcouldbe
by eitheror bothexcludingtemperature
intensified and abandoningtheuse of
fluctuations
a green'safety'light.The capacityforgermination in darknesswas observedin all ofthe
legumesand manyofthegrasses.Dark germination did notoccur in theCyperaceaeand
was uncommonintheCompositae.The inhibitory effectofdarknesswas characteristicof
manyofthespeciesknownto formreservesofburiedseeds,butit occurredalso in certain
specieswithmoretransient seed banks.
(8) There were recurrentassociationsbetweenfeaturesof seed morphologyand of
germination,severalofwhichcoincidedwithparticularecologicalcharacteristics.
(9) The functionalsignificanceof some of thegermination observedin
characteristics
this studyleads us to the conclusionthatcertainregenerative mechanismsin the field
maybe predictedfromthelaboratorycharacteristics oftheseed.

INTRODUCTION

The resultsof manycomprehensive studies(Ridley 1930; Salisbury1942; Isely 1947;


Ross-Craig 1948-74; Baker 1972; van der Pijl 1972; Hubbard 1976; Lhotska &
Chrtkova1978) show clearlythatflowering plantsdifferconsiderably withrespectto the
number,size, shape and dispersalmechanisms of theirseeds and fruits. Althoughsuch
information to plantecologists(Salisbury1942; Harper,Lovell
is of considerableinterest
& Moore 1970; Stebbins1971; Baker 1972), its value is severelylimitedby ignorance
of otheraspects of the regenerative process.In particular,thereis a shortageof data
concerningthe germination biology of floweringplants. The objectiveof the study
describedin thispaperis to describesomeof thegermination characteristicsof a rangeof
flowering in a
plants drawn froma local flora northernEngland.Using standardized
procedure,a series of germinationtests was conducted,to assess germinability
immediatelyfollowingseed collection,afterstorage,and undervarious conditionsof
temperature and lightflux.
The resultshavebeenusedto comparethegermination requirements ofgroupsofspecies
of contrastedecology,and an attempt has beenmadeto relate germination characteristics
inthelaboratory to themechanisms which controlthe time and place at which germination
occursundernaturalconditions.

MATERIALS AND METHODS

Seed* collections
Seeds of 403 speciesweretakenfromnaturalpopulationsduringthe period23 May
1972 to 4 August 1976. Each specieswas collectedin bulkfromplantsestablishedin a
smallarea (10-20 m2). Considerableeffort
relatively was madeto collectonlyripeseed,to
sample both large and small inflorescences, and, as far as possible,to includeseeds
representativeofthefullrangein size and morphology presentat each site.
Details of the collectionsitesare givenin Appendix2. Withtheexceptionof fifty-six
speciesof southern, montaneor coastal distribution,all collectionsweremade withinan
area of2400 km2aroundSheffield.
* Here and subsequentlythe term'seed' refersto the germinating thatin some cases would be
structure
moreaccuratelydescribedas a fruit.

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J. P. GRIME et al. 1019
Seed description
The experiments describedin thisstudyprovidean opportunity to relategermination
characteristicsto other attributesof the seed. Accordingly,a standardizedseries of
measurements was conductedon each seed collection,and the resultsare recorded
in Appendix2. In additionto information relatingto size and structure, of
a description
seed colouris included(based uponmatching withMunsellcolours(Munsell1954) under
a binocularmicroscope).The seeds of certainspecieswerefoundto be multicoloured or
variablein colouror both;data forthesespeciesarenotincludedin Appendix2.

Seed storage
Immediately following the
collection, seeds of fleshyfruitswereextractedand washed
by hand in tap water. Seeds of all species were then allowed to dry at laboratory
temperaturefor 1 or 2 days, mixedthoroughly, and placed in dry storagein plastic
in darknessat a temperature
containers of5 ?C.

Germination
experiments
Initialtests
Beforeplacingthe seeds in drystoragethe capacityof thefreshly-collected seeds for
immediate germination seedswereplacedon WhatmanNo. 1
was tested.Fiftyunsterilized
paperwas moistened
The filter
paperin each of two Petri-dishes.
filter withdistilledwater
and thedishesweretransferred to a growthroomproviding visibleradiationat a fluxof40
W m-2 ('Warm-white' tubes+ tungsten
fluorescent bulbs),overa 15-hday at 20 ?C and a
nighttemperature of 15 OC. Each dish was inspecteddaily,and germinated seeds were
countedand removed.The seed was consideredto have germinated whentheradiclefirst
emerged.The minimum durationof a testwas 30 days and countingcontinueduntilno
germination occurredfor5 successivedays. Furthertestswereconductedon subsamples
removedfromdrystorage3, 6 and 12 monthsaftercollection.

Dormancybreaking
In theinitialtestsmanyspeciesexhibitedlow germination percentages or low ratesof
germination or both.Wherethisoccurred,subsamplesof seedsweresubjectedto various
additionaltreatments in an attemptto facilitate
germination.Because of thelargescale of
theinvestigation it was impracticable to examinetheresponseof each speciesto a wide
range of treatments. Three main treatments were adopted (dry storage,chillingand
and thesewere appliedin a series,progressionthroughwhichdepended
scarification),
uponcontinuedfailureto inducerapidgermination to a highpercentage.
It should be emphasizedthat these procedureswere not designedto provide an
exhaustivestudyof dormancy.The purposewas to identify major typesof dormancy
associatedwithparticular ecologicalgroups,and to obtainseedsin a conditionsuitablefor
laterstudies(see below)oftheeffects oftemperature and lightupongermination.
In thedry-storage treatment, a subsampleof theseeds was removedfromcold storage
(usually within6 monthsof collection)and placed in darknessin dry conditionsat
laboratory temperature (about 20 OC). Aftera minimum of 5 weeks,100 seedsweretested
forgermination as describedabove.
In thescarification
treatment, 100 seedswereremovedfromdrystorageand thetestaof
each was perforated usinga small scalpel or mountedneedle.The germinability of the
seedswas thenmeasuredas before.

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1020 characteristics
Germination in a localflora
a largesubsamplewas immersed
In thechillingtreatment, in moistsand in a refrigerator
maintaininga temperature of 5 OC + 1 OC. Each subsampleoccupieda separatebox of
sand and the seeds were placed betweentwo layersof terylenecloth.At intervalsof
approximately 45 days,seedswereremovedfromthesand and testedforgerminability.
In a smallnumberof speciesthe effectupon the imbibedseed of incubationat room
temperaturewas assessed.The procedurewas identicalto thatforchillingexceptthatthe
seedswerestoredat about20 0C.

Responseto temperature
The influence oftemperature upongermination percentage and ratewas studiedat thirty
constanttemperatures over the range5-40 OC, usingtemperature-gradient bars closely
similarindesignto thosedescribedbyMason (1976).
Each bar was coveredwitha singlelayerof Whatman3MM chromatography paper.
Beforesowing,the paper was thoroughly saturatedwithwaterand rolledto removeair
bubblesand to ensureclose contactwiththebar.The edgesofthepaperdippedbelowthe
sides of the bar into a waterreservoirand acted as a wick transporting waterto the
germination area. At thehotendofthebar an additionalsourceof moisturewas provided
by threefineplastictubesthroughwhichwaterwas deliveredto thepaperby meansof a
peristaltic
pump.
Only species showinghighgerminability (greaterthan75% after14 days incubation)
and rapidratesof germination in theinitialtestswereconsideredforexperiments on the
bars. Preparationof the seed forsowingusuallyinvolvedremovalof the outerinvesting
structures,as far as possible,withoutdamagingthe seed. Whereseeds were knownto
possessa testaimpermeable to watertheywerescarified and allowedto imbibewaterfora
shortperiodbeforesowing.
The seeds weresownin groupsof 30 at intervals of 2.2 cm alongthebar. Each group
was sown in a line transversely across the bar and spanninga widthof about 0.5 cm
(about 0. 13 OC). Thirtygroupsof seeds wereaccommodatedon each bar. Infrequently,
thenumberof seeds in each groupwas reducedto 20 iftheseed was especiallylargeor
was availableonlyin limitedquantity.Each groupof seeds was coveredby a miniature
transparent cloche,designedto restrictevaporativewaterloss fromthe seeds and the
surfaceofthebar.
Germinatedseeds were counteddaily and removed.The minimumdurationof an
experimentwas 14 days, and countingwas extendedconsiderablyin species which
continuedto germinate.Whereseeds remainedon the bars fora protractedperiodwe
sometimeshad to replacethe chromatography paper in orderto avoid heavybacterial
infection.

Responseto light
Onlytheeffectof lightfluxupongermination was studied.Earlierstudieson theeffects
of both fluxand spectralcompositionhave been reportedelsewhere(Grime & Jarvis
1975).
In Experiment 1, seeds wereexposedto threetreatments ('light','shade' and 'dark').In
the 'light'treatment,fivereplicatetransparentdishes,each containing20 seeds, were
placedin a growthroomwitha visiblelightfluxof 40 W cm-2('Warm-white' fluorescent
tubessupplemented by tungstenbulbs),a daylengthof 15 h, and a temperature regimeof
20 OC (day) and 15 OC (night).Seeds were sown on Whatman No. 1 filterpaper
moistenedwithdistilledwater,and germinated seeds were countedand removeddaily.

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J. P. GRIME et al. 1021
Conditionsand procedurein the 'shade' treatment were identicalto thatin the 'light',
exceptthatthefluxofradiationwas reducedto 97 ,uWcm-2(2.4% ofthe'light'treatment)
by meansof a neutralfilter. In the'dark' treatment thedishescontaining theseeds were
placedin a light-proofbox withinthegrowthroom;in orderto recordgermination thebox
was opened brieflyeach day under a low-intensity greenlight.The experiment was
continueduntilno germination was observedforfiveconsecutivedays in any of thethree
treatments.
Briefilluminationofseedsbylow-intensity greenlightmayeventually cause germination
in certainspecies(Grime& Jarvis1975; Blom 1978).Furthermore, Experiment 1 involved
a diurnalfluctuation in temperature (20/15 IC), a conditionknownto stimulatethe
germination of certainspecies even in continuousdarkness(K. Thompson,Grime &
Mason 1977). Accordingly,a second experiment(Experiment2) was conductedon
selectedspecies.Seeds weresubjectedto threetreatments. The firsttreatmentreproduced
the conditionsin the 'light'treatment of Experiment1. The remainingseeds were in
continuousdarkness.In one treatment a diurnaltemperature fluctuationof 20/15IC was
applied,whilstin theotherthedishesof seeds weremaintainedat a temperature of 20 +
0*1 OC withina water-bath.Germinatedseeds could not be removedfromthe dark
treatments, but daily countsweremade in the'light'treatment, and theexperiment was
terminated whengermination theredeclinedto a low rate.
Standardization
Practicalworkextendedover6 years,and involveddailygermination countson a wide
and in particulartherecognition
varietyof species.In orderto standardizeprocedures, of
germinated seeds,one author(JPG) was responsibleforgermination countingon at least2
daysin each weekthroughout theinvestigation.
Data analysisandpresentation
The resultsof all testsare presentedas finalgermination percentageand as the time
requiredfor50% ofthefinalgermination percentage to be attained('half-time',
P5O).
Analysis of the data fromthe temperature-gradient bar experimentfollowedthe
proceduredescribedby P. A. Thompson(1970a, b). For each speciesa graphwas drawn
showingthetimetakenat each temperature to attain50% of themaximumgermination
percentage.Each successive day, plots were made of the maximumand minimum
temperatureat whicha germination percentageequal to or exceeding50% of the final
germinationpercentageattainedat the mostfavourabletemperature was achieved.The
pointswereconnectedby a smoothcurve,fittedby eye. Thus a largequantityof data is
summarizedin one diagram.Specimencurves are shownin Fig. 1, and the essential
ofthecurveforeach speciesaredescribedin Appendix2.
characteristics

RESULTS
The largebody of data accumulatedin thisinvestigationprovidesmanyopportunities for
The originaldata collectedforparticularspeciesare available
analysisand interpretation.
on request.Here we attemptto classifythe specieswithrespectto a varietyof criteria,
life-forms
includingseed characteristics, and fieldecology,and to examinevariationin
germination both withinand betweenthe resultinggroupsof species.A
characteristics
summaryof the resultsof the germination experiments conductedon each species is
used to analysetheresultsis
includedin Appendix2, and a synopsisof theclassifications
presentedin Appendix3.

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1022 Germination in a localflora
characteristics
16 - (a) Type 1 16 ( b) Type II
14 - 0 0 14 l
12 - 2 _e

1O I0 I \
4 4 __-___ __.1

20 ?
2Type I I 6II) (d) Te VType I

44 -
40 -
~~~~~~~~~~~~~~~~~~12-
36 - ~~~~~~~~~~~~~~~~~~10
323628
32 -
28 -
~ ~ ~ ~ ~ ~ ~ ~ ~~ T~~~~~~~~~~~~
24-
~ ~ ~ ~ ~ ~ 4 0 0
2
20 7 I 0
OL I I I I 1

5 10 IS 20 25 30 35 5 10 IS 20 25 30 35 40
Temperature(IC)

FIG. 1. The maintypesofgermination Each curvehas beenconstructed


responseto temperature.
by plottingforsuccessivedays aftersowingthemaximumand minimumtemperatures at which
50% maximumgermination is attained.(a) Ballota nigra(Type I), (b) Koeleria cristata(Type
II), (c) Miliumeffusum (Type III), (d) Senecio squalidus(Type IV).

Capacityforimmediate germination
Distributions of initialgerminabilityin selectedtypesof plantsare shownin Fig. 2, and
a morecomprehensive comparisoninTable 1. Speciesdiffered
statistical considerably with
respectto the capacityof freshly-collected seeds to germinateunderthe experimental
conditions,and the distribution of finalgermination percentageswithinthe 403 species
showeda strongly bimodalpattern. At one extreme were158 species(39%) in whichfewer
than 10% of seeds germinated, whilstat the otherwere 128 species (32%) the seeds of
whichshowedmorethan80% germination. Justoverhalfthespeciesinvestigated failedto
achieve50% germination, and in approximately one-quarternoneoftheseedswas capable
ofimmediate germination.
Differences in germinability were associatedwithlife-form. A highproportion of the
annualand perennialgrasseswas capable of germinating immediately aftercollection,in
markedcontrastto boththeannualforbsand thewoodyspecies.The fivemajorfamilies
represented in the seed collectionsformeda seriesdecliningin germinability as follows:
Gramineae> Compositae> Leguminosaeand Cyperaceae> Umbelliferae.
An extensiveliterature(Brenchley& Warington1930; Chippindale& Milton 1934;
Milton 1939; Champness& Morris1948; Chancellor1968; Wesson & Wareing1969a;
Watkinson1978; K. Thompson& Grime 1979) suggeststhat underfieldconditions
the seeds of manyBritishgrassesgerminatesoon afterreleasein the late summerand
autumn.However,itis also clearfromthesestudiesthatamongthespecieswhichexhibited
high initialgerminability in the presentstudythereare some (e.g. Agrostistenuis*,
Deschampsiacespitosa,Holcus lanatus,Poa annua,P. trivialis)in whicha proportion of
theseeds do notgerminate immediately and may remainlive,in thesoil,foran extended
period.Most of thegrassesin thiscategoryhave seeds whichare smalland compactand
* Nomenclature
followsthatofClapham,Tutin& Warburg(1962).

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J. P. GRIME et al. 1023

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1024 Germination
characteristics
in a localflora
TABLE 1. Numberof species,groupedin variousclasses, with?50% and >50%
germination of seeds immediately aftercollectionand afterdrystorage,and time
(t50)requiredforhalfthefinalgermination to be reached.The valuesin the'>50%'
columnsare based upon themaximumpercentageattainedin any of thefourtests
on dry-storedseeds. The significancelevels betweenthe '?50%' and '>50%'
columnsreferto the probabilitywithwhichthe ratio of low- (<50%) to high-
(>50%) germinating speciesin theclass departsfromtheratioexpectedfromthat
observedfor all 403 species (chi-squaredtest). The significance levels between
freshand dry-stored seed columnsare based upon a comparisonwithineach class
betweenthe ratiosobtainedforfreshseeds and dry-stored seeds. The significance
levelsbetweenthetwo 't50'columnsreferto theprobability withwhichtheratioof
rapid (t50 < 1-4-days)to moreslowly(P0 > 4 days) germinating species departs
fromthe ratioexpectedfromthatobservedforall 267 species. Significance con-
ventionsin thisand othertables:* P < 0.05, ** P < 0 01, ***P < 0.1, - numbers
too smallforreliabletest.Whereno indicationis given,P > 0.05.
Germination
(%) Germination
rate,
Freshseeds Dry-storedseeds t50
<50% >50% ?50% >50% 1-4 days >4 days
All species 211 192 * 144 259 120 147
Life-form
Annualgrasses 3 6 0 9 9 ** 0
Annualforbs 59 * 23 * 34 48 28 21
Perennialgrasses 6 * 39 3 * 42 26 * 17
Perennialforbs 119 116 ** 84 151 56 * 99
Shrubsand trees 24 ** 8 23 * 9 1 * 10
Family
Compositae 12 * 40 4 * 48 30 * 18
Cyperaceae 13 5 8 10 0 * 10
Gramineae 9 * 45 3 * 51 35 * 17
Leguminosae 14 * 5 14 * 5 1 6
Umbelliferae 17 * 1 16 * 2 0 - 2
Britishdistribution
Northern 20 23 16 27 5 * 22
Southern 82 62 ** .53 91 46 47
Ubiquitous 68 69 * 43 94 49 47
Widespread 32 28 28 32 14 21
Local 9 10 4 15 6 10
Frequencyin Sheffield
flora
> 10-0% 1 ** 14 1 * 14 11 ** 3
1-1-10-0% 68 67 * 46 89 45 47
0-1-1 0% 87 78 ** 58 107 44 66
<0- 1% 55 30 * 39 46 18 30
Habitat
Disturbed 53 37 ** 31 59 43 * 16
Skeletal 30 31 11 ** 50 24 27
Marshland 42 42 29 55 17 * 40
Grassland 45 * 64 35 74 33 44
Woodland 41 ** 18 38 * 21 3 ** 20
Heightofinflorescence
(cm)
0-20 62 * 38 * 37 63 25 39
21-40 52 63 38 77 39 41
41-60 32 41 * 17 * 56 27 29
61-80 27 18 19 26 14 13
>80 38 32 33 * 37 15 25

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J. P. GRIME et al. 1025
TABLE 1-continued
Germination
(%) rate,
Germination
Freshseeds Dry-stored
seeds t50
<50% >50% ?50% >50% 1-4 days >4 days

Dehiscence 211 192 *** 144 259 120 147


Passive 191 183 * 126 248 116 137
Explosive 20 * 9 18 ** 11 4 10
Dispersuleshape
Spherical 27 24 25 * 26 6 ** 21
Ovoid,rhomboidalorturbinate55 * 33 * 40 * 48 19 32
Trigonousor triquetrous 21 15 15 21 12 9
Lenticular,reniform or subulate66 * 37 ** 42 61 29 34
Cylindricalor ligulate 12 ** 27 6 * 33
Clavate 10 * 2 6 6 232
Winged 6 4 5 5
Tadpole-shaped 4 11 1 * 14 J
Conical 10 * 39 4 * 45 27 * 19
Dispersuleappendages
Absent 168 123 * 117 * 174 73 107
Straightawn(s)or spine(s)only 13 12 8 * 17 9 9
Hygroscopicawn or spine 4 9 1 * 12 9 * 3
calyx
Pappus or persistent 10 * 41 4 * 47 28 * 19
Dispersulehairsor teeth
Absentor veryinconspicuous188 * 131 * 132 * 187 76 116
Radial or irregular 8 7 7 8 4 6
Antrorse 15 * 54 5 * 64 40 ** 25
Germinuleweight(mg)
<0- 10 19 * 47 * 5 * 61 33 28
0-10-0-99 87 97 * 55 129 63 70
1.00-9-99 86 * 40 * 66 * 60 22 40
>9-99 18 * 7 18 * 7 2 7
Germinulesurfacetexture
Smooth 121 89 * 83 127 52 79
muricate
Rugose,tuberculate,
or reticulate 44 44 * 29 59 32 29
Striate 27 19 19 27 14 14
Hairy 9 12 8 13 9 * 4
Striateand hairy 6 * 17 3 * 20 10 10
Mucilaginous 4 11 2 13 3 11
Germinulecolour(Munsellhue)
5R, 7-5R and IOR 39 28 ** 23 44 18 26
2-5YR 18 23 * 9 * 32 20 * 13
5YR 55 50 * 37 68 26 43
7.5YR 26 33 17 42 18 27
1OYR,2-5Y and 5Y 26 31 19 38 26 ** 12
Black 11 ** 0 7 * 4 2 - 2
Multicoloured 7 14 6 15
Variable 21 11 18 ** 14 10 24
Multicolouredand variable 5 - 2 - 5 - 2

whichappearsto
tendto be withoutappendagesand largehairs-a suiteofcharacteristics
be conduciveto burialinthesoil.
No obvious correlationswere detectedbetweeninitialgerminability and patternsof
withintheBritishIsles. Therewere,however,indicationsof a positive
species-distribution

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1026 Germination in a localflora
characteristics
relationship betweenthecapacityforimmediate germination and species-abundance inthe
local flora.Of thefifteen mostcommonly-recorded speciesin theSheffield area,onlyone
(Rubusfruticosusagg.) failedto exceed 50% germination in theinitialtest.By contrast,
morethanhalfofthespeciesin thetworarestclasses had germination percentages below
50%.
There was also an unusuallyhigh proportionof woodland species withlow initial
germinability; thiswas evidentinbothwoodyplantsandin manyofthewoodlandherbs.
No consistent relationshipwas apparentbetweenthecapacityoffreshly-collected seeds
to germinateand the averageheightof the inflorescences of the species. There was a
tendency,however,for plantswithexplosivemechanismsof dehiscenceto includean
unusuallyhigh proportionof species in whichthe seeds were incapableof immediate
germination.
The majorityofthespecieswithsmallseeds (less than0. 1 mg)attaineda highgermina-
tion percentagein the initialtest.Conspicuouswithinthisgroupwere species such as
Calluna vulgaris,Hypericumperforatum, Juncusarticulatus,J. effusus,J. inflexus,
Origanumvulgareand Thymusdrucei,all of whichare knownto accumulatelargeand
persistentreservoirs ofseedsin thesoil (K. Thompson& Grime1979).
Threecategoriesofseed shape(cylinders, 'tadpoles'and cones)werestrongly associated
withthe capacityforimmediategermination. Cylindricaland tadpole-shapedseeds are
particularly commonin theGramineae,whilstcones are characteristic of manygenerain
theCompositae.The highinitialgerminability ofthemajorityofthespeciesfromthesetwo
familiescan be relatedto further correlationsevidentin the results:the presenceof
hygroscopic appendages (Gramineae), pappus (Compositae) and antrorse (i.e.
backwardly-projected relativeto the directionof radicle extension)hairs or teeth
(Gramineaeand Compositae).In experimental studieswithboth composites(Sheldon
1974) and grasses (Peart 1979, 1981) it has been establishedthatthesetypesof seed
structureare characteristic of species in whichgermination normallyoccurs in seeds
lodgedon the soil surface.Radicle penetration and earlyseedlingestablishment in these
speciesappearsto benefitfromthe combinedeffectsof the seed shape and the antrorse
hairsor teeth,bothofwhichtendto anchortheseedin a relatively upright positioninloose
soil,litteror bryophyte mats,thusbringing thepointof imbibition and radicleemergence
intoclose contactwiththesoil.
It is interesting to note that none of the eleven species with black seeds, and
comparatively fewof thosewithdarkred (Munsellhues 5R-10R) seeds,achieveda high
germination percentage.A highproportion of the specieswhichwerepolymorphic with
respectto seed colour(mostofthemlegumes)failedto exceed50% germination.

Responseto drystorage
Appendix2 gives,formostspecies,estimatesof changein germination percentageand
rateduringdrystoragefor12 monthsin airat 5 'C and forabout5 weeksat about20 'C.
In a large numberof species,storagewas associatedwitha progressiveincreasein
germination percentageand rate. Germinability in some species remainedunaffected,
however,and in a minority therewas a declinein germination percentage.In thislast
categorytwo groupscan be distinguished. The firstcontainsfivespecies (Hedera helix,
Petasiteshybridus,Quercuspetraea,Tussilagofarfaraand Ulmusglabra) in whichhigh
oftheembryo.The second
was followedby desiccationand shrivelling
initialgerminability
and largergroupwas composedmainlyof legumesand specieswithinthe Geraniaceae;

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J. P. GRIME et al. 1027
here,increasingdormancyof viable seeds was associatedwithprogressivedryingand
hardening oftheseed coat.
Comparisonof freshly-collected and storedseeds withrespectto finalgermination
percentage(Fig. 2) revealsa markedincreasein germinability withstorage.Althoughthe
distributionof germination percentages in the403 speciesremainedbimodal,66 (31%) of
the211 specieswhichfailedto achieve50% germination as freshly-collectedseedexceeded
thisvalue afterstorage.The same calculationhas been made foreach of the classes of
species recognizedin Table 1. In certainclasses the effectof storagewas to increase
germination (>50%) in a proportioneithergreateror smallerthan would be expected
fromthevalue of 31% obtainedforall species.Conspicuouslydifferent was theresponse
of the seeds of shrubsand trees,in whichonly4.2% of the speciesthatfailedto attain
50% germination as freshseed were inducedto exceed this percentageafterstorage.
A similarvalue of 7.3% was obtainedforall thewoodlandspeciesofinitially low (<50%)
germinability.
These resultsare in contrastto the high proportion(63%) of the low-germinating
(<50%) speciesfromskeletalhabitatsin whichgermination exceeded50% afterstorage.
Strongbeneficialeffectsof storage upon species with littlegerminationwhen fresh
occurredin the Compositae(67%) and Gramineae(56%), but not in the Umbelliferae
(5.9%) and Leguminosae(0%). A veryconsistent relationship is apparentbetweenseed
weightand theresponseto storageof seeds thatdid not attain50% germination as fresh
seeds.The decliningeffect of storagewithincreasing seed weightis maintained throughout
thefourclasses of seed weight:<0 1 mg,67%; 0 1-0-99 mg,38%; 1 00-9 99 mg,22%;
>9.99 mg, 0%. There are indicationsin the resultsthat dry storageexertedmaximal
effectsuponseedsreleasedfromlow (<20 cm) inflorescences (40%). The failureofstorage
to increasesubstantially thegerminability of seedswhichwerespherical(7.7%), subjectto
explosivedehiscence(10%) or eitheror bothmulticoloured and variablein colour(< 10%)
is correlatedwiththehighrepresentation oflegumesinthesecategories.
Several differences are apparentin the responsesof individualspecies to particular
storagetreatments. In the 5 IC treatment rapid ripeningand germination occurredin
certainannual species (e.g. Aira praecox, Draba muralis,Lapsana communisand
Solanumnigrum)and someperennials(e.g. Hypericum hirsutum, Rumexcrispus,Thlaspi
alpestreand Veronicaofficinalis), butwas comparatively slow in speciessuch as Ballota
nigra,Carexflacca,Plantagolanceolata,Senecioviscosusand Veronicapersica.
In Table 2, listsare providedforthreegroupsof specieswhichrespondedanomalously
to drystorage:eighteenspeciesin whichgerminability increasedat 5 IC but showedno
increaseat 20 IC; twenty-six species in whichthe effectof storagewas much more
pronouncedat 20 ?C than at 5 IC; and seventeenspeciesin whichgermination varied
erraticallybetweentests.Overhalfof thelast groupweremarshplants,whichare known
(K. Thompson,Grime& Mason 1977,AppendixI) eitherto germinate onlyat relatively
hightemperatures or to requirediurnalfluctuations in temperature (or both)to achieve
highgermination percentages.

Germination rate
In mostspecies and tests,germination was discontinuous (sensu Salisbury1942), i.e.
when the numbersof seeds germinating on successivedays is plotteda bell-shaped
distributionis obtained,withthe majorityof seeds germinatingwithina relativelyshort
period.Continuousgermination, involvingradicleemergencein a smallproportion ofthe
seeds each day over a longperiod,sometimes seeds
occurredin testson freshly-collected

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1028 Germination
characteristics
in a localflora
TABLE 2. Threegroupsofspecieswithunusualresponsesto drystorage.
Germination
percentage at5 IC butnotat20 0C
increasing
Annuals Perennials
Atriplexhastata Aquilegiavulgaris
Chenopodium album Bromusramosus
Galiumaparine Galega officinalis
Impatiensglandulifera Galiumpalustre
Odontitesverna Geraniumpratense
Polygonumlapathifolium Trifoliummedium
Polygonum persicaria Trifoliumpratense
Stellaria media Violapalustris
Viola arvensis Ulexeuropaeus
Germination
percentage
increasing
morerapidly
at20 ?C thanat5 ?C
Annuals Perennials
Aphanesarvensis Carex nigra
Erophilaverna Carex otrubae
Geraniumlucidum Carex sylvatica
Geraniummolle Centaureanigra
Juncusbufonius Glechomahederacea
Matricariamatricarioides Hieraciumpilosella
Moehringiatrinervia Polemoniumcaeruleum
Myosotisarvensis Poteriumsanguisorba
Saxifraga tridactylites Rumexacetosella
Spergulaarvensis Rumexsanguineus
Valerianellacarinata Serratulatinctoria
Veronicaarvensis Sieglingiadecumbens
Silene alba
Stellariapalustris
Germination
percentage
varying
erratically
during
storage
Annuals Perennials
Rorippaislandica Atropabelladonna
Brachypodium sylvaticum
Erica tetralix
Eriophorumangustifolium
Eupatoriumcannabinum
Filipendulaulmaria
Hypericumtetrapterum
Irispseudacorus
Juncusinflexus
Linaria vulgaris
Oenotheraparviflora
Pulicaria dysenterica
Ranunculusrepens
Scirpussylvaticus
Urticadioica
Verbascumthapsus

and oftencoincidedwithlow finalgermination percentage.Continuousgermination was


also associatedwiththe consistently low ratesof germination of manylegumesand of
certainspeciesof Geranium,in severalof whichcontinuousand morerapidgermination
was obtainable if the testa was scarified.A second group exhibitingcontinuous
germination includedspeciessuch as Potentillaerecta,P. tabernaemontani,
Ranunculus
acris, R. flammulaand R. repens,in whichmaturationof the embryooccurs during
incubationin moistwarmconditions.
In a verylarge numberof species,dry storageat 5 IC broughtabout a progressive
increasein speedofgermination. Althoughthiseffect ofseed-ripening
oftencoincidedwith

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J. P. GRIME et al. 1029
an increasein numberof seedsgerminating, itwas also observedin specieswhichattained
uniformly highgermination throughout thestorageperiod.Many of thespeciesin which
5 OC storagecaused an accelerationin germination rate showed an even more pro-
nouncedresponseto short-term storageat about 20 OC. This effectis wellexemplified
by some winterannuals (e.g. Cerastiumsemidecandrum, Erophila verna, Myosotis
ramosissima)and perennialforbssuchas Carex echinataand Serratulatinctoria.
Caution mustbe appliedin judgingthevalidityand significance of the comparatively
low (i.e. t50<4 days) ratesof germination observedin certainspecies.In plantssuch as
Calthapalustris,Coniummaculatumand Molinea caerulea,slow germination coincided
withverylow germination percentage, and almostcertainly reflected, at least in part,the
failureto providesuitablegermination conditionsor pre-treatment. Seventy-nine species
are identified in Appendix2, however,in whichconsistently low germination ratein the
dry-storedseeds is associated with high germination percentage.This group almost
certainlyincludes some species in which an invariablylow rate of germinationis
genetically determined; but forsome species,greaterratesof germination wereachieved
underconditionsdifferent fromthose appliedin the initialtests.In forty-three species
(twentyof them marsh plants) a greaterrate of germination was observedin the
temperature-gradient bar experiment (Appendix2). In fourteenspecies (includingnine
marshplants),theoptimaltemperature was higherthanthatused in theinitialtests,and in
fourspecies (threeof themwoodlandplants)maximumgermination rates occurredat
temperatures lowerthanthemeandailytemperature appliedintheinitialtests.
Thus it is apparent that the potentialgerminationrate of certain species was
under-estimated in the initialtests.Despitethislimitation, thetestsprovidea substantial
bodyof comparative data whichallowsa preliminary analysisof among-species variation
in germination rateand someecologicalinferences. In Fig. 3 thedistribution ofmaximum
germination rates in the initialtests in various classes of species is presented.The
distribution forall speciesis skewedto theleft,witha maximumt50at 3-4 days. All the
annualgrassesand a highproportion oftheannualforbsand perennialgrassesgerminated
rapidly.A widerangeofgermination ratesoccurredin theperennialforbs,buttherewas a
clearbias towardslow ratesin theshrubsand trees.Germination rateswerehighin many
Compositae,but low in all the Cyperaceae and many of the Leguminosae.The
distributions in Fig. 3 also provideseveralcorrelations betweengermination rateand field
distribution (Table 1, last twocolumns).Therewas a well-defined tendencyforspeciesof
northern distribution in theBritishIsles to germinate relatively slowly.Rapid germination
was characteristic of the majorityof the speciesof fertile disturbedhabitats.Marshland
and woodlandplantsincludeda highproportion ofslow-germinating species.
Severalcorrelations betweenseed characteristics and germination rate(t50)are apparent
in Table 1 (lasttwocolumns).Withincreasing seed weightthereis a progressive declinein
representation of rapidly-germinating species. Slow germination was common among
species in which the seeds were spherical,multicoloured,variable in colour, or
mucilaginous.Rapid germination was correlatedwiththe presenceof a pappus,conical
seed shape (Compositae),hygroscopicawns (Gramineae)and antrorsehairs or teeth
(Compositaeand Gramineae).
From a previousinvestigation (Grime& Hunt 1975), estimatesare availableforthe
maximumrateofdry-matter production(Rmax)duringtheseedlingphaseofninety-nine of
thespeciesinvestigated here.In Fig. 4 thesespecieshavebeenclassifiedaccordingto Rmax,
and theclasses have beencomparedwithrespectto theproportion ofrapidly-germinating
species.A veryclearpositivecorrelation is evidentbetweenrateofgermination and rateof

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1030 Germination in a localflora
characteristics

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J. P. GRIME et al. 1031

24 ElIt50>4 days
24 -501I 4 daYS
20 _
~~~~ 16 ~ ~ ~ ~ -4dy
20
16 _ ........
....

12
? ..,,,,,
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076 101-125 151-175


076 100 126-150 >175
Rmax(week-l)

FIG. 4. The contribution (t50? 1- days) and slow-germinating


of rapidly-germinating (t50> 4
days) species to groupsclassifiedwithrespectto estimatesof the maximumpotentialrate of
seedlingdry-matterproduction(Rmax).

seedlingdry-matter production.
It is notunreasonableto proposethatin manyspeciesthe
two attributeswillshowparallelresponsesto naturalselection,whichin some cases may
be linkedphysiologically.Need forcautionis suggested,however,by species such as
Helianthemumchamaecistus,Lotus corniculatusand severalothertap-rooted forbsof
infertile
soils,in whichthe capacityof scarifiedseeds to germinaterapidlyis associated
withan inherently lowrateofseedlinggrowth.

Dormancy
Of the403 speciesexaminedin theinitialtests,137 failedto attaina finalgermination
percentageof 50%, and in many other species germinationwas incomplete.Low
germination in some specieswas due to unusuallightand temperature requirements.In
Urticaurens,forexample,the resultsof the light-flux testrevealthatgermination was
almostcompletelyinhibitedby the irradianceused in the initialtests.In Ballota nigra,
Bidenstripartita, Carex capillaris,C. demissa,C. nigra,Moliniacaerulea,Nardusstricta,
Thiaspi arvense and Verbascumvirgatum,low germination percentagesin the initial
testsmay have been relatedto a requirement forratherhightemperature. Seven of the
species exhibiting low germination (Bidens tripartita,Linaria vulgaris, Lycopus
europaeus,Mentha arvensis,Molinia caerulea,SonchusarvensisandStachyssylvatica)
are capable of responding in thelightto amplitudesofdiurnaltemperature changegreater
thanthe5 ?C fluctuation usedintheinitialtests(K. Thompson1977).
The presentinvestigation was notdesignedto providea thoroughanalysisofthecauses
of seed dormancyin each species.Seeds fromthesame populationofplantsor evenfrom
the same individualmay differ in germinationrequirements, and in orderto investigate
theirresponsesfullyit would be necessaryto apply various proceduresin factorial
combination to seeds subjectedto a rangeof pretreatments (Steinbauer& Grigsby1957;
Toole 1973; Vincent& Roberts 1977; Bostock 1978; P. A. Thompson1981). Several
procedureswereapplied,however,in an attemptto stimulate germinationin manyofthe

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1032 Germination in a localflora
characteristics
specieswhichfailedto attainhighpercentagesin theinitialtests,and a summaryof the
resultsis presented in Appendix1.
Thereis evidenceof a positiveresponseto chillingin 75 ofthe 107 speciestested.The
listincludes7 woodyplants,16 woodlandherbs,Umbelliferae and 3 annualhemiparasites
(Euphrasia officinalis, Odontitesverna,Rhinanthusminor).Of the threegrasseswhich
respondedto chilling, one was a woodlandspecies(Bromusramosus),whilsttheothertwo
(Molinia caerulea and Nardusstricta)had seedswhichbeforechilling germinated onlyat
relativelyhightemperature. A patternof responsesimilarto thatobservedin Molinia
caerulea and Nardus strictawas recognizablealso in Carex nigra,Mentha arvensis,
Sonchus arvensis, Stachys sylvatica and Solanum dulcamara, in each of which
germination priorto chillingwas dependent uponexposureto hightemperature or to large
diurnalfluctuations intemperature.
Amongthespecieswhichrespondedto chilling, thereweremarkeddifferences notonly
withrespectto thelengthoftreatment requiredbutalso to thecapacityforgermination at
the chillingtemperature (5 IC). At one extremewerespeciesin whichsatisfaction of the
chillingrequirement was associated with the capacity for rapid germinationat low
temperature. This groupcontained10 speciesin the Umbelliferae, and includeda high
proportion ofspecieswithcomparatively largeseeds.
In 24 speciesno germination was observedduringthe chillingtreatment, but a high
percentageoftheseedsgerminated followingtransferto lightand highertemperature. This
group of plants includedarable weeds and marshplants withrelativelysmall seeds.
Scarificationwas applied to 27 species, and resultedin a markedstimulationof
germination in 19 species(12 legumes).

Responseto temperature
The need forcautionin extrapolation fromlaboratoryto fieldis especiallyclear with
respectto theresultsofthetemperature gradientbar experiments.In orderto summarizea
largevolumeof results,thegermination responsehas beendescribedmainlyby reference
to the upper and lowerlimitsof the rangeof temperatures over whichhigh(>50% of
maximum)germination occurred.This formof presentation
percentages obscuresthefact
thatthe percentagegermination was less outsidethis range.It shouldbe remembered
also thattheresultsareforconstanttemperature, and thatinmanyspeciesthegermination
range(particularly at itslowerextremity) maybe extendedunderfluctuating temperature
regimes.
Despitethesequalificationsthedata showseveralconsistent patternsofpredictive value.
Resultsfor272 speciesare presentedin Appendix2. The majorityof resultswerefor
seeds testedafterdrystoragein thelaboratory.In some speciesthemeasurements were
conductedon pre-chilledor scarifiedseeds, and in two species (Ranunculusacris, R.
repens)seedswereincubatedindarknessat 20 IC priorto use intheexperiment.
In 16 of the species investigated,germination did not occur or was restricted to a
verysmallnumberof seeds.Undersimilarlightflux,all thesespecieshad attainedhigher
percentagesin the initialtests,and it seems likelythat failureto germinateon the
temperature-gradient barswas relatedto a requirementforfluctuatingtemperature.
Among the 256 species which germinatedon the bars, there were considerable
differences inresponseto temperature,someofwhichare summarized in Fig. 5.
In Table 3 comparisonsof temperature-response have been drawnbetweenvarious
classes of plants.Amongannualand perennialgrassesa highproportion of specieswas
capable of germinating over a wide (>20 OC) range of temperature. This featureis

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J. P. GRIME et al. 1033
(a) Lower limit ( b) Upper limit ( c) Range

80

60-

0
40

E
z olL
20-
L-|V 1 --1 1

<5 8-12 21-33 19-22 26-28 32-34 38-40 4 8 12 16 20 24 28 32 36


5-7 13-20 >33 23-25 29-31 35-37
Temperature(?C) Temperaturerange(?C)
FIG. 5. Distributionsof (a) upper and (b) lower limitsof temperature and of (c) range of
temperature overwhich50% maximumgermination was attainedin 256 species.The classes in
(a) are notofequal width.

particularlyevidentin speciesof dryhabitatsand of southerndistribution,and suggests


to temperature
thatrelativeinsensitivity of seeds in whichwatersupply
is characteristic
acts as theprimarydeterminant ofthetimingofgermination inthefield.
Several sources of variationin temperature-response appearedto be associatedwith
taxonomicdifferences. Scarifiedseeds of legumestendedto germinateover a verywide
rangeof temperatures. In boththe Leguminosaeand the Gramineae,wide germination
rangewas relatedto thecapacityforgermination butin the
at highand low temperature,
Compositae wide germination range was mainlydue to high germinability at higher

TABLE 3. Number ofspecies,grouped invariousclasseswithrespectto therange


of constanttemperature overwhichgermination in lightexceeds50% of that
attainedat theoptimaltemperature, and of germination percentagein thedark
treatmentof Experiment 1. The significance levelsreferto theprobabilitywith
whichtheratiofora particular class departsfromtheratioexpectedfromthe
obtained
results forall species.Significanceconventions as inTable1. Superscript
's' = scarified.
Dark treatment
of
Temperature Experiment1;
Lower limit Upper limit Range (IC) germination
?7 IC >7 OC <28 ?C >28?C <20 >20 <50% >50%

All species 97 159 68 188 112 144 161 110

Life-form
Annual grasses 7 ** 1 1 7 0 * 8 0 ** 8
Annual forbs 27 28 21 * 34 24 31 34 21
Perennialgrasses 20 * 15 5 30 7 ** 28 13 ** 25
Perennialforbs 41 ** 109 37 113 76 * 74 110 ** 48
Shrubsand trees 2 6 4 4 5 3 4 8

Family
Compositae 11 35 5 * 41 16 30 33 * 11
Cyperaceae 0 * 12 2 10 11 1 12 ** 0
Gramineae 27 * 16 6 37 7 ** 36 13 ** 33
Leguminosaes 13 * 0 2 11 0 ** 13 0 * 16

Britishdistribution
Northern 4 ** 27 11 20 22 ** 9 20 5
Southern 34 66 20 80 40 60 64 33
Ubiquitous 43 * 46 25 64 33 56 56 48
Widespread 12 13 8 17 10 15 15 16
Local 4 7 4 7 7 * 4 6 6

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1034 Germination in a localflora
characteristics
TABLE 3-continued
Dark treatmentof
Temperature Experiment1;
Lower limit Upper limit Range (IC) germination
?7 IC >7 OC ?28 OC >28 OC ?20 >20 <50% >50%

Frequencyin Sheffield
flora
> 10.0% 10 ** 4 2 12 2 * 12 3 * 11
1.1-10.0% 38 53 29 62 38 53 53 48
0. 1-1.0% 33 76 26 83 51 58 76 40
<0.1% 14 28 11 31 21 21 29 * 11

Habitat
Disturbed 21 38 13 46 24 35 44 17
Skeletal 23 25 18 30 17 31 20 26
Marshland 11 ** 44 5 ** 50 31 * 24 49 I* 11
Grassland 37 41 21 57 28 50 32 ** 48
Woodland 5 11 11 ** 5 12 ** 4 16 8

Heightof inflorescence
(cm)
0-20 34 * 34 25 43 30 38 32 33
21-40 29 44 16 57 31 42 46 34
41-60 15 38 15 38 27 26 30 22
68-80 10 20 8 22 12 18 26 * 9
>80 9 23 4 28 12 20 27 12

Dehiscence
Passive 88 156 64 180 109 135 160 94
Explosive 9 3 4 8 3 9 1 16

Dispersuleshape
Spherical 13 14 11 16 13 14 14 17
Ovoid, rhomboidalor turbinate 19 26 11 34 15 30 35 18
Trigonousor triquetrous 7 14 7 14 12 * 9 18 * 5
Lenticular,reniform
or subulate 31 33 25 * 39 35 * 29 36 35
Elongated 16 38 6 * 48 20 34 27 24
Conical 12 33 8 37 17 28 31 * 11

Dispersuleappendages
Absent 66 104 53 117 79 91 109 80
Straightawn(s) or spine(s) only 10 9 2 17 8 11 10 9
Hygroscopicawn or spine 8 ** 3 3 8 2 9 1 * 8
Pappus or persistentcalyx 10 * 37 7 40 18 29 33 * 10

Dispersulehairsor teeth
Absentor veryinconspicuous 63 122 52 133 89 96 132 74
Radial orirregular 9 * 5 8 * 6 3 11 3 * 8
Antrorse 25 32 7 * 50 20 37 26 28

Germinuleweight(mg)
<0-10 18 40 16 42 26 32 38 19
0.10-099 53 81 35 99 60 74 83 55
1.00-9-99 22 37 14 45 25 34 37 31
>9-99 4 - 1 3 - 2 1 - 4 2 - 5

Germinulesurfacetexture
Smooth 43 79 33 89 56 * 66 87 54
Rugose, tuberculate,muricateor
reticulate 27 31 16 42 27 31 24 33
Striate 11 20 8 23 8 23 20 9
Hairy 8 7 4 11 4 11 5 7
Striateand hairy 6 14 3 17 10 10 14 4
Mucilaginous 2 8 4 - 6 7 * 3 11 3

Germinulecolour (Munsellhue)
5R, 7.5R and IOR 15 24 18 ** 21 27 ** 12 26 21
2-5YR 13 16 4 25 9 20 21 9
5YR 16 * 52 18 50 36 32 42 26
7-5YR 11 28 11 28 20 19 28 * 12
1OYR, 25Y and 5Y 22 * 16 7 31 7 ** 31 22 16
Black 2 - 4 2 - 4 3 - 3 5 - 2
Multicoloured 8 7
Variable 9 11
Multicolouredand variable 18 19 8 29 10 27 0 - 6

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J. P. GRIME et al. 1035
temperature. A quitedistinct patternof responsewas characteristic oftheCyperaceae,in
whichgermination was restrictedto a narrowrangeofrelatively hightemperatures.
Whengroupsof speciesbelongingto variousgeographicalelementswithintheBritish
florawerecompared,severalfeatureswereapparent.The moststriking of thesewas the
failureof the majorityof plantsof northerndistribution to germinateat low (<7 IC)
temperature. This resultis in agreement withearlierstudies(Billings& Mooney 1968;
P. A. Thompson 1968, 1970b; Wein & Maclean 1973), and is consistentwith the
hypothesis thatthe hightemperature-requirement in theseplantsis theresultof natural
selectionat high latitudesfor a germination-response restrictingthe appearance of
seedlingsto theshortbutrelatively favourable summer.
Species concentratedin southernBritainincluded a high proportioncapable of
germinating at low temperature, and tendedto have a temperature rangeexceedingthatof
thenorthern species.Speciesgerminating overa widerangeof temperature reachedtheir
highestfrequencyamong plants which are ubiquitousin the BritishIsles. A wide
temperature rangeforgermination of plantsoccurring
is also characteristic in thehighest
frequency-class (> 10% ofrecords)intheSheffield flora(Table 3).
Relationshipsbetweentemperature responseand habitatare suggestedby the results.
Marshlandand disturbedhabitatsaccountedfora largeproportion of the specieswhich
did not germinateat low (<70C) temperature. Some of thesespeciesare also unusually
responsiveto diurnalfluctuations in temperature (K. Thompson 1977). In temperate
localities,seedlingestablishment in marshesand at themarginsoflakes,rivers,pondsand
ditchesis usuallyrestricted to thelatespring, whenthewater-table recedesand thesurface
layersofmudbecameexposedand aerobic.It seemslikely,therefore, thattherequirement
forhighor fluctuating temperature in thesemarshplantsprovidesa mechanismdelaying
springgermination untilsuchtimeas thedecliningwater-table no longerinsulatesthesoil
fromtheincreasing radiationload (K. Thompson,Grime& Mason 1977).
Germination over a wide rangeof temperature was particularly commonin grassland
plantsand speciesfromskeletalhabitats.In thewoodlandplantsinvestigated, by contrast,
germination was oftenrestricted to a narrowrangeofintermediate temperatures.

Responseto light

Experiment 1
Figure6 describesthedistribution of germinationpercentagesattainedby 271 species
in the 'light','shade' and 'dark' treatments.In both'light'and 'shade', percentageswere
highin themajorityofthespecies,althoughtherewas a tendency forgermination percent-
age and rate to be slightlylowerin the 'light'.Germination was muchreducedby the
'dark'treatment, and failedto exceed10% in 104 species.
Despitethesegeneraltrends,interspecific differencesoccurredin responseto the'light'
treatments. At one extremewere speciessuch as Lycopuseuropaeus,Menthaarvensis,
Polemoniumcaeruleum,and Urticaurensin whichgermination percentagesin the'light'
fellbelowthosein 'shade',whilst,at theother,wereplantssuchas Bidenstripartita, Carex
flacca, Juncusconglomeratus and Scirpussylvaticus,wheregermination in 'shade' was
inferior to thatinthe'light'.
These resultsconfirmthose of the comprehensive screeningexperimentof Kinzel
(1920), whofoundthatplantsdiffered considerablyin theirgermination responsesto light.
In Kinzel's investigation, as in our study,germination in the majorityof species was
promotedin the light.In both studiestherewas a smallernumberof species in which

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1036 Germination in a localflora
characteristics
140 - (a) Light

120 -

100 _

80 _

60 -

40-

20 - -
---
0 -

160 (b) Shade


a 140 -

o 120 -
-E 100_

Z 80 _
60 -

40 -
20-
0

(c) Dark
60-
40 [-7
20 - ___ _

0 1-10 21-30 41-50 61-70 81-90


11-20 31-40 51-60 71-80 91-100
Germination (/)

withrespectto germination
FIG. 6. Distribution of 271 speciessubjectedto (a) 'light'(b) 'shade'
For detailsoftreatments
and (c) 'dark'treatments. see text.

germination in thedarkwas superiorto thatinthelight.The inhibitory effects oflightwere


less pronouncedin our study,however,and thiscan be relatedto thelowerlightfluxand
smallerproportionof far-redradiationin our experiments comparedwiththe natural
daylightusedin Kinzel'sinvestigation.
The tendencyof certainspecies to be inhibitedfromgerminating in 'light'has been
observedin numerousspeciesin earlierinvestigations (Toole 1973).The inhibitory is
effect
mostsevereunderprolongedhighlightflux(Black & Wareing1960; Schulz& Klein 1963;
Chen & Thimann1964; Rollin& Maignan 1967; Kendrick& Frankland1969) and also
at hightemperature (Black & Wareing1960; Chen & Thimann1964). This suggeststhat
underfieldconditionsthe inhibitory effectsof highlightfluxwillusuallycoincidewith
stronglydesiccatingconditions,whichmaypose a severethreatto seedlingestablishment.
It seems possible,therefore, to high irradianceprovidesa safeguard
that sensitivity
reducingtheriskofseedlingdeathcausedbydrought.
Strikingdifferences in responseoccurredin the 'dark' treatment (Table 4, last two
columns).The proportionof speciesexceeding50% germination in the'dark' treatment
was considerably higherin thegrassesthanitwas in theannualand perennialforbs.After
all of theLeguminosaegerminated
scarification, rapidlyto a highpercentagein the'dark'

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J. P. GRIME et al. 1037
treatment, in markedcontrastwiththe Cyperaceae and Compositae.Species withthe
capacityto germinate to a highpercentage in the'dark'treatmentwereespeciallycommon
amongplantswhichare ubiquitousor of widespreadoccurrencein the BritishIsles, and
also formeda highproportion ofthespeciesof commonoccurrencein theSheffield flora.
Whenthespecieswerebroadlyclassified withrespectto habitat,theycouldbe arrangedin
a seriesthroughwhichtheproportion of speciesattaining50% germinationin darkness
declines as follows: grassland> skeletal habitats> woodland> disturbedhabitats>
marshland.
No consistentrelationship was detectedbetweenthecapacityof seeds to germinate in
the 'dark' treatmentand theiraverage heightof release. With one exception(Viola
palustris),species with explosivedehiscence(mainlylegumes) germinatedto a high
percentagein darkness.Small-seededspecies (<0.1 mg) includedmanywhichfailedto
germinate in darkness,and theincidenceof specieswitha dependenceuponlightdeclined
progressivelywithincreasing seed size.

Experiment2
The resultsof the secondexperimentclarifythoseobtainedin the'dark' treatment
of
Experiment1.
Whengermination in thetwodarktreatments ofExperiment2 was comparedwiththat
measuredin Experiment1, a varietyof responseswas apparentamongthe 70 species
examined.In Table 4 fivemaintypesof responsehave been recognized,and thesehave
beenidentified
as Types(a)-(e).
TABLE 4. Theeffectofa diurnalfluctuation
intemperature andbrief dailyexposure
to greenlightuponthegermination indarknessofselected species.In eachcolumn
germination is expressed of thatrecorded
as a percentage in a concurrent test
conducted levelsrefer
inthelightat 20/15IC; thesignificance tothiscomparison.
conventions
Significance 's' = scarified,'c'
as inTable1.Superscripts: = chilled.
Response
type
1 Experiment
Experiment 2 2 (see text)
Experiment
Exposureto green'safety'light + - -
Temperature regime(OC) 20/15 20/15 20/20
Annualgrasses
Airapraecox 91** 97 98 b
Bromuspseudothominii 100 101 100 b
Bromussterilis 100 100 100 b
Catapodiumrigidum 94 102 99 b
Hordeummurinum 100 98 100 b
Vulpiabromoides 103 104 104 b
Annualforbs
Aphanesarvensis 29*** 12*** 2*** c
Arabidopsisthaliana 23*** 3*** 0*** c
Atriplexpatulac 57*** 41*** 26*** d
Blackstoniaperfoliata 16*** 0*** 0*** c
Cerastiumsemidecandrum 98 65*** 56*** d
Chaenorrhinum minusc 59*** 66*** 67*** b
Chenopodiumalbum 8*** 8*** 3*** a
Draba muralis 95 43*** 31*** d
Galiumaparine 91 83 31*** e
Lapsana communis 57*** 0*** 7*** c
Medicago lupulinas 100 99 99 b
Moehringiatrinervia 82*** 53*** 56*** d

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1038 Germination in a localflora
characteristics
TABLE 4-continued
Response
type
1
Experiment Experiment
2 Experiment
2 (see text)
Papaver somniferum 48*** 66*** 16*** e
Saxifraga tridactylites 35*** 0*** 0*** c
Trifolium arvenses 102 102 104 b
Valerianellacarinata 93 2*** 0*** c
Veronicapersica 67*** 12*** 0*** c
Vicia sativa spp. angustifolium' 100 100 98 b

Perennialgrasses
Agrostiscanina spp. canina 102 75** 78** d
Agrostistenuis 71** 68** 36** e
Alopecuruspratensis 98 100 91 b
Arrhenatherum elatius 98 100 100 b
Cynosuruscristatus 103 98 96 b
Loliumperenne 98 102 100 b
Miliumeffusum 34*** 7*** 16*** d
Phalaris arundinacea 83* 33*** 21*** d
Phleumpratenseagg. 50*** 20*** 32*** d
Poa compressa 94*** 100 57*** e
Poa pratensisssp. angustifolia 104 56*** 56*** d
Poa pratensisssp.pratensis 75*** 102 5*** e
Poa trivialis 79 73 18*** e

Perennialforbs
Artemisiavulgaris 44*** 14*** 2*** c
Campanula rotundifolia 25*** 4*** 2*** c
Carex nigrac 9*** 0*** 0*** a
Cerastiumfontanum 95 12*** 0*** c
Chrysanthemum leucanthemum 83*** 44*** 38*** d
Epilobiumadenocaulon 84* 3*** 0*** c
Filipendulaulmaria 97 53*** 13*** d, e
Galiumverum 98 48*** 67** d
Hieraciumpilosella 91** 45*** 43*** d
Hypericumhirsutum 12*** 0*** 0*** a
Hypericummontanum 98 0*** 0*** c
Lathyrusmontanus' 98 88* 92 b
Leontodonautumnalis 22*** 0*** 0*** c
Leontodonhispidus 7*** 0*** 16*** a
Lotus uliginosus5 96 98 96 b
Luzula multiflora 102 92 82** b
Melilotusaltissima' 100 104 104 b
Minuartiaverna 87 88 51*** e
Myosotissylvatica 78*** 71*** 60*** e
Plantagomedia 10*** II*** 5*** a
Reseda luteola 16*** 24*** 16*** a
Rumexacetosa 92* 100 91 b
Rumexhydrolapathum 12*** 8*** 1*** a
Sedumacre 56*** 0*** 0*** c
Silene dioica 89* 60*** 34*** d, e
Silene nutans 97 84** 78*** b
Trifolium pratenses 102 98 94 b
Trifolium repenss 100 100 89 b
Veronicachamaedrys 60*** 0*** 0*** c
Veronicaofficinalis 11*** 2*** 0*** a

Shrubs
Thymusdrucei 78*** 23*** 19*** d
Vacciniummyrtillus 102 0*** 0*** c

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J. P. GRIME et al. 1039
In Type (a), representedby ninespecies,germination remaineduniformly low in all the
dark treatments, indicatinglow responsivenessto both green light and fluctuating
temperature. Type (b) specieshave a highpercentagegermination in all treatments, and
includethe annual grasses and the legumes,togetherwithfourperennialgrasses(Alo-
pecuruspratensis,Arrhenatherum elatius,Cynosuruscristatusand Loliumperenne)and
threeperennialforbs(Luzula multiflora, Rumex acetosa and Silene nutans).It seems
reasonableto conclude that in these species no evidenceof a lightrequirement was
obtained.Response-type(c) was that in which considerablegermination in the dark
treatment of Experiment1 was associatedwitha value at or near to zero in bothdark
treatments of Experiment 2. This effectwas clearlyrelatedto theexclusionof greenlight,
and was observedin seven annual forbs,nine perennialforbs and one undershrub
(Vacciniummyrtillus). A similarbutless dramaticeffect oflight(Type (d)) was evidentin
15 species,wherea proportionof the seeds continuedto germinatein the absence of
greenlight.This typeof response,indicating withinthe seed populationin
heterogeneity
respectto lightrequirement, classesexcepttheannualgrasses.
occurredin all life-form
In a fifthpatternof response(Type (e)) dark germination was unaffected by the
exclusionof greenlight,but declinedmarkedlyat constanttemperature. This dependence
of germination in darknessupon diurnalfluctuations in temperature was apparentin nine
species,two of which(Filipendulaulmariaand Silene dioica) also showedevidenceof a
responseto exposureto greenlight.
The resultsconfirm theneedforcautionin interpreting theresultsobtainedin thedark
treatment of Experiment 1, and suggestthatin orderto detectall thespecieswhichmaybe
incapableof germination in darknessit is necessaryto deviseexperimental treatments
whichexcludebothvisibleradiationand fluctuations intemperature.
The capacityof the seeds of certainspeciesto germinate in darknessin responseto
diurnalfluctuations in temperature appears to be relevantto an understanding of the
mechanismsof depth-sensing and gap-detectionby buried seeds. Measurementsin
grassland (K. Thompson,Grime & Mason 1977) and scrub (Panetta 1979) have
confirmed that the amplitudeof diurnalfluctuations in temperature withinthe soil is
greatestnear to the soil surfaceand in local-situationswhereremovalof the insulating
layersoffoliageand litteradmitsa higherfluxofradiation.

DISCUSSION

The measurements in each rowofAppendix2 referto seedsfroma singlepopulation,and


furtherstudiesare requiredin orderto establishthe e%tentto whichthese resultsare
forthe species.This appliesparticularly
definitive withrespectto responsesto lightand
temperature,as it has been established(Koller 1962, McCullough& Shropshire1970;
1973; Gutterman1974; Cresswell& Grime1981) thatthesemaybe influenced
Junttila by
theconditionsexperienced duringseedmaturation.
The experimentalconditionsused throughoutthis study were extremelysimple
comparedto the complexfluctuations of manyfactorsexperiencedby seeds in natural
environments.The screening approachadoptedrestsuponthreeassumptions:(a) thatlittle
maybe achievedby transferring to thelaboratorycomplexities whichhave alreadydefied
analysisin the field;(b) thatthe prospectsforreliableextrapolationto thefielddepend
upon examinationof an adequate range of species and recognitionof the causes of
response,ratherthan the use of experimentalconditionswhich closely
differential
approximate to thoseoccurring ofecologicalimportance
in nature;and (c) thatdifferences

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1040 Germination
characteristics
in a localflora
are oftendetectablein the responsesof groups of plants of contrastedecology to
standardized experimental conditions.
The results in this paper reveal numerousinstances where the same group of
seed-characteristicsrecurs in association with species of similar ecology, and the
information has alreadybeenused (Grime1979) to considertheroleofmorphological and
physiologicalattributesof seedsin particular
regenerative
strategies.
The mostsatisfactory
analysis,however,is thatin whichthelaboratoryresultsare complemented by studiesof
theproductionand fateof seedsundernaturalconditions. This approachis adoptedin the
sequelto thispaper(Grime& Hillier,unpublished) in whichseed-bankmodelsare used to
assess thefunctionalsignificanceofsomeoftherecurrent groupingsofseed-characteristics
emerging fromthepresentstudy.As a preliminary to thisanalysis,theremainder of this
discussionconsiderssomeproblemsof interpretation associatedwithparticularaspectsof
thedata.

Capacityforimmediate germination
Withtheexceptionof speciessuch as Petasiteshybridus and Tussilagofarfara,whose
seeds are exceptionallyshort-lived,it would be unwiseto assume that the abilityof
freshly-collected
seedsto germinateinthelaboratory is a reliableindication
thatunderfield
conditionsgermination occurssoon afterseed release.Many ofthespecieswhichshowed
the capacity for immediategerminationhad other characteristics(e.g. specialized
temperature requirements)whichsuggestthatgermination offreshly-dispersedseedsmay
be prevented by theintervention
oflimitingfactorsoperating inthefieldbutexcludedfrom
the initialgerminationtest.This hypothesisis supportedby the factthatmanyof the
species whichdisplayedhighinitialgerminability are knownto accumulatelarge and
persistentburiedseedbanksinthefield.

Responsetodrystorage
Beneficialeffectsof dxy storageupon germination percentageand rate were most
conspicuous among winterannuals of shallow or sandy soils (e.g. Aira praecox,
Arabidopsis thaliana, Draba muralis, Erophila verna, Myosotis ramosissimaand
Saxifraga tridactylites).Seeds of thesespeciesare shed duringthe earlysummer.From
field studies (Ratcliffe1961; Newman 1963) it seems likelythat the after-ripening
functions as a mechanismpreventing premature germination in thedryhabitatsexploited
by these plants. The same explanationmay be applied to the characteristic but less
pronouncedresponsesto dry storage evidentin certainof the autumn-germinating
perennialgrassessuchas Festucaovina,KoeleriacristataandPoa compressa.
Of theperennialspecieswhichdisplayeda markedimprovement in germinability
during
drystorage,the majorityare small-seededand manyare knownto formpersistent seed
banks in the soil. The possibility
mustbe considered, therefore, thatin certainspeciesa
majoreffect ofdelayedripening and germinationis to facilitate
seedburial.

Germination rate
Success in seedlingestablishment almostinvariablydependsuponrapidexploitation of
temporarily favourableconditions, and theprospectsforsurvivaland reproduction during
laterstagesofthelife-historymaybe strongly affectedbythespeedofgermination and the
rateof seedlingdevelopment. Thus itis tempting
to proposethatall specieswillbe capable
of rapid germinationin their natural environments, and that where low rates of
germination were observedin the presentstudytheyarose fromthe failureto provide

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J. P. GRIME et al. 1041
appropriategerminationconditions or pretreatments.The ratesrecordedin theinitialtests
and in most of the studiesof responseto temperature and lightwere based upon
experiments in which germination was initiatedby hydrationof air-driedseeds. This
procedurereproducesthe common circumstancein which the germination of many
grasses,membersoftheCompositaeand winterannualsat theendofthesummercoincides
withtheonsetof wet conditions, but it is scareclyrelevantto thevarietyof instancesin
whichfully-imbibed seeds are inducedto germinateby environmental stimulisuch as
changesintemperature or illumination.
The ratesof germination reportedin thispapermaythusproveto be of doubtful value
as an indexofthegermination ratesachievedinthefield.

Responsetochilling
The resultssuggestthatthespecieswhichrespondedto chillingtreatment fallintotwo
categories,in the firstof whichthe seeds are comparatively large and afterchillingare
capable of rapidgermination at low temperature and in darkness.This groupincludesa
varietyof plantsof widespreadoccurrencein northern Europe.In Britainthemajorityof
thesespeciesare knownto producelargeeven-agedpopulationsof seedlingsin theearly
spring.Thereis no evidenceto suggestthatany of theseplantsmaintainpersistent seed
banksin thesoil,and themostprobableexplantion forthispeculiargermination biologyis
thatit allows springcolonizationof gaps in perennialvegetation. It seemslikelythatthe
largeseeds and capacityforearlygermination allow seedlingsofthesespeciesto compete
effectivelywithneighbouring perennials.
established
A differentsignificancemay be attachedto thechillingresponsesnotedin someof the
smaller-seeded arableweedsand marshplants.Heretherequirement forchillingis alliedto
a need forsubsequentexposureto lightor highertemperature or both.As severalof the
speciesin thissecond categoryare knownto formreservesof buriedseeds (Brenchley
1918; Chippendale& Milton1934; Champness& Morris1948),we may suspectthatthe
chillingrequirementsareinvolvedin mechanisms ofdelayedgermination and seedburial.

Responsetoscarification
Althoughthe role of impermeable seed coats underfieldconditionsis stilluncertain
(Ballard 1973), therecan be littledoubt that this characteristic is oftenconduciveto
delayedgermination and incorporation intopersistent seed banks.A further insightinto
thepossibleecologicalsignificanceofthehardseedcoatsoflegumesis providedby studies
such as thoseof Hamly (1932) and Hagon & Ballard (1969) and Hagon (1971), which
have shownthatpermeability dependsupontheconditionofthestrophiole, theonlypoint
of entryof waterintotheunscarified seedsof manylegumes.More suggestive stillare the
resultsof Quinlivan (1961, 1968) and Quinlivan& Millington(1962), which have
establishedthatin someAustralianlegumespermeability ofthestrophiole can be increased
underfieldconditionsby diurnaltemperature fluctuations of the magnitudecommonly
experienced by buriedseedslyingon or close to a soil surfacedevoidofvegetation. From
these sources, therefore, there is circumstantialevidenceto suggest that seed-coat
impermeability in thelegumes,and perhapsalso in otherspeciesexaminedin thisstudy,
thepersistence
facilitates of seedreservesin thehabitatand mayalso providea mechanism
of gap-detection(K. Thompson,Grime& Mason 1977; Grime1979) by surface-lying or
buriedseeds.

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1042 Germination
characteristics
in a localflora
Responseto temperature
The time-course of the germination responseto temperature in each specieshas been
describedinAppendix2 byreference to theclassification
ofcurve-types illustrated
in Fig. 1.
The most commontype of curve (Type I) was that in whichthe upper limitof the
temperature rangebecame constantsoon afterthe commencement of germination, and
therewas a progressiveextension,withtime,of the lower limit.Type I curves were
particularlycharacteristicof grasses and legumes.In the second most commonly-
occurring typeof response(Type II) boththeupperand lowerlimitsof thecurvebecame
fixedaftertheinitialburstof germination. This compactformof responseoftencoincided
withgermination over a narrowrange of relativelyhightemperatures, and was more
frequent in forbsthanin grasses.It wouldappear(K. Thompson1977) thatin manyofthe
speciesexhibiting thisresponse,thelowertemperature limitmay be extendedby exposure
to diurnalfluctuations in temperature. Althoughfurther workis requiredto clarifythe
significance of theType II responseto temperature, we suspectthatin manyspeciesit is
associated withthe capacityto formpersistentseed banks and with a tendencyfor
germination to occurinthespringand earlysummer.
The temperatureresponses classifiedas Types III and IV, togetherwith two
intermediate types(TypesI-Ill, 11-111), are characterized
by delayedgermination ofsome
seeds at moderatetemperature. The group of species in whichthis pehnomenonwas
observedis ratherheterogeneous. It includesmanymembersof the Compositaeand a
numberof speciesin whichtheseed is largeor has a surfacetexturelikelyto impairthe
conductionof waterfromthe surfaceof the temperature-gradient bar to the seed. In a
numberof elegantexperimental studies(Harper et al. 1965; Harper & Benton 1966;
Sheldon1974; Peart 1979), it has been shownthatthecontactestablishedbetweenseed
and substrateis oftena criticalfactordetermining imbibitionand germination. Thus it is
notimmediately obviouswhysome seeds shouldhave a structure whichimpairsmoisture
uptake.However,underwarmsummerconditionsrapidevaporationofmoisture fromthe
soil surfaceis a severehazardto thoseseedlingswhichappearin responseto temporary
moistening of thesoil,and a selectiveadvantagemay be expectedformechanisms which
restrictgermination to circumstanceswheremore constantmoisturesupplyor lower
evaporative losses,or both,arepropitious forseedlingsurvival.

Responseto light
The presenceof a light-requirementin thefreshly-collected seedsofmanyofthespecies
includedin thisinvestigation poses severalinteresting problems.Why is thisbehaviour
foundin some speciesbut not in others?In manyspeciesthedifference appliesto seeds
withinthe same seed collection,and it is known(Cavers & Harper 1966) that seeds
removedfromthesameinflorescence mayexhibitmarkeddifferences in light-requirement.
It is also necessaryto considertheinfluencewhicha light-requirement mayexertuponthe
subsequentfateoftheseed.
Since the studiesof Flint& McAlister(1935, 1937) and Borthwicket al. (1952), it
has been establishedthat the responsivenessof seeds to changes in illumination is
mediatedbythepresencein theseed ofphytochrome (P), a compoundwhichexistsintwo
interconvertible forms,one of which(P650)absorbslightin theredregionwhilsttheother
(P730) absorbsin thefar-redregion.Providedthatotherfactorssuch as watersupplyand
temperature are notlimiting,
germination appearsto be inducedwhentheconcentration of
P730 in the seed rises above a criticalthreshold. In these circumstancesgerminability

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J. P. GRIME et al. 1043
depends upon the ratio of red:far-redin the lightimpingingupon the seed. From
laboratorystudiesand experiments withnaturalleafcanopies(Pigott1971; Gorski1975;
Grime & Jarvis 1975; King 1975; Panetta1979; Fenner1979, 1980; Silvertown 1980),it
has beendemonstrated thatsunlightwhichhas beenfiltered througha leafcanopymaybe
depletedin red lightto an extentthatcauses germination to be inhibited.In viewof this
evidenceof post-dispersal of leafcanopiesupon germinability,
effects it seemslikelythat
similareffectscould occur at an earlierstageduringthe maturation of the seeds on the
parentplant.Recentstudies(Cresswell& Grime1981) suggestthatinductionof a light
requirement in developingseeds occursnot onlyin inflorescences shadedby neighbouring
plantsbutalso in circumstances whereseed maturation is completedwithingreenparental
structures.It seems,therefore,thatthemostlikelyexplanationforthelightrequirement
notedin manyof thefreshly-collected seeds examinedin thisstudyis to be foundin the
properties
light-filtering ofthetissueswhichsurround thedevelopingseedsofthesespecies.
Failureto germinate in darknesswas mostcommonlyobservedin small-seededspecies
of disturbedhabitatsand marshland.Many of thesespeciesare knownto developlarge
and persistentseed reservesin the soil. This suggeststhatthe light-requirement of the
freshly-dispersedseed (perhapsin some cases reinforcedby theinfluence of leaf-canopies
priorto burial)preventsthegermination oftheburiedseed and initiatesgermination when
theseedis eventuallyunearthed by someformofdisturbance.

ACKNOWLEDGMENTS
We are gratefulto manypast and presentmembersof the Unit of ComparativePlant
we thankDr J.C. Hodgsonforhishelp
Ecologywho assistedin thisproject.In particular
in collectingseeds; A. H. Lumb, W. Hall and W. Ross-Fraserwho constructedthe
temperature-gradient bars; and MrsN. Ruttle,Miss S. H. Hillierand R. F. Grimefortheir
skilledassistancein thepreparationofthemanuscript. This researchwas conductedwith
thesupportoftheNaturalEnvironment ResearchCouncil.

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(Received1 May 1981)

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1046 Germination in a localflora
characteristics
APPENDIX 1

Effectsof chillingand scarification upon germination percentageand rate; the


controlpercentagerefersto the maximumobtainedforfreshor dry-stored seeds.
For the species subjectedto chilling,the values in parenthesesafterthe name of
the speciesindicatethe durationin monthsof the chillingtreatment (5 IC, moist
storage).Wherechillingwas precededby moistincubationat 15 IC, an additional
numberhas been insertedto indicatethe lengthof this pre-treatment. For the
thevalue in parentheses
speciessubjectedto scarification, refersto thedurationof
the periodof drystoragepriorto scarification. The significancevalues are based
upon comparisonofthegermination percentageofthetreatedseedswiththatofthe
controls.Germinationpercentagesin bracketsare estimates(to 5%) forspecies
in which substantialgermination occurredpriorto removal of seed fromthe
chillingtreatment. conventions:
Significance NS, P > 0.05; * P < 0 05, ** P < 0-01,
*** p < 0*001.

Control Chilled Scarified


t5O t5O t5O

% (days) % (days) % (days)


Annualforbs
Aethusacynapium(1) 22 10 52*** 6 - -
Anagallisarvensis(13) 7 - 58*** 2 - -
Atriplexhastata (3) 51 4 86*** 9 - -
Atriplexpatula(2) 0 - 90*** 2 - -
Capsella bursa-pastoris (10) 1 66*** 1 - -
Chaenorrhinum minus(1) 0 - 70*** 9 - -
Chenopodiumalbum(2) 24 5 73*** 5 - -
Chenopodiumrubrum(3) 100 3 98NS 10 - -

Erodiumcicutarium(3) 32 4 - - 80*** 2
Euphorbiapeplus (2) 4 - [201 - -
Euphrasia officinalisagg. (3) 2 - 17*** 2
Galeopsistetrahit(10) 0 - [251 -
Galiumaparine(1) 36 20 100*** 5
Geraniumdissectum(2) 25 5 - - 100*** 1
Geraniumlucidum(11) 92 4 - - 98NS 5
Geraniummolle(2) 20 3 - - 100*** 1
Impatiensglandulifera(1) 35 4 99NS 1 -

Impatiensparviflora(4) 0 - [100] - -
Linumcatharticum (2) 4 - 90*** 6 -
Medicago lupulina(10) 13 15 - - 98*** 3
Odontitesverna(3) 9 12 92*** 4 - -
Papaver argemone(15) 3 - ONS - -

Papaver dubium(2) 1 - [801 - -


Papaver rhoeas(3) 3 - [75] - - -
Polygonumaviculare(3) 2 - 44*** 6 - -
Polygonumconvolvulus(3) 0 - [1001 - - -
Polygonumhydropiper (3) 7 1 65*** 4 - -
Rhinanthusminor(3) 0 - 66*** 4 - -
Sisytnbrium officinale(2) 10 5 16NS 5 - -
Torilisjaponica(8) 0 - 74*** 11 -
Trifolium campestre(7) 72 5 - - 100*** 2
Urticaurens(3) 2 - 1NS - -

Vicia hirsuta(3) 20 4 - - 100*** 2


Viciasativa ssp. angustifolia(10) 88 24 - - 100*** 2
Viola arvensis(3) 20 4 [100] - -

Perennial
grasses
Bromusramosus(2) 22 30 52*** 2
Molinia caerulea 11 13 64*** 7
Nardusstricta 25 32 100*** 4

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J. P. GRIME et al. 1047
APPENDIX 1-continued
Control Chilled Scarified
t5O tSO t5O

% (days) % (days) % (days)

Perennialforbs
Aegopodiumpodagraria(12) 0 - [100] - - -
Agrimoniaeupatoria(3) 0 - 100*** 3 - -
Alchemillavestita(19) 4 - 22*** 7 - -
Alismaplantago-aquatica(1) 4 - 8NS 2 - -
Alliariapetiolata(10) 0 - [1] - - -
Alliariapetiolata(3, 2) 0 - 0 -
Alliumursinum(6) 0 - [5] - - -
Alliumursinum(3, 2) 0 - 0 -
Anemonenemorosa(3) 0 - 0 -
Anemonenemorosa(3, 2) 0 - 2NS -

Angelicasylvestris (3) 12 30 65*** 8


Anthriscus sylvestris(3) 0 - [100] -
Anthyllis vulneraria(7) 20 5 - - -100*** 1
Aquilegiavulgaris(6) 53 18 38* 13
Arummaculatum(6) 0 - 25*** 6
Baldellia ranunculoides(3) 13 1 6NS - - _
Calthapalustris(3) 31 25 83*** 3 -
Campanulalatifolia(1) 10 9 65*** 7 -
Carex demissa(2) 21 19 14NS 19 -
Carex laevigata(5) 0 - 47*** 15 -
Carex nigra(1) 46 12 90*** 9 -
Carexpanicea (18) 0 - 70*** 15 -
Chaerophyllum temulentum (3) 15 23 [100] - -
Chelidoniummajus (3) 25 29 66*** 6 -
Conopodiummajus (9) 0 - [90] - -
Convallariamajalis (3) 0 - 0 - -
Daucus carota (3) 49 8 84*** 5 -
Drosera rotundifolia (4) 0 - 30*** 18 -
Echiumvulgare(3) 12 4 8NS 5 -
Eleocharispalustris(19) 0 - 46*** 17 -
Endymionnonscriptus (2) 0 - 2NS 0 -

Endymionnonscriptus (3, 2) [100]


Galeobdolonluteum(6) 0 - [90] - -
Galiumpalustre(3) 42 23 17*** 11 -
Gentianellaamarella (8) 0 - 0 - - -
Geraniumpratense(6) 96 27 - - 75*** 5
Geraniumsanguineum(126) 54 16 - - 100*** 4
Heracleummantegazzianum(2) 0 - 40*** 12 -
Heracleumsphondylium (3) 0 - [50] - - -
Irispseudacorus(12) 48 15 - - 70** 9
Lathyrusmontanus(6) 46 6 - - 98*** 6
Lathyruspratensis(8) 41 9 - - 100*** 5
Lotus corniculatus(4) 49 9 - - 86*** 3
Lotus uliginosus(8) 23 3 - - 100*** 2
Lycopuseuropaeus(9) 0 - 60*** 4 -
Melilotusaltissima(0) 6 - - - 98*** 1
Menthaaquatica (1) 17 10 86*** 4 -
Menthaarvensis(18) 1 - 49*** 4
Mercurialisperennis (10) 0 - 0 - - -
Mercurialisperennis (3, 5) 0 - [15] - - -
Myrrhisodorata (1) 0 - [1001 - - -
Oxalis acetosella(6) 0 - [50] - - -
Pimpinellamajor(1 1) 21 24 [100] - - -
Pimpinellasaxifraga(2) 26 38 82*** 4 - -
Pinguiculavulgaris(2) 6 - 49*** 11 - -
Plantago major(3) 31 4 100*** 3 - -

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1048 Germination
characteristics
in a localflora
APPENDIX 1-continued
Control Chilled Scarified
t5O t5O t5O

Perennialforbs-continued % (days) % (days) % (days)


Primulaveris(6) 0 - [60]
Resedalutea(9) 0 - INS

Sanicula europaea (1) 0 - 0 -


Saponaria officinalis (2) 1 - 57*** 7
Silaum silaus (10) 8 - [100] -
Smyrnium olusatrum(1 1) 0 - [80] - - -
Sonchusarvensis(1) 0 - 72*** 1 - -
Sparganiumemersum(1) 7 - 30*** 11 - -
Sparganiumerectum(6) 15 20 30* 6 - -
Stachyssylvatica(8) 1 - 65*** 4 - -
Tamuscommunis(11) 0 - 0 - - -
Thalictrum minusssp. montanum(5) 12 15 40*** 8 - -
Trifolium medium(1) 18 5 -
Trifoliumpratense(3) 46 2 - - 100*** 1
Trifolium repens(16) 21 3 - - 99*** 3
Trolliuseuropaeus(8) 0 - [1001 - -
Verbenaofficinalis (3) 5 - 86*** 3
Veronicamontana(3) 0 - 0 - -
Vicia cracca (10) 60 22 - - 85*** 3
Viola hirta(14) 0 - 30*** 4 -
Viola lutea (5) 28 [100] - - -
Violapalustris(3) 23 18 92*** 8 - -
Viola riviniana(3) 0 - 90*** 2 - -

Shrubsand trees
Alnusglutinosa(3) 29 4 38NS 3 - -
Clematisvitalba(2) 1 - 50*** 10 - -
Crataegusmonogyna(2) 0 - 0 - - -
Empetrumnigrum(1) 0 - 2NS - -

Empetrumnigrum(3, 2) 0 - 85*** 23
Frangulaalnus (3) 0 - 24*** 25 - -
Fraxinusexcelsior(1) 2 - ONS - -

Helianthemum chamaecistus(6) 23 15 - - 86*** 2


Ligustrumvulgare(3) 20 71*** 19 -
Lonicerapericlymenum (4) 0 - [1001 - -
Lupinusarboreus(3) 10 - - - 92*** 2
Prunusspinosa(30) 0 - [101 - -
Rosa pimpinellifolia (13) 0 - 9** 4
Rubusfruticosus (3) 0 - 0 - - -
Rubus idaeus (6) 0 - 8* - -
Rubus saxatilis(3) 0 - 0 - - -
Sambucusnigra(4) 2 - [25] - - -
Sarothamnusscoparius(2) 90 11 - - 100** 3
Solanumdulcamara(6) 18 100*** 3 -
Sorbus aucuparia (6) 0 - [75] -
Symphoricarpos rivularis(12) 0 - [21
Taxus baccata (12) 0 - 0

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J. P. GRIME et al. 1049
APPENDIX 2
and germinationdata for 403 seed collections.Key to
Species-characteristics
notes:
parenthetic
(1) Based upon Perring& Walters(1962): u = ubiquitous,w = widespread,1 =
localized, s = southern,n = northern.
(2) Values referto the% occurrencein 2748 1-M2 quadratsdistributed withinthe
major terrestrialhabitatsoccurringin an area of 2400 km2(J. P. Grime&
J.G. Hodgson,unpublished results).
(3) s = skeletalhabitats(rockoutcrops,screes,cliffs, walls),d = disturbedfertile
habitats(arable, paths, soil heaps, manure and sewage waste, etc.), m =
marshland,g = grassland,w = woodland (includingplantations,scrub and
hedgerows).
(4) Based uponClapham,Tutin& Warburg(1962).
(5) 1 = nearlyspherical,2 = ovoid, rhomboidalor turbinate,3 = trigonousor
triquetrous,4 = lenticular,reniform or subulate,5 = cylindricalor ligulate,
6 = clavate,7 = winged,8 = tadpole-shaped, 9 = conical (includingall seeds
witha pappus).
(6) 1 = absent, 2 = straightawn(s) or spine(s) only, 3 = hygroscopicawn or
spine,4 = pappus or persistent calyx, 5 = large hook(s) or barbedspine(s),
6 = eliasome, 7 = wing.
(7) 1 = absent or inconspicuous, 2 = radial or irregular,3 = antrorse.
(8) 1 = smooth, 2 = rugose, tuberculate, muricate or reticulate, 3 = striate,4 =
hairy,5 = striateand hairy,6 = mucilaginous.
(9) Seed coloursare describedusingtheMunsellSystem(Munsell1954).
(10) Germination is expressedas the final% germination and as t5O (thetimein
days requiredfor50% oftheseedsto germinate).
(11) * = % germinationsignificantly (P < 0.01) differentfromthat of freshly-
collectedseeds.
(12) Pre-treatments (p): d = dry storage at 20 IC, c = chillingat 5 IC, s
w = moist storageat 20 OC in darkness,f= freshly-collected
scarification,
seed.
(13) Range is describedin termsof thelowestand highesttemperatures at which
germination attained50% ofmaximum.
(14) A keyto curvetypesI-IV is includedinthelegendto Fig. 1 (p. 1022).
(15) * = % germination significantly fromthatof seeds in the
(P < 0.01) different
light,t = additionalmeasurements of germinationin darknessare presented
forthisspeciesinTable 4 (pp. 1037-1038).

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1050 Germination in a localfiora
characteristics
APPENDix 2-continued

-
E~~~~~~~~~~~~~~~~~~~
~~~~~~~~~~~~~~SeedcollectioE E

National Weight

(a) Annalgrase
1.-A tr crvphivea r 0.2 2.18.74 Oldcolier~y
spoilheap SK 555685 20 p 1 55 0.40 8 3 3 0.09
2A tr praecocGr 0.8 3.7.73 Oldstonequarry heap SK 248803 10 p 2-20 0.80 8 3 3 0.18
3 Avena faw,aGr <0-.1 d 4.8.76 Fallowarble SK 522861 60 p 24.10 2.10 8 3 3 14.13
4Brmspsuohonii Gra <0.1 g 14.873 roadveg
Disturbed SK 291838 40 p 11.65 1.90 8 2 3 3-15
5. Briu trh r s 0-8 d 10.8.73 Heapofminerasoil SK 293984 65 p 37.-55 1.100 8 2 3 8.37
6Ca pdti iidi r s 0.4 20.7.73 Deeitlmsoeqar flo 5K 505826 10 p 1.80 0.50 4 1 3 0-19
7. Hodii niiii r 03 d 20.7.73 brick
Urbarn. rubble SK 349863 45 p 3775 2 70 9 2 3 6.55
8 Poaaiu r 12.2 d I 7 73 Cultivated
garden soil SK 333874 15 p 2.60 0.85 2 1 1 0.26
9V lt boioie r s 0.3 21.673 Rockoutcrop inlimestonequary SK 555678 15 p 13-70 1.00 8 2 3 0.30

10 A tis ciati Umb s 0.4 d 5 1073 Ditre odiebn SK 342752 40 p 2-85 2 05 4 1 1 1.06
11A iigli rvni r 09 d 24.8.73 Flor fdisusd sndquary SK 564819 15 p 1.45 1.05 3 1 1 0.40
12. A phaies nensis Rs 13 15.673 Sandyradsidebank SK670918 10 p 2 30 1.10 4 1 1 0 18
13A raiopi thalina ru 0-5 226.72 Derelictlimestone quaryheap SK 165661 25 p 0 50 0.50 4 1 1 0.02
14 A eai eriIioi Car 1.9 108.73 Oldcide heap SK 293985 10 p 0.50 0.50 4 1 1 0.06
IS Atriplehastaa Che 13 d 8.10.73 Hollowsubjecttoflood1ing SF 595000 60 p 2.00 1 85 4 1 1 0.86
16. Atriple.cpawula Che 30 d 7.873 Trampled wstegroud SK 375750 60 p 1.80 1.55 4 1 I 1.33
17 Bidenstrprl Com s 0.2 m 4 10.73 Lakesidemarhhand SK 455644 35 p 9 05 2.40 5 5 3 2.72
18Blcslii perdoial Gn 0.3 24.873 Flooro diue lietn quarr SK 555827 30 p 0.35 0.30 4 1 1 0.01
19 Capsellabtrapsoi Car 2.6 d 1.8.73 Urbanwateground SK 355835 25 p I .00 0-50 4 I 1 0.11
20 Cadiin isl r 0.8 s 22672 Deeitlmson uryha SK 165661 20 e 1.00 1.00 4 1 I 0.09
21. Ceiarui rire e 07 8.10.73 Ditre atgon SF 598028 20 p 0.35 0.30 1 1 I 0.01
22 Ceattmsindcnri Car 0.3 12673 Dereictli,metoequaryheap SK 555678 10 p 0.50 0 50 1 1 I 0.03
23. Chaenor, un iu Scr 0.3 20.9.73 D,ereictrailway cinders SK 261655 15 p 1.10 0.65 2 1 1 0.07
24 Cheiiopodiui alu Che IS11 d 7.873 Urbanwatgon SK 375750 50 p 1.50 1.35 4 1 1 0.77
25. Cheiiopodiui rubru,ni Che 07 d 8.1073 Hollowsubject toflooding SE 596008 40 p 0.60 0.55 4 1 1 0.09
26 Coi'zacana~denss Cms <0. I d 8.10.73 Refusetip SF 593014 50 p 3.55 0.50 5 4 3 0.05
27. Crepis cailat Com u 2.0 s 10.8.73 Cinder andslagheap SK 293985 55 p 5.75 0-50 9 4 1 0.21
28. Draba Inrls r 0.2 s 22.6.72 Derelictlimestonquarr heap SK 165661 20 p I 00 0.55 4 1 1 0.05
29. Erige,ron acer Com s 0.6 8.9.73 Derelictra~iway cidr SK 136733 20 p 6 40 0 45 9 4 3 0.1 I
30 Erdii iulrui e <0.1I s 8 10.73 Ditre snybn SF 598027 30 e 17.10 1.30 6 3 3 1.25
31. Erophilavra r 0.3 6.6.73 Shallowsoi onlimetoneoutcrop SK 184716 10 p 0.63 0.50 4 1 1 0.03
32. Etiphorbkapeplus Eup 0.2 d 7.8.73 Trmldwatgon SK 375750 20 p 1.70 1.00 2 6 I 0.48
33 Etprai o/Jkicinai Scr 1.4 5.10.74 Floorofdereict limestonquarr SK 150612 15 p 0.95 0.60 2 1 1 0.13
34. Galeopsis letrahit Lab 1
i.3 d 16.8.73 Disturbed wasteland SK 436863 55 p 2.95 2.30 4 1 1 4.83
35 Gaisg prflr Com s <0.1I d 2.10.75 Ditre rasd SF 558034 40 p 2.50 0-55 9 4 3 0-18
36. Gaii aparine Rub 5.8 w 31.10.72 Hedgero SK 232748 65 p 2.33 2.00 1 1 2 7.25
37 Geaii iseti e <0.1I d 18.7.74 Soilepa deoito site SK 545913 35 e 2.00 1.60 2 1 2.47
38. Geaiunlcdi e 0.3 11.7.74 Shallowsoilonsae ietn SK 135825 25 e 2.15 1.35 2 1 1 1.09
39. Geaiii iol e 0.7 28.8.73 Distubedwateland loutcropSF 615003 25 e 1.85 1.45 2 1 I 1.24
40 Geraiiitiroetaii Ger 4.7 3.10.73 Lietn ala als SK 145733 30 e 1.25 110 2 1 1 1.14
41. inpawiens glandulifera Bal 0.9 m 23.10.74 Stremside SK 527837 150 e 4.75 3.45 4 1 1 7.32
42. liptei atilr Bal I <0.1 S.10.73 Distubedwodlad SK 265565 65 e 4.45 2.15 4 1 I 6.91
43 inci biftiu Jun 1.0 m 5.10.73 Shallowpoolindisturbed ground SK 362774 10 p 0.60 0.35 2 1 1 0.02
44 Lasn oiiui Com u 1.1 d 28.8.73 Ditre rasd SF 615002 55 p 4.50 1.35 5 I 1 1.27
45. Liuncthriui Lin 3.7 9.8.74 Raiwy embarnkment SF 603003 15 p 1.40 0.85 4 I I 0.15
46 Marcraia(iaiie Com u 47 d 7.873 Ditre atgon SK 523649 15 p 1 20 0 50 5 1 1 0 08
47. MeiaoIpln Leg 2.5 148.73 Lietn otcrops SK 156732 25 p 2.85 2.40 4 1 2 2.01
48 Mohigatii i a 0.8 w 19.673 Scrubindereictquary SK 554677 25 p 0.95 0.75 4 1 1 0.22
49 Mvo~sosavniBr 0.8 d 28.8.73 Wasteland SF 600009 20 p 1.75 0.95 4 1 I 0.29
50. Mrosolis cespilsaBo <0. I m 26.7.73 Pondmargin SK 486790 30 p 1.55 1.30 4 1 1 0.24
51 Mrslsrioisii o 0.6 4.6.73 Limestonotcrops SK 183716 10 p 1 00 0.75 4 1 1 0.11
52. Odo,,itites en Scr <01I g 5.10.73 Lightly trampled pathmagi SK 381660 25 p 1 65 0.95 2 1 1 0.IS
55. Papaveraiemone Pap s <0.1I d 29.7.74 Ditre rasd SK 633927 30 p 1.00 0.60 4 1 1 0.17
54 Papavrdbunla <0.1I d 8 10.73 Ditre snybn SK 658964 40 p 0.95 0.75 4 1 1 0.12
55 Papaverhoeas Pap s 0.7 d 5.1073 Ditre odiebn SK346745 40 p 0 90 0.65 4 1 I 0.09
56 Paae oinfri Pap <0.1I d 7.873 Riverbank SK 375749 65 p 1.00 0.90 4 1 1 0.25
57 Porou vclr Pol 61 d 78.73 Trmldwatgon SK 373750 10 p 3.75 2.10 3 1 1 145
58 Pol,hgonincovovuu Pol 2.0 d 7.8.73 Ditre wstelad SK 522649 75 p 4.00 2.35 3 1 1 1 28
59 Poli'gonum hvdropiper Pol 0.5 m 168.73 Lakemargin SK435863 50 p 3.50 2 00 3 1 I 1.24
60 Polivgonui lapathifobhtti Pol 05 d 7.873 Fallowarable SK 508649 60 p 3.00 2.55 4 I I 2.25
61 Po'gni priai Pol 31i d 317 73 Ditre odiebn SK495851 50 p 3.20 2 30 4 1 1 2.12
62. Rauciisselrtt a 0.6 m 7 873 Fecr SK583578 40 p 2 20 1.23 4 I 1 0.16
63 Rh,nathiis innrScr 0.4 g 12.7.73 Floorofderelct limestone qurr SK 142732 20 p 4-85 3.75 7 7 1 2.84
64 Roip sadc r 0.5 m 31.873 Pondmargin SK 453645 35 p 0.90 0.60 4 I 1 0 07
65 Saginaapewala Car 03 12673 Derlictlimestoequaryheap SK 555678 10 p 0 36 0.26 4 1 I 0.001
66 SafrgaU ridacQ,vifes Sax s 04 22772 Deeitlmson uryha SK 164662 10 p 0.47 0 25 2 1 1 0.01
67 Sceaihsaiiu a <0.1I 15673 Sarndytrack SK669921 S p 3 90 1.35 9 I 1 1.16
68 Se,wecisqualidii Com s 3.3 d 107 73 Disturbed wateland SK 332873 25 p 7.50 0.75 9 4 3 0.21
69 Seicw s'vIvaicus Com w 0.2 28.8.73 Disurbedgrasslad SF 595011I 50 p 5 80 OSO0 9 4 3 0.23
70. Seei vsou Com w .0 28873 Rala ine rc SF603003 35 p 9.00 OSO0 9 4 3 0 60
71 Seei vlai Com u 3.9 d 76.73 Cutvtdgadnsi SK33l1870 20 p 4.10 0 50 9 4 3 0.25
72 Siai reni r 07 d 1.8.75 Margin ofarablefield SK 542802 SS p 1.85 I 25 I I I I. IS
73 Sivnru lisiii r I <0Ol d 810 73 Dereictrfuetip SF 593014 60 p 130 0-95 4 I I 0 25
74. Sivnru officinale Cu 0.3 d 31.7.73 Roadside SK 461845 60 p 1.20 0.95 4 I I 0.23
75,Sola u ngugin Sol <0. I d 8 10.73 Disturbed sandvbarnk SK 658964 30 p 6.40 5.10 4 I I 0 76
76. Sonchusaper Com u 07 d 3l17.73 Roa~dside SK461845 8S p S 30 I IS 9 4 I 0.32
77 Socu oleraceus Com u 31i d 178.73 Roadside SK 311850 8S p 6S50 0-8S 9 4 1 0.27
78 Spruaavni a IS d 317 73 Disturbd rodside SK 495851 20 p I 45 1.30 4 I I 0.42
79. Siellari,iwdiaCa 6.3 d 5.10.73 Margin ofarblefield SK 353738 IS p 1.30 100 4 I I 0 35
80 Thap res r <0 I d 15.875 Arablefield SE53l10l5 35 p 2 IS ISO0 4 I I 0.95
81 Toii aoic Umb u 0.7 d 19.9.73 Roadside SK 239761 65 p 4 00 2 40 4 5 2 I 98
82 Trflun res Leg <0 I 24.8.74 Floo ofad lvb SK 671918 IS p I 20 I 00 4 I I 0 31
83 TrifoliiicamipesIre Leg <0OIl 14.873 Lietn uco SK 157732 IS p 1.40 0.90 4 1 I 0.36
84 Trperseiuniaiinii Com u 2S5 d 16.873 Lakemargin SK 435863 35 p 2 00 IlO0 S I I 0.29
85. Uria rn Urt 04 d 15.673 Floo ofsadyI ay-bv SK 671918 35 p 2 12 1.50 4 I I 0-50
86. Vaeinla aiil Val <0OIl 226.72 Deeitlmson uryha SK 164662 20 p 2.10 IS1 2 I I 0.66
87. Veoic avni Scr 0.8 226.72 Deeitlmsonuryha SKl164662 IS p 1.12 0.8l 4 I I OIl1
88 Jetonc pe-swa Scr 2.0 d 78.73 Trampled watelad SK 373750 2S p 1.50 0-75 4 I I 0S52
89. Vkciahirsula Leg <O.l d 2 874 Deeitcllevha SK 555683 2S 2.05 2.00 I I I 2l19
90. Vicia5aiva ssp.anqs~oi Leg s Os- d 17.773 Wasteland SK 157732 55 2.70 2.30 I I I 1306
91 Vkiolaarensis Vio 0.9 d 15.673 Marginf sndvarblefield SK 6679l18 20 C ISS 1.00 2 I I 0.40

(c) Perenialgase
92. Agoirncniiii r 0.5 w 3.10.73 Shadedrodsidebank SK 144734 65 p 10.45 2 00 5 I 3 4.04
93 Agropiron repe'ns Gra 9.2 d 5 1073 bewe aralefieds~
Wasteland SK 356742 6S p 1290 1.80 5 2 3 2.02
94 Agotsciiiaspcnn Gra u) 3 g 288973 Dereictwtgrasslnd SE59501I 35 p 1.70 0 3S 2 1 I 0.OS
95. Agro,iicarninsp iniai Gra n 36 7 1174 Acidicheeppastur SK155741 3S p 250 0.50 2 I 3 0 06
96 Agotsggne Gra - d 0 10.74 SK504923 60 p 2 3S 0 60 2 I I 0 09
97 A rsi tlnfr r 18.0 d 288.73 Hollowubjet toflooding SF 605002 70 p 1 6S 0 46 2 I I 0.02

NI-irgmd
on'aabl fied

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All use subject to JSTOR Terms and Conditions
J. P. GRIME et al. 1051

Colour (9) 2!griainatrdysoaeI1


2
-~~~~~~ 5 -C 20 'C~ totetmperature ~ Respornsetotlight15)
E2
2
%
~
O~~~~~
l50
3 months
l%l l'oo
6 months 12 montths
I
>I month
p(12) ~
Rantge(13)
C i Curve 14) p 12) %
Light
lo %
Shade
10
(,
Dark
1

- - 96* 5 d <5 27 2 11 d 72 9 96* 5 (00* 5


2 5YR/4/6 17 (1 82* 6 94* 4
17 . 79* 7 96* 13 99* 4 98* d 10 31 4 11 d 100 4 100 3 91* 40
2 5YR/5/4
2 5YR/4/2 42 3 70* 3 - - - - 70* 2
9 99 (1 97 6 99 2 too d <5 34 2 11 d 100 2 100 1 100 29
4 2 5Y/7/2 98 it
tOO 8 100 5 100 2 d <5 37 1 11 d 100 I 100 I 100
5 5YR/6/2 100 9 8 100
3 100 3 d 6 33 2 11 d 100 3 98 3 94 39
4 lOYR/8/2 97 (4 100 5 98 3 92
d <5 31 1 11 d 100 2 100 I 100 29
3 5Y/8/3 96 8 98 (1 94 (5 90 3 99 2
99 4 (00 3 d 7 31 2 11 d 100 3 100 3 8l* 3
I 5YR/5/3 96 (1 99 4 (00 4 4
70* 5 10 3 d 6 30 3 11 d 97 (00 2 (00 30
4 2-5Y/8/6 92 (9 75* 29 90 8

0 13* (8 3 14 22* (0 5 9 c (0 25 5 11
1 75YR/8/4
0 0 I . 6 8 7 4 c <5 35 1 11
2 IOR/2/2
80* 5 d <5 25 4 It d 87 7 94 7 25* 69
I lOYR/8/4 0 6 23 4 9 48* S
lo* (2 4( 8 84* 5 93* d 7 31 2 II-IV d 84 3 87 4 19* .0
I 2.5YR!6!6 0
99 2 d 7 28 3 11 d 99 2 (00 2 90 2
2 IOR/2/1 92 (1 99 (1 (00 (0 99 5
4 31 5 d (9 33 3 1 d 40 5 32 6 4*
1 Black I 31* 7 39* 6 51*
0 I 0 C 10 23 3 1-111 C 84 2 90 1 48* 2?
I Black 0 0
20* 21 21* 24 d 24 31 3 1 d 68 30 24* 30 2*
5 5YR/4/2 4 25 19* 27 15 25
99 8 97 6 97 7 90 7 d 11 28 6 IV d 91 8 92 7 15* 12?
2 7-5YR/4/2 93 10
0 I 0 0 -
I IOR/5/6 0
92 6 92 5 d (1 28 6 I-TI d 51 10 74* 6 72* 7
I IOR/4/6 90 30 93 36 86 17 1
9 93 8 97 8 - 4 (3 29 6 IV d 97 10 92 8
2 IOR/3/6 99 11 95
24 68 19 84 10 (00 4 d <5 31 2 II d 82 4 88 4 80 4?
2 2-5YR/6/6 82 35 76
0 0 0 0 c (2 35 4 II c 83 5 97* 5 49* 5?
3 5YR/4/3
4 24* 5 lo* 5 d 8 36 2 II d 66 6 48 6 5*9
I Black 0 2 12*
4 99 4 (00 3 - 4 25 35 <1 I d 100 5 (00 4 P*
I 7-5R/2/4 98 -5 (00
2 97 1 (00 2 d 13 30 2 I-TI d 100 2 100 3 2*
I 7-5YR/6/4 98 3 99 2 (00 8*
3 88 11 96 6 85 6 99 3 (00 4 d 10 36 4 1-11 d 99 5 100 3
5YR/5/3
95* 4 96* 3 97* 3 97* 3 d 10 29 2 1-11 d 88 4 91 4 84 5?
2 2-5YR/6/6 0
4 (00 4 99 3 d 13 31 3 TI-TV d (00 6 100 4 97 5
4 2-5Y/7/4 100 4 (00 4 (00
4 7-5R/4/2 10 6 23 2 32* 4 13 3 (6 2 - - -
52* 4 91* 4 d 6 25 4 1-11 d 95 4 97 4 94 4
I IOR/5/6 0 21* 23 28* 3-5
2 0 4 0 4 - -
2 2-5Y/7/0
3 5YR/6/2 0 0 0 2 -
2 5YR/3/2 0 0 0 0 0- -
I-TI - -
4 2-5Y/2/0 52 3 49 3 72* 2 86* 2 - d 8 37 2
2 I 36* 20 0 d 6 26 5 IV d 67 14 95* 8 61 12?
4 5YR/3/1 0
2 2-5YR/3/2 0 lo* 27 25* 5 <5 29 2 IV
. 4 17 45* 14 92* 4 d <5 34 2 IV (00 7 100 6 100 2
I 2-5YR/6/8 2
I 0 0 I 20* 3 85* 3 -
IOR/5/3
25* 19 88* 15 88* 10 - d 57 13 61 8 64 13
I 7-5R/4/4 59 15 59 13 -
I 5YR/3/3 0 0 l5* 4 35* 4 - - - --
3 5YR/3/3 0 0 0 - -
. 26* 13 25* 20 78* 13 99* 7 - - - - d 98 4 97 4 0*
6 2-5YR/6/6 3
30 92* 4 8(* 4 91* 18 d 13 35 4 1 d 98 4 (00 4 56* 4?
3 IOYR/7/6 7 18 96* - -
I I 4 9 3 ii 0 0 c 7 36 3 II
2-5YR/6/6
10 3 93* 3 d (1 23 4 1 d 47 6 58 6 3*
3 5YR/5/1 5 7 54* 11 72* 50*
2 2 It
I 2-5YR/5/4 8 7 (3 14 5 7 (3 15 3 <5 36 2 II ( 00 (00 (00
11 66* 8 89* 6 94* 6 d 9 21 5 II d 98 6 98 6 80* 6?
I IOR/2/1 I 14*
899 4 - - d <5 27 3 II d 94 4 99 4 4*
I IOR/2/1 37 11 57* 7 88* 6
6 (00 6 (00 4 - - - d 100 6 (00 5 3*
I IOR/2/2 95 7 (00 6 (00
98 11 100 5 - - - d 98 7 100 7 98 9
I 2-5YR/4/8 (00 31 (00 22 (00 28
9* - - c 7 25 3 II - -
2 IOYR/7/3 0 0 3 19 12
2 IOR/2/2 3 . 0 0 0 - -
2 IOR/2/2 I 0 0 0- - - -
2 IOR/2/2 0 0 0 3 4
0 2 93 3 d <5 24 3 II d 97 5 99 4 47* 4?
2 IOR/2/2 I 0
2 0 0 0 - -
2 5R/2/3 I
I Black 0 0 0 0 0-- -
I 0 0 2 0 7 1 - -
7-5R/3/2
29* 12 20* 17 94* 5 62* 9 d 28 39 3 1 d 77 7 85 6 P*
I 7-5R/3/2 0
5 19 41 30 40 2 1 d 78 2 72 1 52 1
I 7-5R/2/2 0 II* 10 22 10 759
? ? ? ? ? ?- ? - 36* 8 - - - d 31 10 35 6 0*
2 7-5YR/8/4 85 8
0 0 0 0 - -
I IOYR/4/2 0
29 17 76* 13 20 12 32 4 60* 25 d -35 39 4 I-ITT d SI 4 72* 12 0*
2 5YR/5/6
2 IOR/4/4 81 44 96* 12 96* 12 92 11 98* 12 - -
97* 22 96* 12 97* 7 d 7 25 7 IV d (00 7 98 6 359 9?
2 5R/2/1 0 83* 27
47* 7 54* 6 d <5 25 5 II d 57 7 64 6 28* 5
3 IOYR/7/6 25 10 29 5 41 6
4 91* 3 d 7 36 2 TV d 70 5 97* 3 459 3
S IOYR/4/4 68 5 78 5 86* 4 73
4 99* 3 d (3 33 3 TV d 94 4 100 4 3*
S IOYR/3/1 52 8 91* 3 (00* 3 (00*
S 0 55* 5 67* 4 13* 6 d 11 31 4 TI-TV d 95 5 98 4 0*
7-5YR/4/2 I
99* 3 d II 28 3 II d 79 11 (00* 3 9* 5
S 7-5R/3/4 82 11 76 9 97* 6 99* 10
I 2 2 56 3 66* 2 d ge~rmnationt-0 d 72 3 98* 3 36* 2
7-5R/3/2 39 4 35 19*
1 IOYR/6/8 (4 19 4 6 14 3 16 0O* - - - -
I 5YR/4/4 0 I I 3 7 lo* 5 - -
8 0 - - - - - - d (00 7 100 5 (9* 4
I 2-5Y/7/4 0 80*
91 5 2 99* 2 96* 3 d 9 35 2 IV d (00 3 100 2 4*
3 2-5YR/4/4 79 5 92
74 3 69 3 77 3 83 3 d 6 34 2 II d 57 3 68 2 28* 4
3 2-5YR/4/6 73 5
22 3 21 3 29 3 68* 2 d <5 35 1 II d 21 4 (7 4 22 3
1 Black 37 2
14 4P 6 52* 2 68* 2 42* 9 d 10 28 2 I-TI d 89 2 99 1 83 2
2 7-5R/4/2 5
8* d 22 36 2 I-ITT - - -
3 7-5R/2/3 26 4 3 9* 4 26 4 35 3
S IOYR/7/4 0 0 0 0 - - c <5 25 5 TI-TV
6 20 0 12* 7 5 21 <5 36 2 1-11 92 6 98 4 (00 2?
I I
5 68 8 69 14 68 10 13* 5 <5 29 2 ITT-TV 96 2 93 2 90 2
I 72
21 69 11 61 12 77 12 82P 12 d (3 35 6 ITT-TV d 89 8 89 9 0*
3 61
2 I 2 0 d ge~rmnation-0 d 3 6 89* 6 29* 6
I 5YR/6/2 I
9 92* 8 93* 5 d <5 27 3 II d 91 6 92 6 85 5?
3 7 5YR/5/4 0 80* 13 92*
9 96* -5 97 5 d 7 23 5 1-11 d 92 8 98 6 48* 7
I 7.5YR/5/6 0 56* 27 72*
11 70* 9* 3 92
3 d 7 35 3 1-11 d 100 4 (00 3 67* 4?
2 IOYR/6/4 I 21* 39

~ 4 10 - - <5 32 1 II ( 00 2 100 2 94 2
I v and m 0 20* (0
24 86 33 59* 13 88 24 <5 27 2 II (00 8 99 9 (00 8?
1 and m 95 52 85
0 20* 4 - - - - -
I 7-5YR/5/6 I 0

7 11 (00 6 - - - - - - d 96 9 96 6 80* 12
2 IOYR/7/3 96 13 92 (00
- - 7 70* 7 48* 6 d germination- 0 d 72 4 92 2 88 2
2 5YR/6/2 I (2P
94* 5 91* 5 899 4 d 9 30 2 II d 89 3 94 2 91 3?
1 7-5YR/5/6 65 6 78 5
4 70 6 60 6 86 4 - - - - d 78 9 80 6 30* 5
2 5YR/6/6 72 3 90*
3 78* 3 97 4 99 3 d 7 34 2 II d 93 3 96 2 73* 2
1 -SYR/5/6 98 3 92
8P* 99* 5 95 .5 96* 5 d II 36 3 1-11 d 98 4 (00 3 72* 4
1 5YR/5/6 61 7 5

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All use subject to JSTOR Terms and Conditions
1052 Germination in a localfiora
characteristics
APPENDIX 2-continued
Dehiscene Dispersle

cr
Date Habitat
~~~~~~~~~~~~~~~~~~Natiornal Wih
(mg)
Species U. Gridref.

98. Agrostistenuis Gra u 8.3 g 31.10.72 Acidicsheeppasture SK 223745 35 p 1.85 0.50 2 I 1 0.06
99. Alopecurus pratensis Gra 1.0 g 1.8.73 Semi-derelict
pasture SK 178745 60 p 5.50 1.95 4 3 3 0.71
100.Anhxnthui odoratum Gra 6.7 g 11.8.72 Wasteland SK 356656 35 p 4.30 1.25 8 3 3 0.45
101.Arhenathe~rum elatius Gra 14.1 g 29.8.72 Derelict1limestone
grassland SK 175655 90 p 10-75 1.80 5 3 3 2.39
I02. Brc ,oimpnau Gra 2.5 g 25 10.73 calcareous
Derelict pasture SK 500666 45 p 12.25 1.85 5 2 3 2.85
103. Brcioims,iaiu Gra 3.3 w 16.10.72 Wasteland SK 153732 60 p 19.70 1.60 8 2 3 0.62
104. Brizainedia Gra 1.7 g 24.8.72 Derelict
calcareous pasture SK098714 35 p 2.95 1.75 4 1 1 0.23
lOS. Bromserectus Gra 1.0 g 24.8.73 Derelict
limestonequarry SK 555825 80 p 17.45 1.80 8 2 3 4.23
106. Brmsriou Gra 2.0 w 16.1072 Shadedroadside bank SK 153732 125 p 14-45 2.15 5 2 3 7.37
107. Caars autc Gra <0.1 m 26.6.75 Marsh SK 274590 35 p 2.75 1.15 2 1 3 0.46
108.Cinsrs rsau Gra 2.8 g 26.7.73 Dam,pcattlepasture SK 361775 45 p 3.45 0.75 5 2 3 0.70
109. Dat,i gonrt Gra 17.9 g 20.773 Wasteland SK 324810 65 p 6.70 1.35 3 2 3 0.51
110 Descha,inpsia cespitosa Gra 6.3 m~ 27.10.72 Margin oflake SK 435863 125 p 2.85 0.85 2 3 3 0.31
III Decansafeus Gra n 20.3 g 22.8.72 Acidicgrasslnd SK 305832 30 p 3.50 1.25 2 3 3 0.43
112 Festucaaru,dinacea Gra 0.3 g 16.10.72 Wasteland SK 155732 130 p 7.80 1.60 5 1 3 1.26
113. Festuca giganuea Gra 3.4 w 16.1072 Shadedrodsidebank SK155732 100 p 19-50 1.50 8 2 3 3.-12
114 Festucaovin Gra 11.2 g 31.7.72 Deeitlmstn rsln SK 175655 30 p 4.25 1.05 5 2 3 0.38
115 Festuwapratetisis Gra 0.7 g 7.8.75 Wasteland inderelictquarry SK 563795 60 p 7.45 1.45 5 1 3 1.53
116. Fetc ur Gra 19.4 g 11.8.72 Wasteland SK 356655 40 p 7.30 1.05 5 2 3 0.79
117. Gli'ceriafluitans Gra 1.6 nm 31.8.73 Flodedcattlepasture SK 243578 55 p 6.90 1.55 5 1 3 1.20
118 Heittiho rtts Gra 2.0 g 14.8.73 Deeitclaeu rsln SK 155732 45 p 18.45 1.55 8 3 3 2.08
119. Helictotrichornpubhescens Gra 1.5 g 9.7.75 Roadsidebank SK 493637 50 p 14.90 1.05 8 3 3 1.92
120 Hoilcus lanatus Gra 18.1 g 11.8.72 Wasteland SK 356655 40 p 4.80 1.95 4 3 3 0.32
121 Ko~elericisat Gra 2.7 g 23.7.73 Derelictcalcareousgrassland SK 223743 25 p 4.90 1.00 5 1 3 0.30
122. Loliiicn
pere,ine Gra 7.8 g 16.8.73 Pathmargin neararablefield SK 439863 35 p 6.60 1.80 5 1 3 1.79
123. Vfelicatiutns Gra n 0.3 23.7.73 Hazelscu SK 224742 30 p 2.65 1.35 4 1 1 1.96
124. Afiliu,,n
effusu,'n Gra 1.4 3.7.73 Mix~eddeciduouwodland SK 323813 85 p 2.70 1.20 4 1 1 1.20
125. Afoliniacaerulea Gra n 0.7 g 5.10.73 Heathland SK 295675 80 p 4.10 1.10 2 1 1 0.53
126 Nardusstricta Gra n 3.2 g 14.8.72 Acidicwasteland SK 546915 20 p 10.40 0.85 5 2 3 0.38
127 Phalartirudinacea Gra 1.2 nm 16.10.72 Riverterrace SK 150732 90 p 4.15 1.20 2 1 3 0.67
128. Phei rtts agg. Gra 2.1 g 12.10.74 Derelictarablefield SK 366683 75 p 2.25 2.00 2 1 3 0.45
129. Po onpes Gra 0.2 20.9.73 Topofwall SK 284630 30 p 2.55 0.50 5 1 2 0.20
130. Po rtei ssp angustf/olia Gra 1.1 g 31.7.73 Derelictgrassland SK 533912 45 p 3.00 0.75 3 1 2 0.19
131. Po rtni s.paeii Gra 15.0 g 23.773 Roadside bank SK 223743 45 p 3.50 0.95 3 1 2 0-25
132 Poatrivialis Gra 22.0 d 14.8.72 Hedgebank SF 512002 40 p 3.25 0.90 3 1 2 0.09
133. Piicci,iellia
distans Gra I <0.1I d 8.10.73 Tram~pledlimetnet quarryheap SF600003 35 p 2.00 0.75 2 1 1 0.28
134. Sesleriaalbicans Gra I <0.1 g 24.7.75 Sheeppasture NZ 838283 25 p 4.45 1.20 2 1 3 0.32
135 Sieglingia decu,inbens Gra n 0.9 g 11.8.72 Wasteland SK 356655 25 p 3.20 2.05 4 6 3 0.87
136. Trisetinmifkaesce,ns Gra 5-3 g 7.7.75 Disusedrailway cutting SK 485755 35 p 5.70 0.65 8 3 3 0.18

(d) Perennial
forbs
137. Acilaiileoit Co 3.5 g 27.10.72 Wasteland SK 438863 25 p 2.05 1.05 5 1 1 0.16
138. Achilleaptar,nica Com, n 0.4 nm 15.9.75 Ditchinhaymeadow NZ 862291 40 p 1.80 0.75 5 1 1 0.22
139. Adoa nschatellina Ado 0.4 w 3.7.73 Scrubonriverbank SK 151731 10 p 10.00 8.00 1 1 1 0.63
140.Aegpodiumpodagraria Unmb u <0.1 g 31.8.73 Roadverge SK 246577 70 p 3.90 1.40 4 1 1 2.73
141. Aginoi euaoi Ros <01 g 16.10.72 Wasteland SK 150733 45 p 7.90 5.70 2 5 2 23.78
142 Ajugareptans Lab 0.2 w 23.7.73 Hazelscrub SK 224743 20 p 2.25 1.50 2 1 1 1.47
143. Alche,inillavstt Ros <0.1I 23.7.73 Hazelscrub SK 224743 20 p 3.00 1.10 2 1 1 0.47
144. A iiapatg-qitc Ali 1.0 nm 27.10.72 Marshland atnmargin oflake SK 437863 60 p 2.35 1.60 4 1 1 0-27
145. A lai eilt Cru 0.8 d 4.7.73 Shadedroadside bank SK 147734 70 e 3.15 1.00 5 1 1 2.25
146. Aliuhnursnui Lil 1.9 w 20.7.72 Mix~eddeciduouwodland SK 531831 25 p 2.45 2.35 4 1 1 4.27
147. Anemone ne,inors Ran 3.7 w 10.6.74 Oak-birch wooxdland SK 325813 15 p 4.90 1.90 4 5 2 0.99
148. Angelica sYvestris Um,b u 3.2 m~ 27.10.72 Dampwasteland SK 437863 115 p 4.55 3.40 7 7 1 1.15
149. Anthriscus s Nvestris Um,b u 3.9 g 23.7.73 Roadside SK 360774 80 p 3.40 1.35 6 1 1 5.18
ISO. A,ith'llisvunrra Leg 0.4 g 24.8.72 Derelictlimestone grassland SK 098715 15 p 3.90 2.40 4 1 1 2.76
151 Aquilegiavulgaris Ran <0.1I 29.6.73 Mixeddeciduous woodland SK 524784 70 p 2.55 1.30 4 1 1 0.78
152. Arabiscacasica Cru I 0.2 4.7.75 Roadsidebank SK 295575 20 p 1.60 1.25 7 7 1 0.26
153. Arabishirsula Cru 1.0 5.8.73 Derelict
limestone pasture SK 175655 35 p 1.65 0.85 7 7 1 0.09
154. Achrelun ninus Comn u 0-5 d 20.9.73 Cindersinderelict railwaysiding SK 261653 90 p 11.45 3.70 1 5 1 11.47
155. Ar,neria inaritima Plu I <0-I1 24.7.75 Derelict
nmineral working NZ 756425 15 p 5-50 1.30 2 1 3 0.03
156. Arte,nisia absirnhiu,n Com s 0.5 d 8.10.73 Derelictrefusetip SF 593014 60 p 1.17 0.70 4 1 1 0.10
157. Arfe,nisia vulgaris Com s 2.4 d 8.10.73 Derelictrefuse tip SF 593014 90 p 1.34 0-45 4 1 1 0-12
158. Ariniaclaun Ara 1.7 14.8.73 Mixeddeciduous woodland SK 231748 40 p 8-10 5-85 1 1 1 4-54
159. Asternovibelgii Comn w <0-I d 7.11.74 Urbanwasteground SK 349864 100 p 7.80 0.60 9 4 3 0.42
160. A rp elaon Sol <0-I 5.10.73 Wasteland SK 523650 90 p 16.50 16.50 1 1 1 0.08
161. Baldelliaan,cukoides Ali <01I nm 26.7.74 Pondnmargin SK 595195 10 p 2.10 1.30 2 1 1 0.45
162. Baillotaigr Lab s 0.2 g 29.6.74 Hedgebank TF 670410 70 p 2.25 1.55 3 1 1 0.73
163. Bellisperenrnis Comn u 3.0 g 22.6.72 Cattlepasture SK 160662 10 p 1.70 0.75 4 1 1 0.09
164. Beil erecta Unmb s <0- I nm 23.10.74 Marshland at margin oflake SK 437863 65 p 2.00 1,50 1 1 1 0.49
165 Beto,,ica offici,tialis Lab s 0.6 g 31.10.72 Cattlepasture SK 224745 35 p 2.60 1-75 3 1 1 1-37
166. Brioiadiica Cuc <0-I 288.73 Hedgerow S5F99010 250 p 9-00 9-00 1 1 1 13-37
167. Cailtha palu~stris Ran 0-6 nm 2.7 73 Marshatriverside SK 243753 25 p 2-SO 1-30 2 1 1 0-99
168. Canaua aioi Can, w 0-2 w 18.1174 Mixeddeciduouwooxdland SK 530652 8S p 2-10 1-20 4 1 1 0.06
169 Canauartnioi Cam u .1 g 15.10.74 Limestone screeandgrassland SK 155732 25 p IS1 0-50 2 1 1 0-07
170. Cadiiiifeus Cru 2.4 nm 22.6.73 Sha~dedroadside SK 175887 30 e 1.30 0-55 4 1 1 0.13
171. Cadiii rtni Cru 1.9 nm 4.7.75 Smallmarsh SK 285613 45 e 1.70 1-15 4 1 1 0.60
172. Carduusacanthoides Comn s <0-I d 3.10,73 Disturbed roadside SK 223743 75 p 5-15 1-75 9 4 1 2-49
173. Care.,catifoi,-nis Gyp s 0.6 nm 31.8.73 Pondmargin SK 454644 lOS p 3S50 1-65 3 1 1 I'll
174. Cae aplai Gyp n <o-I n 31.7.74 Montane grassland NY 839283 IS p 2-65 1-10 3 2 1 0.51
175. Carexde,nissa Cyp n 0.2 nm 26.7.73 Shallowpoolsinfloor ofoldquarry SK 490785 15 p 3-70 1-45 3 2 1 0.66
176. Care,cechi,iata Cyp n 0-1 nm 29.7.73 Streams~ide flush SK 18S810 25 p 3.70 1-60 3 2 1 0.83
177. Carerfacca Cyp 2.3 g 15.8.73 Derelictpasture SK 289818 25 p 2.55 1.45 2 1 1 0-37
178. Car-er laev-igata Cyp 0-3 nm 26.7.73 Wetcattlep~sture SK 362774 65 p 3-5 1-25 3 1 1 2.69
179. Car,er igra Cyp n 1.4 nm 15.873 Snmall
Sphagnum~ boginacidic SK 258777 40 p 2.65 1-95 2 1 1 0-81
180. Car,ertr-tbae Cyp <o-1 nm 16.8.73 Ditch [grassland SK424880 75 p 4-65 2-10 6 2 1 1-28
181. Carer.ovalis Cyp n 0.3 nm 28.8.73 Dampgrassland SE59501 SS5 p 4-SO 2.00 4 2 1 0-50
182. Cae aie Cyp n <O-1 nm 1.8.73 Damplimestone pastre SK 175745 25 p 1.55 0-8S 2 1 1 1188
183 Carerpaiculata Cyp s 0.2 nm 7.8.73 Carr SK 583579 105 p 3-SO 1.75 6 2 1 0,82
184. Cae, uiai Cyp n <0-1 nm 1.8.73 Dam,plimestone pasture SK 175745 20 p S-SO 0-75 4 1 1 1-46
185. Cae, sYihatica Cyp s 0-4 w 26.7.73 Wetcattlepasture SK 362774 35 p S.00 I-SO 3 2 I 1-65
186. Carlinavulgaris
187. Centaurea
188. Centaurea
nigra ~
scabiotsa
Co
Co
Co
0.3
3-7
0-9
g
g
1.1075
19.9.73
6 10.72
Limestone
Roadside
Limestone
otcrop atedgeofquarry SK 550789
wasteland
SK 240758
SK 154732
35
SS
60
p
p
p
3.20
3-85
10-25
1-20
1.70
2-40
5
2
9
4
4
4
3
1
1
1-53
2-55
7-46
189 Centrath,is ruber- Val <0-I 10.8.73 Cindertip SK 293984 SS p 7S50 1-65 9 4 1 1-66
190 Ceatunfttiii Car 9.2 g 17.7.73 Limestone sheeppasture SK 135825 20 p 0.91 0-75 1 1 1 0-16
191. CheohIu tueti Unmb s 0.2 g 7.11.74 Disturbed roadside SK 223743 65 p 5-20 1-35 6 1 1 1,44
192 Chawnaenerion angusti/blhu,n Ona 12.6 d 10.8.73 Shadedroadside SK 297988 75 p 12.60 0.46 9 4 1 0-05
193.Chelidot,,iurn in'Pap <0-1I 30,8.74 Gardenwall I F 660240 60 p 1-35 0-55 2 6 1 0-81
194. Chnpdunbn~hniu Che <0-I d 8.10.73 Derelict
refuse tip SEFS93014 40 p 1-95 1-80 1 1 1 1-91
195. Chn,santhemum' leucanthe,nu,n Co 3.2 20.7.73 Derelict
limestone quarry heap SK 505827 45 p 2-40 0-50 5 1 1 0-33
196. Chr,isanthemum,n,naxi,iiu, Co <0-I d 20.9,73 Cindersinderelict siding
railway SK 261654 SS p 2-65 I-OS S I I 0-31
197. Chr,vsosplenium oppotsidtifliun Sax w 1-3 nm 15.6.73 Streanmnmargin SK 531831 10 p 0-66 0-46 2 1 1 0-04
198. Cir-siu'nacuo o <0-I g 24.8.73 Roughgrassland in golfcourse SK 546828 15 p 25.00 1-75 9 4 1 3-71

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J. P. GRIME et al. 1053

Colour (9)Intates
gecrcicaticn aftercdrystoragec(Il)

E
a cO Se~ ~
~~~~~~~~~~C 20 IC cae
Response
eaaeResponsectoalght (I5)

cc 3cmonths 6cmonths 12cmonths >1cmacth RangecS13) Light Shade Dack


/
~>5u %a c P (12) c cc Curve 14) P(52) % %c a %c c

S 5YR/5/6 84 6 88 4 92 4 89 5 90 3 d 7 31 2 II d 76 6 83 4 541 58
4 IOYR/8/2 96 5 100 3 100 2 96 2 100 2 d <5 35 1 11 d 100 2 ISO 2 98 28
I IOR/3/4 78 53 94* 7 96* 12 92 7 92 5 d 7 31 5 TV d 86 7 94 6 82 6
I IOYR/7/3 92 8 91 4 95 4 96 5 99 3 d <5 30 3 TV d 96 3 94 2 94 49
2 I OYR/7/2 90 10 92 11 92 11 100 4 - d 6 34 2 11 d 90 7 88 7 16*
4 7.5YR/7/2 0 40* 25 50* 32 37* 19 72 17 d 9 24 5 11 d 94 53 90 13 22*
I 5YR/5/6 95 50 93 7 500 6 91 5 96 4 - - - - d 79 6 93* 4 85 5
2 7.5YR/6/2 83 53 85 15 87 13 93 7 84 55 4 7 35 2 15 d 91 8 90 4 87 4
4 IOYR/7/1 8 8 11 50 39 48 19 3 20 - - - -
I 2-5YR/3/2 78 4 188* 3 99* 4 97* 8 500* 4 d gccccicatonc-0
2 IOYR/6/3 94 8 97 5 97 4 500 5 98 4 d <5 33 3 51 d 100 4 108 4 588 58
4 7-5YR/8/2 77 8 88 9 88 9 74 7 98* 2 4 <5 36 1 55 d 588 8 87 8 39* 8
1 2-5YR/3/6 588 3 99 3 97 3 588 4 98 2 4 7 33 3 II-IV d 96 4 98 3 24* 8
I 7-5YR/6/2 99 12 99 8 96 6 97 6 100 4 d 6 28 7 II-IV d 85 7 92 5 87 5
2 97 4 92 3 99 3 92 3 92 3 d 50 32 3 551-TV d 93 4 88 4 90 4
2 7.5YR/6/3 69 32 17* 19 lo* 15 32* 18 18* 18 - - - -
2 7.5YR/6/2 85 19 98* 6 94 5 - - 97* 4 d 7 33 4 ST d 86 7 82 7 98 8
2 IOYR/7/3 98 5 95 4 98 3 98 3 96 3 d 8 30 2 IT-TV
2 IOYR/8/2 97 7 100 3 98 3 98 4 100 3 d 6 35 2 ST d 98 3 98 3 98 4
I 5YR/3/3 78 2 84 2 77 2 96* 2 92* 2 d 8 34 2 ST d 96 2 84 2 38 3
2 52 20 84 16 74 20 96 9 62 22 d 8 26 7 III-IV d 92 6 92 4 90 3
2 7.5YR/4/4 100 19 97 6 100 6 94 6 96 7 d 11 30 5 1
I 5YR/6/3 100 3 97 3 98 3 99 3 98 3 d <5 35 3 ST d 100 3 98 3 58* 3
2 7.5YR/7/4 85 11 60* 7 56* 6 88 4 52* 3 d 6 31 2 TI d 84 S 84 S 78 S
2 IOYR/7/2 95 4 96 3 98 3 100 3 96 2 d <5 34 2 II d 100 2 100 2 98 2t
I 5YR/3/3 100 57 100 30 100 21 98 23 100 15 - - - - d 500 19 98 11 lo*
I IOYR/6/3 96 36 100 27 99 25 99 27 99 27 d II 22 22 I1I d 100 22 88 22 34* 23?
I IOYR/4/4 3 13 8 52 It 13 7 13 - - c 19 39 2 5 c 50 4 71* 3 40 3
2 38 32 25 30 56 57 8* 32 25 33 d 17 36 12 15 d 76 52 70 38 22* 41
4 5YR/3/3 82 9 87 7 77 12 65 9 73 8 d 7 35 3 15 d 84 3 89 2 70 3?
4 IOYR/5/4 100 2 100 2 100 2 100 3 100 2 d 11 36 2 1-11 d 100 2 100 2 50* 4t
2 7-5YR/6/2 53 9 83* 4 92* 4 100* 3 79* 3 d 10 31 5 11 d 94 2 98 3 88 3t
2 7-5YR/7/2 96 4 99 4 91 3 90 3 83* 5 d 10 31 3 55 d 69 7 94* 5 72 9?
2 80 23 91 14 95* 12 78 52 59* 7 d 7 25 4 55 d 93 23 92 8 70* 38t
2 2-5YR/6/6 93 5 95 2 89 2 82 2 91 2 d 6 30 2 55 d 96 4 100 3 76* 5t
I 5YR/5/1 93 4 94 5 95 6 91 5 100 - - - - d 100 6 100 5 20* 26
I 5YR/6/2 90 11 98 7 92 7 98 6 100 8 - - - - -
I 2-5YR/3/6 0 0 31* 25 84* 16 95* 15 d 13 29 5 55 d 97 14 93 12 1
I IOYR/7/3 100 5 96 5 100 3 100 3 90* 2 d <5 34 2 15

I 63 2 87* 2 87* 2 70 3 - - d 6 33 5 51 d 62 2 100* 2 68 4


I 82 7 77 5 89 7 88 5 - - d 16 37 2 1-55 d 60 7 89* 6 42 6
- - - 0 0 0- - - -
3 IOR/2/1 0 0 0 0 0 - - - -
4 5YR/3/2 0 0 0 0 - - c 6 28 3 5
2 IOYR/4/2 34 13 35 19 37 17 36 17 - - - - - d 34 20 38 17 2*
I IOYR/5/4 4 0 2 0 4- - - -
I 7.5YR/8/4 4 I I 3 4 - - - -
3 IOR/3/2 0 0 0 0 0- - -
2 5YR/2/1 0 0 0 0 0- - - -
4 7,5YR/5/4 0 0 0 0 0- - -
3 9 20 7 15 8 19 6 17 12 30 - - - -
I 5YR/2/1 0 0 0 0 0- - - -
I 8 6 10 14 18 5 20 5 8 3 <5 37 1 55 s 00 2 100 1 100 I
I Black 4 38* 16 30* 15 53* 18 20* 13 d -0
germirnation d 25 17 51* 56 P*
I 2.5YR/3/4 80 33 84 6 90 5 99* 5 96* 5 d 12 30 3 5-55- - -
I 2.5YR/3/4 95 11 100 4 95 3 97 3 100 3 d 10 31 5 TV d 100 3 100 3 42* 7
3 60 42 9P* 11 91* 16 100* 17 - - - - - - d 90 18 94 16 0*
5 2-5YR/4/4 80 6 86 5 92 4 64 5 - - d 6 29 3 I-IV
3 2S5Y/4/4 78 6 89 5 99* 4 98* 4 100* 3 d 12 31 3 III-IV d 95 4 99 14 12 5
3 2.5Y/3/2 67 4 94* 4 89* 2 96* 2 90* 1 d 7 34 2 55 d 90 2 97 2 40* 2t
2 5YR/6/6 0 0 0 - - - -
S IOYR/4/3 90 11 91 8 98 3 84 7 98 6 d 17 37 2 IIs-IV d 92 7 92 6 2*
2 5YR/4/2 12 33 100* 28 53* 29 91* 27 10 28 c 7 29 3 S-ST d 52 26 42 26 0*
3 7.5YR/6/4 4 5 12 5 - - 13 1 10 12 - - - -
I 5YR/2/1 - - 3 23* 5 46* 5 -. - d IS 36 3 5 d 35 6 81* 6 49 6
I 7-5YR/6/2 100 11 95 7 96 5 99 5 96 3 d 6 35 6 III-IV d 100 5 100 5 100 S
3 91 8 90 8 86 7 84 8 78 8 d 10 24 8 51 d 77 8 77 8 0*
I 5YR/4/2 79 12 77 13 88 9 59* 11 88 10 d 12 25 7 s-IV
2 m 84 8 89 9 90 6 88 7 - - d 14 33 5 TV d 86 10 83 10 3*
I IOR/2/1 17 12 5 13 31 25 19 13 - - - - - -
I 5YR/5/6 0 0 0 lo* 9 - - 12 28 5 sV
I 5YR/4/4 99 6 76* 6 96 5 93 8 92 5 d 10 28 6 I-TV d 81 6 74 7 20* 6?
I 5YR/5/6 100 4 100 3 99 3 97 4 79* 6 d <5 30 2 551 d 91 4 90 4 59* 6
I IOYR/3/4 98 15 93 9 92 9 97 8 90 8 d 12 28 5 551 d 99 6 100 6 66* 7
I 7.5YR/7/2 80 7 79 4 70 3 86 3 - - d II 35 2 551 d 97 4 92 3 14* 4
I 5YR/4/3 10 20 40* 11 52* 18 40* 10 37* 11 - - - -
I 5YR/4/1 23 13 6* 16 18 17 10 12 32 15 d 16 27 10 S-ST
I 5YR/3/4 0 0 0 0 21* 19 d 21 36 9 II-TV
I 7.5YR/5/6 92 22 94 11 99 14 78 18 97 13 d 17 32 7 S-ST d 88 19 97 18 0*
I 5YR/3/3 8 23 10 22 30* 17 63* 18 49* 16 d 16 28 7 s-IV d 50 13 22* 14 0*
I 2.5YR/1/4 0 0 0 0 0 - - - -
I 7.5YR/3/2 0 2 I 12 14 46* 12 19 37 4 S-ST 60 9 72 9 6* .?
I 5 11 26* 17 38* 15 26* 17 97* 7 d 19 31 6 5 d 94 11 77* 11 0
5YR/4/6
I 5YR/4/6 26 15 52* I/ 48* 10 96* 13 71* 15 d 16 30 7 sV d 84 12 78 13 0*
I 5YR/3/4 0 0 0 0 0 - - - -
I 5YR/4/2 8 9 28* 13 32* 13 42* 15 85* 9 d 14 33 6 sV d 86 10 86 10 0*
I 2.5YR/4/4 70 47 I00* 40 100* 29 - - I00* 31 d 10 29 17 S-ST d 100 22 95 21 22P
I IOYR/4/4 0 0 0 0 45* 33 d germination -0 d 60 37 47 34 0*
4 7.5YR/8/0 97 4 86 3 82* 4 88 4 - - d 8 30 4 I-SIT- - -
I IOYR/8/2 36 35 64* 5 60* 8 80* 5 90* 5 d 11 34 3 5 d 90 3 95 3 3*
I 52 4 53 3 70 3 45 9 68 4 d 13 34 3 55 d 64 8 64 5 18* 6
2 83 16 88 9 78 11 78 10 - - d 8 25 4 s-IV d 86 8 70 8 76 8
2 2.5YR/4/6 98 7 98 5 99 3 98 4 97 3 d <5 30 2 55 d 100 3 100 2 95 4?
3 5 17 0 8 13 15 23 - - - - -
I IOYR/5/8 79 6 95* 4 97* 5 94* 5 94* 4 d <5 35 1 55 d 93 6 73* 4 7*
2 5YR/2/1 24 25 - - 10 19 25 29 10 20 - - - -
I 5YR/2/1 95 7 91 7 89 6 99 6 - - d 10 22 5 55 d 94 9 94 7 3*
3 m 100 4 98 4 99 3 84* 3 99 3 d 9 31 2 S-ST d 100 3 100 3 83* 3?
3 m 92 8 80 10 70* 10 80 11 - - - - - -
I 2.5YR/2/2 94 11 5I* 17 31* 17 2* 96 15 d germination- 0
I 32 10 50 7 62* S 5I* 6 - - d IS 29 4 II1 d 53 7 49 6 3*

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All use subject to JSTOR Terms and Conditions
1054 Germination in a localfiora
characteristics
APPENDIX 2-continued
Dehiscene Dispersue

Seedcllection
E ZNational - dh
Species U. Date Habitat Gridref (rn~g)
Cm 6.8 d 28.873 Wasteland SE 610003 60 p 14.35 1.55 9 4 1 1.17
199.Cirsim rvns
200 Cisu ausr o 1.8 m 15.8.73 Sml panm o naii SK 258777 90 p 12-65 1.55 9 4 1 2.00
o 3.0 d 10.8.73 Cindertip Igra~ssand SK 293984 90 p 25-95 1.50 9 4 1 2.64
201. Cirstum vugr 0.49
202. Clhnopodirum ugare Lab 0.3 g 16.10.72 Roadsidebank SK 149734 55 p 1.50 1.10 2 1 1
r <0.1I 11.7.74 Dam,plim,estoe otcrop SK 135825 30 p 1.15 1.00 4 1 1 0.32
203 Coheraofiiai
Umb s 0.2 m 16.8.73 Lakemargin SK 424880 125 p 3.40 1.60 6 1 1 2.25
204. Coniummaultu 2.26
205. Conopodium majus Umb u 1.7 g 15.8.73 Derelict pasture SK 152730 40 p 4.60 1.35 6 1 1
206. Covlai aai Lii 0.4 w 3.10.73 Screeitnhazel crub SK 224743 15 p 8.10 7.00 1 1 1 16.02
o <0.1I d 23.10.74 Cinder tip SK 437862 50 p 5.05 1.50 3 1 1 34-69
207 Covluu rvni
Cm <01I 23.8.74 Wetoerhanging cliff SK 119869 60 p 11.75 0.60 9 4 1 0.69
208. Crepispaludos
209. Daucuscarota Umb s 0.4 25.10.74 Railway cinders SK 577617 65 p 2.65 1.90 6 5 2 0.88
210. Digitalispurpurea c 1.3 w 10.8.73 Shadedrie bank SK 297990 00O p 1-00 0.50 2 1 1 0.07
C <0.1I d 17.9.75 Colliery heap SK 261633 55 p 1.35 0.95 4 1 1 0.32
211 Diplotaxis tenuifolia
s 0.1 d 20.9.73 SK 261654 125 p 4.50 1.40 5 1 3 2.26
212. Discsflou Dip Deeitriwysdn 0.27
213. Donicumpardalianches Com w <01I 4.7.75 Riverbank SK 261633 60 p 5.00 1.05 5 4 3
Dro n <01I m 15.9.74 Bogneareservoir SK 259931 15 p 1.52 0.26 7 7 I 0.01
214 Droerartudifolia 2.76
215. Echium vlgare Bor s <0.1I d 28.8.73 Railway embatnkment SE 605004 60 p 2-80 1.80 3 1 1
216. Elohrs auti Gyp 1.0 nm 16.8.73 Marshm,arginal to ake SK 435863 35 p 2.40 1.45 4 1 1 0.96
217. End,monnonscriptu~s Lili 9.1 20.7.73 Mix~ed deciduouwodland SK 324813 35 p 2.80 2.05 1 1 1 6.17
218. Epilobiuadencaulon Ona 1.6 d 7.8.73 Disturbed wasteland SK 375750 75 p 1.25 0.50 9 4 1 0.06
219 Epilobumhirsuhtumr Otna 3.2 m~ 3.10.73 Marshatriverside SK 151731 115 p 4.05 0.50 9
9
4
4
1
1
0.05
0.13
220 Epilobiumr monta,num Ona 4.7 w 5.8.73 Edgeofdeciduouwodland SK 176654 40 p 7.00 0.50
221. Epilobiumr nerterftoides Ona n <01I m 31.8.73 Wetfloorofdisusedstoequarry SK 209868 10 p 3.70 0.25 9 4 1 0.02
222. Epilobiumpalu~stre Ona 0.5 m~ 26.7.73 Snmallpond SK 487789 35 p 7.30 0.50 9 4 1 0.04
223 Erohrmanutflu Gyp n 1.0 m~ 2.7.73 Moorland SK 258777 40 p 60.50 1.35 9 4 1 0.44
m~ 1.02
~
224. Erio~phouvagintumn Gyp n 0.4 22.6.73 Moorland SK 260877 40 p 21.72 1.21 9 4 1
225. Eupatorium cannbinum Com s 0.3 nm 20.9.73 Riverbank SK 260657 75 p 3.50 0.65 9 4 1 0.27
226. Filipendulalai o 1.5 m~ 16.10.72 Riverterrace SK 156731 90 p 4.20 1.90
8.90
6
1
1
1
3
1
0-99
0.31
227 Frgai esc~a Rs 0.8 22.673 Derelict stonequarry SK 209868 15 p 10.50
8.23
228. Gailega officinalis Leg I <0.1 d 2.10.75 Wasteland SF 560047 150 e 3.75 1.20 2 1 1
229. Gaeboonltu Lab 3.7 w 296.73 Mixeddeciduouwoodland SK 530832 40 p 3.50 1.85 3 1 1 1.98
230. Galiumr Rub <01 g 15.10.75 Disued railballast SK 497553 70 p 1.50 1.25 1 1 1 0-67
molugo 0-91
231. Galiumpalustre Rub 1.5 nm 27.10.72 Marshat m,argin of1ake SK 437863 20 p 1.65 1.50 1 1 1
232. Galiumr s~axatile Rub n 6.0 g 29.8.72 Derelictacidicgrassland SK 176654 15 p 1-50 2.20 2 1 1 0.56
233 Galium sterneri Rub n 2.0 28.8.75 of
Outcrop sgar limestone NZ 838283 25 p 1.10 1.00 I I 1 0.39
234 Galium rermRub 09 g 16.10.72 Derelictlineson grassland SK 149733 55 p 1.50 1.40 1 1 1 0.40
235 Gentianella amarella Gen 0.4 19.9.73 Broken turfon lietn outcrop SK 155732 15 p 0.70 0.60 1 1 1 0.12
236 Geaimpaes Ger 0.4 g 15.8.73 Roadverge SK 259784 55 e 3.55 2.15 2 1 1 6.37
237 Geranium Ger I <01 g 6.10.72 Derelictlinmestone grassland and SK 155732 25 e 3.30 2.35 2 1 1 1.02
saguineumn
238 Geumrivale Rs n 02 m 237.73 Dampareainhazel crub scree SK 232748 40 p 10-50 1.35 8 5 3 1.22
239. Geumr banum os 1.4 w 23.773 Shadedroadside SK 223743 40 p 8-50 1.30 8 5 3 0.73
240. Glechoma hederacea Lab 1.7 19.6.73 Hedgerow SK 668918 20 p 1.90 1.10 2 1 1 0.69
241 Heracleumr Unmb w <01 d 15.8.75 Derelictgarden SEF546007 250 p 11.90 7.70 4 1 1 5.73
mategazzianum 5.52
242 Heracleusphondv'lium Unmb u 8.5 g 16.10.72 Roadside SK 149733 125 p 8 60 6.30 4 1 1
243 Hircun ioel Com u 4.5 25.772 Stabilied limestone sce SK 175655 15 p 7.65 0.50 9 4 1 0.15
244. Hi'pericum hirsutum Hyp 0.5 g 19.9.73 Derelictlinestone grassland SK 155732 70 p 1.30 0.50 5 1 1 0.07
245 Hi'perncummntanumn Hyp 0.4 g 8.10.73 Derelictlim~estone grassland SF 595008 60 p 0.96 0.36 5 1 1 0-07
246. Hi.per,cumpetrforau Hyp 0.9 g 25.10.73 Deeitlmstn rsln SK 155732 60 p 1.30 0.55 5 1 1 0.10
247 Hipericumpulchru'n Hyp n 0.2 g 31.10.72 Senmi-derelictacidicpasture SK 224744 45 p 1.25 0.50 5 1 1 0.08
248. H'pericumtetrapterumr Hyp s <0.1 3.10.73 Marshland nearriver SK 241754 50 p 0.90 0.44 5 1 1 0.05
249. Hiohei radicata Comn u 2.8 g 24.8.72 Railway cutting SK 153629 40 p 13.70 0.75 9 4 3 0.96
250. Inulaconi,a Com, s 0.8 26.974 Disued railballast SEF596004 75 p 13.80 0.50 9 4 3 0.26
251 Irspeuaou Irn <01 10.10.75 Siltedpond SK 442697 95 p 8.40 5.80 3 1 1 46-67
252. Juncus ariculatu Jun 1.4 nm 5.10.73 Shallowpoolsindistrbedwasteland SK 262807 40 p 0.70 0.33 1 21 00
253 Jucuscoglomeratus Jun 0.3 nm 18.10.74 Wasteland SK472822 90 p 0.69 0.25 2 .2
254. Juncus effusus Jn 3.4 m 3 10.73 Streanmside SK 262807 90 p 0.52 0.25 2 1 1 00
255 Juncus infleus Jun 0.3 m 23.10.74 Marshland adjoining lake SK 437863 40 p 0.65 0.42 2 .3
256. Jucssurou Junn 0.5 g 3.10.73 Moorland SK 288819 30 p 0.80 0.50 2 .5
257. Lath',rus mntanu~s Leg 0.6 g 24.7.73 Railway cutting SK 390398 25 2.55 2.20 1 249
258. Lath 'rspratensis Leg 2.3 g 10.8.73 Cindertip SK 293985 75 3.25 2.75 1 285
259. Leontodon Com u 2.1 24.8.72 Derelictlimestone quarry heap SK 165662 30 p 9.75 0.80 9 4 3 0.7
auntumnalis
260. Leontodon hispidus Comn s 4.0 g 24.8.72 Derelict limestone grassland SK 155732 35 p 12-45 0.95 9 4 3 0.85
261. Linariavulgaris Scr s 0.7 d 8.10.73 Oldrefuse tip SF593014 55 p 2.10 1.76 7 .4
262. Lou oniuau Leg 5.6 g 16.10.72 Derehict lim,estone grassland SK 155732 IS 1.85 1.50 1 .7
263. Lou liiou Leg 0.5 nm 8.10.73 Hollowindisturbed wasteland SK 682994 35 1.13 1.00 I I 1 0-4
264. Luul camrpestris Ju g 5.7.73 Acidiccattlepasture SK 198797 10 p 1.65 1.05 I .4
. 1.00 I .7
265. L~uzul muliflraI~ Jun n J 14.8.72 Derelict grassland SK 548915 30 p 1.60
266. Luzlapilosa Jun 0.7 w 22.6.73 Oak woodland andscrub SK 209868 20 p 1.35 1.00 I I 1 08
267. Luuas'vvtc Jun n 0.3 w 20.7.73 Oak woodland SK 323813 55 p 1.70 1.05 1 .9
268. L 'chnisf1os-ecucli Car 0.1 nm 14.8.73 Marshland adjoining river SK 243754 50 p 0.88 0.75 4 1 2 0.21
269. L epseuoau Lab s 0.5 nm 4.10.73 Pondnmargin SK 473645 65 p 1.50 1.25 3 1 1 0.28
270. Litrmsaiai Lyt s <01I nm 15.10.73 Edgeoflargeditch SF 713159 90 p 1.15 0.50 6 1 1 0.06
271. Mfelilotusaltissima Leg s 0.2 d 18.10.74 Railwayenmbanknment SK 468821 90 p 3.10 2.75 2 1 1 3.72
272 Mfentha aquatica Lab 1.4 nm 27.10.72 Marshland adjacent tolake SK 437863 50 p lOS5 0.60 2 1 1 0.14
273. Mfentha arensis Lab s 0.4 nm 8.10.73 Reedbed SF 596008 35 p 1.00 0.90 2 1 1 0.27
274. Mfercukialis perennis Eup 7.0 w 14.6.73 Mixeddeciduoswoodland SK 752731 30 p 2.70 2.60 1 1 1 2.15
275 Minatia erna Car I 1.0 31.10.72 Leadm,ine poilheap SK 223742 10 p 1.00 0.65 4 1 2 0.08
276. Mfycelis mra~~lis Conm s 1.1 15.8.73 Shadedlimestone cliff SK 151731 60 p 8.90 0.90 9 4 3 0.34
277. Mfvsotis Bor 1.5 nm 25.10.73 Riverbank SK 151731 30 p 1.95 1.20 4 1 1 0.28
scorpioides
278. Mf'vsostis Bor I 0-5 w 2.7.73 Hazelscrub SK 152731 30 p 2.00 1.40 4 1 1 0.56
s~'vIvtica SK 295988 80 20.80 3.00 5 1 1 35.01
279. MVrrh isodorata Unmb n 0.5 g 10.8.73 Shadedroadside p
280. Oenotherapariflora Ona I 0.2 d 3.11.75 Disturbed wasteland SK 583864 75 p 1.80 1.30 3 1 1 0.56
281. Origanumr vulgare Lab 1.3 g 16.10.72 Derelict limestone grassland SK 155732 55 p 1.00 0.70 4 1 1 0.10
282. Oxalisacetosella Ox~a 3.5 3.7.73 Hazelscrub SK 152731 10 p 2.55 1.60 4 1 1 1.01
283. Parnassiapalustris Par n <01I nm 15.10.74 Derelict limestonegrassland SK 162728 20 p 1.60 0.60 7 1 1 0.31
284. Petasites hibridus Comn w 08 nm 29573 Shadedriver terrace SK 245754 25 p 11-50 0.55 9 4 1 0.26
285 Pimrpinellamajor Unmb s 0.2 g 17873 Derelictlimestone quarry heap SK 144731 85 p 3 55 1.45 6 1 1 2.12
286. Pimpinrella Unmb S1. g 25.10.73 Broken turfon lim,estoneoutcrop SK 155732 65 p 2.65 1.65 2 1 1 1.19
sairg 0.03
287. Pigicl gradiflora Len I <01I nm 23.8.74 Wetcliff SK 119870 IS5 p 0.85 0.37 5 I 1
288. Pinguicula vulgaris Len n <01I nm 24.7.75 Derelict mineral working NZ 756425 10 p 0.85 0.30 5 1 1 0.02
289. Platagocoronpus Pla <01 2.8.74 Snmallletone outcrop SK 557682 10 p 1.15 0.65 2 1 1 0.15
290. Plantagolaceolata Pla 9.6 g 27.10.72 Wasteland SK 435863 35 p 3.55 1.75 2 1 1 1.90
291. Platago maj.or Pla 5.6 d 27.10.72 Tranmpled wasteland SK 435863 15 p 3.09 1.82 2 1 1 0.24
292. PIatago media Pla 0.4 g 7.11.74 Floorof derelict limestone quarry SK 132733 25 p 2.60 1.95 2 1 1 0.40
293. Polemonium, caeruleum Pole I 0.1 g 24.8.72 Ungrazed lim,estone grassland SK 165660 60 p 3.00 1.20 3 1 1 0.77
294 Pol'gala vlgaris Poly 0.8 g 31.7.75 Grazedlimestone grassland SK 174745 20 p 2.50 1.50 2 6 2 1-72
295 Poaoeo Pot 0.2 nm 8 1073 Ditch SE 594012 <10 p 2.25 1.55 2 1 1 1.12
olgnflu 2 1 1 0.58
296. PotentillaerectaRo 2.8 g 3.10.73 Acidicroadverge SK 290819 20 p 1.55 1.05
207. PoterntiIa
tabernemntan~i Ros I <01I 27.7.72 -im,estone ou~tcrop SK 088723 10 p 1.80 1.35 2 1 I 0.91
298. Poternumsaguisorba o 1.5 g 23.773 Deeitlmson he atr SK 223743 30 p 3.15 2.14 2 1 I 4.11
299 Primula veris Prn 0.5 g 17.8.73 Floorofderelict limestone quarry SK 143731 20 p 1.65 1.35 3 I I 0.69
Prn <01 247.74 Ashwoodland SD 274740 5 p 1.75 1.35 3 1 1 -
300. Primulat vulgaris
301 Prunella vulgaris Lab 2.7 g 31.10.72 Disturbed wasteland SK 230748 IS5 p 2.00 1.00 3 1 1 0.73
302. Puiai i,etrc Com s <01 I nm 8.10.73 Marshland SE 596011 40 p 9.70 0.35 9 4 3 0.06

This content downloaded from 69.167.65.194 on Mon, 2 Dec 2013 13:19:33 PM


All use subject to JSTOR Terms and Conditions
J. P. GRIME et al. 1055
Germinule
Colour(9)
3 ~~ ~~~~~grmnto afterdrystorage(I11)
E
to
~~~~~~~~~~~~~~~~~~~~~Response
~~~~~~~3 5 IC 20 IC temperature 1 5)
Responeeetolight
a 3 3 months 6 months 12 months >1I moth Range(13) Light Shade Dark
5
a >C.5 % 5 0 %5 0 0 p(12) IC t p(12) %
Cureae(14) 5 0 %

1 7.5YR/5/4 I 3 7 3 9 4 4 - - - -
I 7.5YR/8/4 91 8 50. 9 88 6 79 7 - - d 12 33 4 11 d 90 11 86 9 36* 16
a h3 4 51 3 73 2 70 3 88* 2 d 11 30 2 1-11 d 71 4 75 3 19* 4
I IOR/3/2 98 17 97 22 92 7 100 11 97 9 d 7 30 5 11 d 96 8 94 7 4*
I IOR/3/2 92 23 92 5 98 4 98 4 - - d 10 28 2 II-IV d 93 S 100 3 54* 4
3 v andm 10 19 34* 18 35* 18 41* 16 68* 11 d 10 30 9 IV d 94 14 88 13 70* 11
I 5YR/3/3 0 0 0 0 0 - - - -
I 2.5YR/4/6 0 0 0 - - - -
2 5YR/2/2 16 20 10* 17 40 4 60* 1 30 3 - - - -
3 7.5YR/7/4 76 8 62 10 68 8 30* 15 76 8 d 14 33 5 11
5 a 34 10 30 10 42 10 49 8 45 9 a 6 29 4 11 d 42 3 64 4 0*
2 2S5YR/2/2 99 8 91 7 99 7 98 6 97 6 d 10 29 6 Itt-tV d 96 7 100 6 56* 7
a 15 3 1* P* 6 9 0* - - - -
S 7.5YR/7/2 54 8 85* 5 60 6 48 6 - - d 11 30 S III-IV d 71 6 49* 7 5*
5 90 6 64* 12 50* 9 18* 7 100 5 - - - -
I 7.5YR/5/2 0 0 0 0 - - - - - -
2 a 2 3 12 4 5 6 - - - - - -
I IOYR/3/4 0 0 0 0 0- - - -
2 Black 0 0 0 0 0- - - -
I 5YR/3/3 100 3 99 3 97 3 100 2 100 3 d 8 36 2 It-tV d 98 3 98 3 82 St
I 2.5YR/4/2 94 4 93 4 98 2 96 2 98 2 d 12 33 3 II d 100 3 96 3 2*
I 7.5YR/5/2 98 6 100 4 98 4 100 4 100 4 d 8 22 4 II d 100 4 98 4 1
I 7.5YR/5/6 98 4 96 3 94 4 84* 3 80* 3 d 11 30 4 II d 95 4 100 4 19*
I 5YR/5/3 99 4 96 3 100 3 95 3 93 3 d 6 34 3 It-tV d 100 3 100 3 86* 8
I 5R/3/1 74 6 59 9 90* 7 58 7 - - d 16 31 4 1-11 d 72 8 57 7 0*
I a 96 5 88 6 95 5 91 7 95 7 d 10 33 3 It-tV d 92 8 82 7 31* 13
3 7.5R/2/0 63 36 38* 12 21* 13 61 18 - - d 23 30 6 II1 d 92 15 100 15 0*
I 5YR/4/2 50 20 48 20 29* 18 38 30 51 14 11 30 3 It-tV 80 5 66 .4 78 71
I a 97 6 67* 15 94 7 97 9 96 10 d 11 32 6 1 d 97 10 92 10 0*
I 7.5YR/5/6 I 70* 3 80* 4 98* 5 6 4 - - -
I IOR/3/1 0 0 0 0 0 - - - -
2 5R/2/1 100 6 95 4 96 3 97 5 - - d 9 29 4 IV d 100 4 96 3 100 3
14 34 II - - -
2 5R/2/1 6 24 2 11 16 42* 23 10 17 6 2
2 5YR/2/1 46 24 20* 2S 27* 28 9* 23 34 22 d 7 22 13 IV
2 5YR/2/1 88 11 96 8 88 9 45* 11 50* 6 d 12 29 4 IV
2 5YR/2/1 71 22 60 31 65 10 75 13 72 8 d 11 29 7 IV d 59 8 71 7 58 7t
I 5YR/4/6 0 0* 0 - - - -
I 5YR/3/2 8 5 92 34 86* 43 96* 27 20 11 6 33 2 II1 d 99 29 100 24 83* 3S
I 7.5R/2/2 54 16 36 12 - - - - 45 - - - - 96 3 92 4 54* 5
4 2.5YR/3/2 73 15 37* 12 57 14 53* 18 66 14 d 17 30 7 1 d 93 10 93 10 4*
4 7.5YR/4/2 79 16 73 13 71 10 66 14 84 7 d 10 22 5 II1 d 82 6 81 6 4*
I IOR/3/4 45 9 34 6 44 10 44 6 70* S - - - - d 32 5 36 6 0*
I IOYR/7/4 0 0 0 0 - -
I v n 0 0 0 0 - -
3 Black 47 13 83* 7 72* S 81* S 92* 5 d 6 27 3 II d 100 3 97 3 91 3t
I 7.5R/3/4 6 31 95* 6 98* 8 88* 7 95 d 12 29 5 1-11 d 94 7 99 6 H*1 -
I IOR/2/1 100 13 100 5 100 7 100 7 99 8 d 10 34 5 II d 100 6 100 6 98 9t
I 5YR/2/1 90 11 - - 97 6 97 7 - d 13 35 3 II d 90 10 92 9 P*
I 5YR/4/2 60 14 73 12 91* 9 76 12 81* 12 d 11 23 8 1 d 75 12 65 13 0*
I 7.5YR/4/2 25 21 54* 17 71* 15 44* 13 64* 14 d 20 38 5 IV d 38 13 44 12 0*
S IOR/3/2 99 9 95 6 72* 7 97 7 91 5 d 11 30 3 III d 97 4 97 4 81 4
S 5YR/3/1 94 14 98 13 100 7 31* 9 94 6 d 13 30 3 1-11 d 91 5 90 7 2*
I 2.5YR/2/2 48 15 8* 20 30 14 40 12 42 13 d germintion 0
6 5YR/6/6 98 12 100 8 98 8 100 7 100 9 d 17 31 4 II d 94 11 93 13 0*
O
6 7.5YR/5/6 91 2S 92 22 85 20 - - 81 23 d germination-0 d 72 14 2*
6 7.5YR/5/6 75 8 71 7 84 6 87 5 93 11 - - - - d 99 7 63* 8 7*
36 4 d 46 5 16* O
6 5YR/4/6 73 5 19* 5 72 4 55 4 71 5 d 14 II
6 2.5YR/3/2 99 5 100 4 100 4 100 3 100 4 d 10 30 3 II d 100 4 100 4 2*
I a 46 16 12* 7 8* 16 23* 6 lo* 19 - - - - 98 6 92 6 96 6t
I v n 57 20 34* 10 41 19 6* 9 O* <5 28 4 IV 90 5 100 4 96 4
S 2.5YR/3/4 42 5 79* 2 74* 4 50 4 75 3 d 18 32 3 III d 41 5 71* 4 9* St
S 5YR/3/4 80 7 90 8 95* 6 74 10 94* 7 d 12 32 4 IV d 98 6 95 6 7* *t
I IOR/3/1 6 10 34* 7 3 9 40* S 59* 6 - - - - d 66 6 68 6 0*
I v n 46 16 48 3 49 9 47 9 13* 2 <5 34 1 IV 100 3 100 3 100 3
a 12 4 6 4 7 3 23 3 8 3 7 37 1 II 96 2 97 2 92 3t
6 IOR/3/4 90 12 100 12 97 11 100 11 - - d 7 24 8 It-tV d 82 11 100* 10 71 12
6 IOR/3/4 99 9 97 8 100 8 100 7 - - d 8 24 7 IV d 98 7 100 6 100 lit
6 IOR/3/3 89 15 99 13 96 13 76 16 - - d 9 23 7 It-tV d 100 12 100 13 72* 16
6 IOR/3/1 94 19 98 14 96 14 95 11 99 1 d 6 20 10 IV d 100 16 91 14 0*

~
2 5R/3/1 81 9 71 11 88 7 88 7 97* 5 d 17 30 4 1 d 96 5 84 5 6*
I 5YR/4/4 0 0 0 - - 0 C 22 34 2 1 a 12 4 74* 3 2
I 5YR/4/6 60 9 89* 9 66 6 76 5 87* 11 d 19 37 3 1-11 d 65 6 68 7 0*
I a 0 I 2 6 6 1 <5 39 1 II 100 1 100 1 100 it
I lOR/4/4 17 10 P* 4* 2* 10 17 - - - - a 24 5 58* 4 2*
I 5YR/5/2 0 I 0 - - - a 36 40 2 III a 13 4 63* 3 0
2 5Y/5/2 0 0 0 0 0 - - - -
2 5YR/3/4 89 12 96 10 98 9 98 9 98 7 d 10 27 11 II d 90 9 92 7 78 9t
S IOR/2/1 98 5 94 4 89 4 92 7 94 5 d 10 24 5 1 d 89 5 97 5 30* 6
I 5YR/3/1 0 52* 29 64* 12 62* 10 66* 11 - - - - d 92 8 96 9 20*
I 5YR/2/1 88 2S 79 12 97 15 90 13 88 11 d 9 22 12 I-tV d 96 9 91 8 75* 91
3 5YR/3/2 0 0 0 0 0 - - - -
2 IOR/3/2 32 5 lo* 5 20 5 56* 4 - - d 22 34 2 II d 78 4 72 4 2*
I 5YR/3/2 89 4 95 3 87 3 91 5 95 3 d 13 33 3 1-11 d 91 4 96 4 90 4
3 IOR/3/2 0 0 0 0 - -
I 5YR/4/6 93 8 84 9 81 8 15* 11 90 8 d 13 33 4 II-tV d 92 8 87 6 lo*
3 83 1 2* O* O* - - C 6 34 1 II C 100 1 100 1 100 1
3 5YR/3/2 12 21 18 17 20 19 21 24 15 19 - - - -
3 5YR/2/2 16 24 26 38 22 21 15 17 - - - - - - 86 8 74 5 20* 6
I 5YR/4/2 69 19 74 18 55 18 74 23 83 15 d 13 29 10 II-tV d 81 18 82 17 0*
I 2 I 2 2 6 29 10 30 4 II - -
2.5YR/4/4
6 2.5YR/4/4 12 17 37* 5 50* 10 - - 40* 7 - - - -
6 37 14 46 5 62* 8 73* S 60* 5 - - - - d 64 5 88* 4 25* 6
6 5YR/4/3 20 4 6* 4 9 5 17 4 31 4 15 34 2 1 d 36 5 68* 4 0*
6 5YR/3/3 71 5 80 4 90* 4 97* 4 88* 4 d 15 33 2 1 d 81 4 87 4 8* *t
I 2.5YR/3/4 58 10 39 4 35* 8 39 8 70 7 d 7 27 4 1-tV d 18 7 72* 9 P~
4 IOYR/2/1 10 17 40* 10 25* 10 30* 8 - - - - -
2 IOYR/2/2 60 17 56 12 26* 11 - - - - - - - - 62 6 39* S 22* 6
I 7.5YR/5/4 48 29 40 21 49 26 62 24 57 24 - - - - d 54 27 77* 29 5* 33
I 7.5YR/5/4 3 I 54* 28 75* 37 50* 29 - - - - d 36 21 37 26 P*
3 5YR/4/3 44 21 56 24 82* 22 64* 16 86* d 8 30 6 11-IV d 95 8 96 7 75* 9
2 7.5R/2/2 0 0 0 0 0 gerination--0 100 21 96 19 2*
2 7S5R/2/2 86 31 85 21 75 19 - - 92 20 - - - - d 90 19 92 18 P* 1
6 5YR/4/4 91 7 97 6 98 5 85 7 88 6 d 13 34 5 1 d 73 5 59 5 2*
5 7.5YR/4/2 17 37 52* 7 28 6 28 6 60* 14 d 24 35 4 I-Itt d 27 6 24 6 0

This content downloaded from 69.167.65.194 on Mon, 2 Dec 2013 13:19:33 PM


All use subject to JSTOR Terms and Conditions
1056 Germination in a localfiora
characteristics
APPENDIx 2-continued

S~~~~~~dIIed" ~ ~ ~ ~ DeiceeDiprsl

306. lwe~
Resed~ Re <01 d 20.9.73 de~eIictall~ii~g
Ci~e~ 1S80d1.35l2c1i1n07 SK 291653 50 p

g 21.9.72 15 p 1550 1.00 3 1 1 0400


309. RanuculusekapisRa 4.6 Cate atueSK3089668 1.33
Pa 0.2 d 28.80.73 Mas adSE159501 753 p 3.15 2.65 3 1 1
304. Ranuncuuss faml 1 1 2.61
311. Rauclsrpn 0
82 82.10.73 ReedurbedwseadSK595009 120 p 8.405 2307 3
Ran 1315 3 1 1 1.10
312. Reseabutea Ress s 4O.3I d 27.0.972 Cindersainderlcaiiig SK 291653 75 p 370
395 2.15 3 I 1 1.133
313. Re~sedalueol Res s 1.1 16.8.73 Wahasteland~ Ik SK 436863 750 p
0.02
2.2 d 241.6.73 Derlithagradssland SK 3328748 50 p 0.50 0.25 3 1 1
314.umxacts Pola u 1 2.97
315. Rurex 0.86 g 16.873 Abandoedstoneqduarry SK 5308338 40 p 4*O0 2.80 2 5
cetosel Pol 1 1 1.46
316. Rurnex~ <01 g 3.10.73 Abandoead ratlwedgetrat
de k SF5851543 60 p 3195 1.650 4
crfispus.iPosCa 0.05
31II. Rumexhvdrlpathum Poa, 0.1 31.7.73 Reedbedl, s~~,eSK 183715 120 p 0.66 0275 2 1 2
1.32
1
~
318. Runxotsflu Dip 4.2 g 26.90.72 WatladSK 217653 40 p 2.60 31.50 9 3
o 1 1.62
319. cRumex <01 91.10.73 Mashatkmrin ofldakel pt SK 4602963 200 p 3205 2185 4 2
kcsangines GPo 3 1 1 0.11
3204.Saiapoubn Gyp 0.2 146.975 Cu~ltatedgresolSK 3463287 65 p 1150 0725
SKF610803 75 p 40.0 0.55 2 1 1 0.07
3215.Saniculaieuropaea Smb 0.4 281.8.73 Mixdecdoula odad 0.08
0.4I d 31.10.72 Wastbelad S5K23174 60 p 1.00 0.25 2 1 1
322. Sa~ponaiakfcinli Car s ~ateIadgofpt 2 0.03
0.8 Shadedliestoncree SK 155732 50 p 0.85 0.45 2 1
323. Saxirag hvnode S~ax 6.17.23
10 1.10 0.50 2 1 31 .3
3248.Sdabioa olmara Ci 1.<01 31.9.73 eelc lietn grassland SK 172804 p
g SK 151730 400 p 0.95 0.50 6 2 1 0.062
325.~ Seirpu lacustis Cypa <01 25.10.73 Shllowpool ainoldgravelapt
2.30 0.85 9 4 1 0.37
326. Scirps tqvticus cThm 032 m 810973 Marshland SK 596012 650 p
9 4 1 0.07
327. Scrjophlri qutc Scr 6304 248873 Mashang,dI~,d a SK5610823 90 p 7490 0.50
30.0
024 g 19.90.73 Ditre atln SK 2315732 60 p 9.60 050
1 9 4 1
3228 cohuai noos Scr 1 1.09
329. Sedumkacr <0.1 g 810.73 Limpgtneoucros K
5E597312 65 p 4085 2 10 4
Crab 4 1 1 09
Cra <03 d 718.73 Lietn otrpi ol5uar K 5076046 65 p 1148 1 50
3304.Sedumalbum 10.5 4 1 1 0069
eehu Cra 23.7073 Lieson riwa blls SK2237430 60 p 1.25
331. Seu 370.I
4 4 1 0287
Com u 013 1681072 MashanK 555712 50 p 1.40 0.80
3326.Secioauatcu
02 d SK525773 55 p 1905 170 4 1 1 1.09
333. Silau slaus Cmb 267073 D~ampraslan
<01 g Ditrbed rasd ak S5H07646 100 p 7.8 4 2
70 4 1 1 20463
334. SilenealaCarb 598.73
1 0069
<01 d 2311757 Hdazel
scrba t SK2626743 150 p 1240 1035 9 4
3351 SilenedigoicatCarm 075 1 02
336. SieenuasCarm 0.6 g 16.10.72 Deeitlmstn rsln SK 155732 40 p 7640

105 10 40 1
15 9 4 I 0.60
337 Socu avni Com s 03 d 16 873 Edeo l rfsi SK 420878 p
11 50 250 2 2 1 43
338. Spraim mru Spa 0.3 m 29 975 Rie SK 437768 <10 p
9.30 585 2 2 187
339 Sparganiueetum Spa 17 m I110 75 Margin fp~ond SK 554777 45 p
14
95 1 50 3 1 1
340 StahYs vlvatir Lab 28 w 19.973 Roadside SK 235760 65 p
00
0 75 050 3 1 1
341 Stellaria lsine Car 16 m 12.6 73 Masln agia osra SK 585626 15 p
04
40 p 1.45 1.25 1
342 Stellaiapalustris Car I 04 m 28 873 Damp hollo i wasteland SE608005
3 15
09 g 3 10.73 Deeitlmstnrsln SK 155732 55 p 450 160
343. Sucs rtni Dip 3.20 1 1 183
344. Tamscmuis i 08 w 5.1073 Hegeo SK 358741 300 p 320
Derlictral sidings SK 260655 65 p ( 0 75 I 1 01
345 Tanctumvlgare Com w 0.3 d 209.73
SK 331870 35 p I1870 1.25 9 4 3 0-64
346. Taaau ffcnl g Com u 120 d 235.72 Unutvtdadnpo I 1 08
3 10.72 nce SK 175665 25 p 150 1 00 I
347. Terium oodonta Lab 27 Deeitlmsoegrsln 1 1 03
24.975 lietn
Derelict grassland SK 154728 35 p 4 35 1.80 4
348. Thlctu miussp motnm Ran <0.1 4 1 1 03
I 01 107.75 Leadminepoil heap SK 287561 15 p 1.55 1.10
349. Thlaspilpestre Cru 4.69
350 Trgpgn rtni Com s 05 g 4.7.73 Road vege SK 236762 50 p 45 04 1.85 9 4 3
SK 498554 25 9.30 2 80 4 1 1 23
351. Trfliumdium Leg 07 g 15 10.75 Disjsd rhway ballast p
21 g 11773 Mowngraslandin park SE 310340 25 p 2.10 1.50 4 1 1 1.35
352 Trifliumpratens Leg 4 1 1 05
353. Leg 96 g 24.8 72 Copceietoesolha SK 164662 15 p 10 1OI00
Trflumrpn 1.95 1 25 2 1 1 10
354 Trliseuoau Ran <0 I 14.774 Stream bank SD 315860 35 p
4 1 02
SK341875 IS5 p 16 40 0.50 9
355. Tuslgoafr Com u 57 d 3 573 Brick andmortar rubble
200 8 10 0 30 5 4 1 00
356 Tvpha latifolia Typ 0 7 m 23.10 74 Marshland nea lake SK 436863 p
SK 174654 90 2.00 1 35 4 1 1 01
35 7 Uriadoc IJ,t
r 112 d 8 11 72 Stemie erc p
SK 172804 85 p 7 10 1 85 9 4 1 09
358 Vaein fiiai Val 0.8 g 29 773 Roadside bank
SK 261654 115 0.90 0.55 2 1 1 00
359 Vebsu thpu Scr <01 d 20.9.73 Disusd rail siig p
<0. d 8.10.73 Ditre atln SK 648985 70 p 1.00 0 70 2 1 1 01
360 Vebsu vrau Scr 1 03
atln SK 615992 45 p 1.25 0 70 5 1
361 Vebn Ver <01I d 9 10 73 Edeo yei
fiiai
19.9.73 Streamside SK 263805 15 p 1 75 1.20 4 1 1 03
362 Veoiabcaug Scr 08 m
0.50 4 1 1 00
Scr <0.1I m 26.7.74 Ditch SE 5951 95 25 p 0 75
363 Veroic catenata 0 95 4 1 1 01
Scr 2.4 17.8 73 Lietn uryha SK 144732 20 p 1 30
364 Veoiachmer, 4 1 1 03
Mieeiuoswoln SK 531831 20 p 165 1.60
365 Veoia otn Scr 05 w 16 8.73 01
15 1 25 1.05 4 1 1
Scr 0.4 3 18.73 Old heap~insnsoeqar SK 208868 p
366 Veoiaofiiai
SK 375748 90 e 2 80 2 60 1 1 142
367. Vii cac Leg 0.6 g 7.8.73 wasteland
Trampled
1I0 g 29 673 Lietn rslndiofcus SK 546828 5 e 400 1 35 2 1 1 28
368 Viola hirta V,o 1 1 04
SK 163729 10 e 2.00 1 35 2
369. Violklte Vio 0 2 g 4 7.73 Lead mine spoil heap
10 e 2 10 1.15 2 1 1 06
370 Vil auti Vio 0.3 m 15 974 Marshland SK 259876
10 e 1.85 1 45 2 1 1 10
3 71 Violark iin V,o 4.1 22.6.73 Birch scrub SK 208868

(e) Shrubs
Moorland SK 222903 30 p 0.68 0.46 2 1 1 0.03
372. Callun vulgaris Er 49 g 30 1074 1.27
Ran <0I1 11174 Deeitlmsoeqar SK 259778 1000 p 1920 1.50 9 4 3
373 Clemtisvitlba 2 1 1 21.97
374 Crteu ooin o 53 g 26.9.74 Diuealayebnmn SE 596004 600 p 11 35 8.90
8 7.73 Roadsidebank SK 172804 30 p 5.90 5.30 1 1 1 0.75
375 Empetrum nigrum Emp n 07 g 0.01
Eri n 04 m 13973 Oldpeatcuttingin
bog SE 720164 20 p 0.44 0 30 2 1 2
376 Erc etai 1 1 1 20.43
377 Heer helix Ara 60 w 265 74 w~all
Sandstone SK 329844 500 p 6 60 4.60
Deeitlmstn rsln SK 155732 5 p 1 75 10-0 2 1 1 0.32
378. Heinhmmcaacsu Cis 1.3 g 15 873
SE 599010 350 p 4.00 4.00 2 1 1 14-46
379 Liutu ugr Ole 01 8.1073 Hedgero 5.21
0.7 5.973 Fie addn SH 320298 300 p 9.80 9.80 1 1 1
380. Loieaprcveu Cap 1 22.93
Leg I <0 I g 8 10.73 Diue reus tip SE 602007 ISO 4.70 3 50 4 1
381. Lupinusaroru 1 1 118.13
05 w 5.9.73 Fixedsand-dune SH320298 250 p 14.95 13.45 2
382. Pruu pinosaRo 15-84
Ros <01 g 59.73 Fixedsand-dune SH 320298 25 p 13.00 12-50 2 1 1
383. Rosapipnlioi - - I I 1 2.49
384. R bsfuiou agg. Ros 11.5 .w 28.8.73 Wasteland SE 596008 ISO p
SK 32381 1 130 1225 11-80 1 1 1 1.15
385 Rubusidaeu Ros 1.2 20 773 Railw~ay
cutting p
8.09
0.2 w 23 7 73 Stabilizedlimestone sce SK 223743 25 p 8.50 7 50 1 1 1
386 Rubussxth Ros 1.90
1.3 3 1073 Unaae hdeo SK 231748 500 p 7.00 7 00 I I 1
387. Sabuusnigra Cap
Leg 0.3 g 7.8 74 Roadside SK 556584 130 e 3.70 2 25 4 6 i 8.55
388.Saohmu cpru 1.49
1
389.Soau uemr So~l 18 m 28 873 Hedgero SE 6000 10 115 p 10.00 1000O 1 1
1 1 6.97
<0I1 3 1 873 Deeitlmsoeqar SK 359238 ISO p 16.70 15-60 1
390 Svpoiapo iuai Cap 0.11
18 g 15 8.73 Deeitlmstn rsln SK 155732 5 p 0.50 0.50 4 1 1
39 1 Th,musdrucei Lab 6.20
04 g 14.8 72 Seidrlc atr SK 223745 130 e 3 25 2.30 4 6 1
392 Ulexeuopaeu Leg 0.26
44 g 3.773 Oak wodland SK 257799 30 p 6.00 5 30 1 1 1
393. Va,enim i'tillu Er
I I 0.37
Eri <0I1 m 19.9.73 SmallSphagnum
bog SK 263805 5 p 7.00 7 00 1
394. Vacinu oyoco 1 1 0.26
Er 0.9 g 10873 Blanket
bog SK 278836 20 p 7 70 6.90 1
395. Vacrnium vitis-idaeat

(f) Tree
Woodland onrvr erc SK 325815 1500 p 2.70 2.70 4 1 1 1.30
396 AIusglutinosa Bet 08 m 20 10.74
w Watln n cu SK 360774 1000 p 3.35 2.25 7 7 1 0.12
397 Beuapueen Bet 3.7 5 1073 17-05
Rha <0.1I 24.8.73 Mixeddeiuu wodland SK 562821 450 p 8 00 8.00 1 1 1
398 Fragula alnus
399. Frainusxcelior Ole 66 w~ 25.10.73 and
limestongrassland
Derelict SK 154732 ISOO
1750
p 28 15 7.90 7
2
1
1
1
1
46.19
744.70
400. Quecu ptrae Fag 1.8 ,w 24 10 74 Oak wodland IsLrub SK 325815 p 20.00 12.88
2.58
IS ,w IS 8.73 Stemieinmoln SK 289819 1750 p 11.25 7.80 1 1 1
401 Sobsauuai Ros
eiuoswoln SK 348623 1000 p 10-15 6.65 1 1 1 43.02
402. Ta usbacat Tax s 0.1 2.11.74 Mie
SK 530832 2000 p 20S55 15-95 7 7 1 9.93
403 Ulmsglabra Ulm 1.7 , 10.6 74 Mie eiuoswoln

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J. P. GRIME et al. 1057

Colou (9)

9 9 ~~~~
~~~5
IC ~ 20 IC ~ totmpertur ight (15)
Responseto
1-- ~~~3months
6months 12months >Imonth ( 3)
Ran,ge Light
l
Shade
t5
Dark
> % 5 5 t1 t1 % p(12) IC t 0Curvet14) p(12) % % % t5

I 5YR/3/1 54 31 45 42 60 53 35 41 57 42 germintiontt--t0
I 5YR/3/1 61 20 84* 13 78 13 82* 12 79* 14 - - - - d 63 22 57 25
I 2.5YR/3/4 0 5 27 75* 35 29* 24 18* 19 germintio-0 c 80 62 66 39 0*
I Black 0 0 0 0 0 - -- -
I 80 4 76 4 17* 4 6* 4 51* 4 c 11 34 1 11 d 98 2 96 2 16* 29
I IOR/3/4 90 3 93 3 98 2 100 2 97 2 d 6 24 3 II-TV d 99 3 100 3 91 39
I lOR/4/3 5 18 0 7 17 3 80* 4 d 16 36 2 1 d 48 7 67 5 58 5
I 2.5YR/3/4 15 5 95* 4 89* 3 99* 3 - - d 15 23 4 1 d 93 4 100 3 P*
I 2 5YR/4/4 99 8 100 5 00o 6 00o 3 00o 7 d germintiontt-0 d 100 4 98 3 12* 41
I 2.5YR/3/4 89 4 86 7 84 6 87 4 86 7 d 15 28 3 1 d 87 4 98* 4 0*
I 7.5R/2/2 2 16* 13 11 17 98* 6 100* 5 d -0
germintion d 100 5 100 4 0*
I 5YR/4/8 100 6 100 4 100 9 100 4 100 3 d 11 29 8 1 d 100 5 100 4 1
4 m 0 0 0 0 0- - - -
2 Black 0 0 I - - 0 c 18 36 3 1-111 c 79 5 99* 3 80 4
2 2.5YR/3/2 100 7 100 6 91 5 94 6 88* 5 d 7 28 5 II-TV d 98 6 100 5 76* 6
5 2.5Y/8/4 73 14 74 13 84 13 71 19 80 11 - - - -
I IOR/3/1 0 15* 29 14* 29 14* 24 - - d germintion0 c 74 20 57 22 2*
I 75YR/7/2 97 16 50* 16 77* 27 95 21 61* 23 d 31 36 7 1 d 55 12 14* 10 0*
2 5YR/3/1 99 4 100 3 100 4 100 4 99 3 d 10 33 2 11 d 99 4 95 4 0*
2 5YR/3/2 97 4 90 4 96 4 74* 5 86 3 d 16 29 3 11 d 72 6 80 5 34* 6
I 5YR/4/6 72 19 89* 11 100* 7 94* 16 100* 9 d 13 27 3 1 d 95 6 97 4 53* 51
I 7.5YR/5/6 8 10 56* 7 50* 6 76* 6 - - - - - -
I 2.5YR/3/4 92 18 86' 3 99 6 95 4 - - d 13 36 2 11 d 88 5 95 6 0*
3 lOYR/7/4 84 6 95 5 98* 4 93 5 88 5 d 12 34 2 II-TV d 99 4 96 4 2*
3 IOYR/7/1 94 4 95 3 96 3 78* 4 86 4 d 7 28 4 11 d 100 4 100 3 8*
3 33 49 26 21 37 38 44 19 71* 8 d 5 33 5 II-TV d 62 45 56 28 0*
3 8 21 0 0 0 - - c 6 24 3 11
2 5YR/7/1 20 6 7 7 8 6* 31 4 81* 3 d 12 35 2 III d 86 2 92 2 43* 3
2 7S5YR/4/0 80 6 94* 5 96* 3 99* 3 98* 3 d 7 31 2 II-TV d 97 3 97 3 86 41
2 5R/5/I 100 3 98 3 99 2 99 3 96 2 d 9 25 3 1 d 97 3 98 3 94 49
2 5YR/4/1 51 6 80* 6 74* 5 84* 4 88, 4 d 10 27 4 TV d 93 3 94 3 37* 3
3 5YR/2/1 0 0 0 0 - -
3 5YR/7/3 96 5 70* 6 95 5 100 6 - - d 8 37 2 II-TV
S 5YR/4/2 99 6 80* S 71* 7 36* 8 74* 6 - - - -
3 2.5YR/3/4 0 0 0 0 0 c 15 36 1 1 c 60 2 67 2 44 2
1 7 11 4 15 5 21 4 - - c 11 31 6 11
I 15 20 8 22 14 22 0 - -
I 5YR/2/1 0 3 I 0 0 - - - - c 72 11 76 9 0*
2 7.5R/4/4 90 6 100* 4 100* 3 95 3 100* 3 d 6 30 8 III-TV d 100 3 100 3 7*
2 lOR/4/I 0 4 31* 12 60* 7 90* 6 d <5 29 4 11 c 81 4 87 4 23* 4
S 7.5YR/5/2 70 13 71 12 56 12 7* 10 39* 14 d 12 31 14 TV d 62 14 55 18 48 21
1 0 0 0 - -
I 2.5YR/5/2 60 13 94* S 82* 10 - - 94* 3 - - - - d 94 3 100 3 6*
S IOYR/7/6 87 4 74* 4 63* 4 90 4 91 4 d 7 34 3 II-TV d 89 4 88 4 65* S
I IOYR/2/2 72 12 62 13 66 11 40* 11 60 12 - - - - d 37 12 41 11 17* 12
3 lOYR/3/I 5 23 9 10 9 17 12 15 5 19 - - - -
I 40 8 100* 2 100* 3 100* 3 100* 3 d <5 31 3 II-TV
S 5YR/4/2 90 15 93 10 48* 8 88 6 77 5 d 11 29 4 1-111 d 86 5 100 4 90 4
I 2.5Y/5/6 0 I 3 6 18* 5 1 <5 36 1 11
I vadm 11 1i 28* 7 26 5 46* 2 32* 2 5 35 1 11 s 98 1 100 1 100 it
I 7 6 3 22 14 2 21* 3 17 3 <5 39 1 11 100 2 100 2 100 21
I Black 0 0 0 0 0 - - - -
3 5YR/6/4 99 1 82* I 40* 4 0* 24* 4 f <5 34 <1 11 f 100 1 100 1 100 1
I 56 4 74 4 I00* 3 87* 4 86* 5 d 15 37 3 11 d 44 6 66 7 2*
I lOYR/4/4 60 7 39* 9 66 7 71 16 67 11 d 22 35 6 1 d 80 8 99* 6 0*
3 7.5YR/5/4 60 21 64 19 60 17 I00* 20 87* 17 d 19 27 10 I d 89 23 83 11 28* 9
3 7.5YR/3/2 26 7 I00* 4 61* 5 94* 4 94* 4 d 17 39 1 1-11 d 99 7 99 6 0*
3 7.5YR/3/2 I H* 6 8 4 I9* 4 21* 9 d 17 39 1 1 d 28 6 7* S 0*
2 5YR/3/2 0?-- - - - - 5 - - - -
I 5YR/4/2 100 14 100 11 100 10 100 6 98 9 d <5 24 5 11 d 100 9 94 8 94 24
I 7.5YR/5/6 86 6 I00* 3 - - 98* 4 I00* 3 d 7 34 2 11 d 99 4 97 4 0*
I 7.5YR/5/6 92 13 93 9 97 8 100 9 96 8 d 9 28 7 II-TV d 98 8 100 7 59* 119
I 5Y/7/8 0 0 0 0 0 - - - -
I 2.5Y/7/6 9 11 87* 9 69* 9 91* 9 - - d 10 31 6 11 d 88 10 I00* 10 lo* 10t
I 51 12 60 22 32* 11 34 10 20* 12 - - - -
- - 0 4 0 0 - -
5YR/5/6 0 28* 33 10 31 5 12 - - - - - -
I 5YR/2/2 20 11 4* 14 22 21 23 18 - - c 16 30 2 1 c 68 9 64 8 4*
I 7.5YR/5/4 0 0 0 0 - --

2 7S5YR/5/6 88 18 73 15 82 18 66* 20 92 15 d 10 28 14 TV d 70 15 62 15 0*
4 IOR/3/2 0 0 0 I - - - - - -
I 5YR/4/8 0 0 0 0 - - - - - -
I 0 0 0 0 0- - - -
2 5YR/4/6 8 27 35* 29 42* 35 4 26 60* 39 c II 31 5 11
2 7.5YR/S/4 90 12 70* 8 20* 9 - - - - f 6 29 6 11 f 76 10 84 7 84 8
I IOR/3/3 23 15 10 21 10 2 13 5 6* 8 <5 36 2 11 98 2 94 2 86 2
I 5R/2/2 0 I0* 27 20* 31 - - - -
I 7.5YR/S/6 0 0 0 0 - - - - - -
I 6 21 8 15 10 9 - - 4 - - -76 1 84 2 88 2
2 m 0 0 0 0- -- - - -
I 0 0 0 0 - - - - - -
2 0 0 0 0 - - - - - -
2 lOYR/8/2 0 0 0 0 0- - - -
2 7S5YR/7/4 0 0 0 0 - -
2 7S5YR/S/6 2 0 0 - - - -
I 73 10 85 9 90* 11 47* 31 2* - - - - 98 5 100 6 98 5
2 IOYR/6/4 2 13* 31 14* 22 I8* 21 - - c 19 35 2 1 c 76 6 I00* 4 70 3
1 0 - - 0 0- -- - - -
I 92 5 93 4 90 3 96 3 99 3 d 9 28 5 II d 96 4 98 4 75* 49
I 32 14 42 9 41 8 50 9 12* 10 - - - - 98 6 100 6 100 6
I lOR/4/6 95 20 78* 14 90 15 97 15 - - f 12 27 11 III-TV f 93 14 100 13 95 131
I 2.5YR/3/4 0 72* 15 88* 12 78* 10 - - f germintion0 f 64 10 80 11 23* 12
I 5YR/4/4 60 21 92* 19 97* 19 91* 19 - - d 18 28 11 11 d 98 20 87* 17 4*

I 5YR/4/4 18 4 12 4 5* 3 28 3 29 4 - - - -
I 7.5YR/6/8 6 4 7 4 22* 3 17 4 II S5
I - - - 0 0 0 - - - -
I 7S5YR/S/6 2 0 0 - - - -
I 5YR/4/3 95 11 0* 0* 0O -
I 7.5R/3/4 0 0 0 0 - -
I 5YR/6/3 0 0 0 0 - -
I m 82 10 89 8 87 7 46* 8 - - - - - - d 88 6 96 6 4*

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1058 Germination
characteristics
in a localflora
APPENDIX 3
Synopsisoftheclassificationsusedto analysetheexperimental data. Classifications
(a)-(g) referto characteristics
of the species,(h)-(m) referto the seed. The four
columnsrelateto different typesofgermination result.The valuestabulatedare the
numberof speciesforwhichresultswereobtained.For keyto parenthetic notes,see
Appendix2, exceptthat(*) = does notincludesixteenspeciesin whichmostofthe
seedsfailedto germinateat constanttemperature(see Appendix2)
Initialtestson
freshly-collectedMeasurements Measurements Measurements
and dry-stored ofgermination ofresponseto ofresponseto
seeds rate temperature light

Total speciestested 403 267 256 (*) 271

(a) Life-form
Annualgrasses 9 9 8 8
Annualforbs 82 49 55 55
Perennialgrasses 45 43 35 38
Perennialforbs 235 155 150 158
Shrubsand trees 32 11 8 12

(b) Family
Compositae 52 48 46 44
Cyperaceae 18 10 12 12
Gramineae 54 52 43 46
Leguminosae 19 7 13 16
Umbelliferae 18 2 6 4

(c) Britishdistribution
(1)
Northern 43 27 31 25
Southern 144 93 100 97
Ubiquitous 137 96 89 104
Widespread 60 35 25 29
Local 19 16 11 12

(d) Frequencyin Sheffield


flora(2)
>10 15 14 14 14
1.1-10.0% 135 92 91 101
0- 1-1-0% 165 110 109 116
<0- 1% 85 48 42 40

(e) Habitat(3)
Disturbed 90 59 59 61
Skeletal 61 51 48 46
Marshland 84 57 55 60
Grassland 109 77 78 80
Woodland 59 23 16 24

(f) Heightofinflorescence
(cm) (4)
0-20 100 64 68 65
21-40 115 80 73 80
41-60 73 56 53 52
61-80 45 27 30 35
>80 70 40 32 39

(g) Dehiscence
Passive 374 253 244 254
Explosive 29 14 12 17

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J. P. GRIME et al. 1059
APPENDIX 3-continued
Initialtestson
freshly-collectedMeasurements Measurements Measurements
and dry-stored ofgermination ofresponseto ofresponseto
seeds rate temperature light
(h) Dispersuleshape
Spherical 51 27 27 31
Ovoid,rhomboidalor turbinate 88 51 45 53
Trigonousor triquetrous 36 21 21 23
Lenticular,
reniform or subulate 103 63 64 71
Cylindricalor ligulate 39
Clavate 12
Winged 10 59 54 51
Tadpole-shaped 15
Conical 49 46 45 42
(i) Dispersuleappendages
Absent 291 180 170 189
Straightawn(s) or spine(s)only 25 18 19 19
Hygroscopicawn or spine 13 12 11 9
Pappus or persistentcalyx 51 47 47 43
(j) Dispersulehairsor teeth
Absentor veryinconspicuous 319 192 185 206
Radial or irregular 15 10 14 11
Antrorse 69 65 57 54
(k) Germinuleweight(mg)
<0- 1 66 61 58 57
0 1-0 99 184 133 134 138
1-00-9 99 126 62 59 68
>9-99 25 9 5 7

(1) Germinulesurfacetexture
Smooth 210 131 122 141
muricateor
Rugose,tuberculate,
reticulate 88 61 58 57
Striate 46 28 31 29
Hairy 21 13 15 12
Striateand hairy 23 20 20 18
Mucilaginous 15 14 10 14
(m) Germinulecolour(Munsellhue)
5R, 7-5Rand IOR 67 44 39 47
2-5YR 41 33 29 30
5YR 105 69 68 68
7*5YR 59 45 39 40
1OYR,2-5Y and 5Y 57 38 38 38
Black 11 4 6 7
Multicoloured 21 15
Variable 32 J34 37 20
and variable
Multicoloured 7 6

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