Chapter (non-refereed)

Park, D. G.. 1982 Seedling demography in quarry habitats.

In: Davis, B. N. K., (ed.) Ecology of quarries: the
importance of natural vegetation. Cambridge, NERC/ITE,
32-40. (ITE Symposium, 11).

Copyright © 1982 NERC

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 32

SEEDLING DEMOGRAPHY IN QUARRY HABITATS

DAVID G PARK
Institute
of TerrestriaZ
Ecology,
MonksWoodExperimental
Station,
Huntingdon
INTRODUCTION

Seedand seedlingecologyhavebeen largelyoverlooked in the studyof

quarries,but are areasof importance when considering
thenaturalcolonisation
of thesehabitats.Quarrying initiates a primarysuccession
with the removal
of existingvegetation, themineralised soillayerand the seedbank.
Colonisationby plantsof thesedenudedareaswillbe dependent on the influx
of seed/propagulesfromexternal sourcesand the actualrateof colonisation
willbe dependent upon the outcomeof seedlingestablishmentand subsequent
seedlingsurvival.
A studyhas beenmade of bothnaturaland experimental seedling
populationsof severalcommonplantcolonists.Particular attention
has been
givento ratesof recruitment andmortality,and to ways in whichtheserates
may be modifiedby experimentaltreatments.Resultsfrom thisworkwillbe
reportedin the paper.
GENERAL ASPECTSOF SEEDLINGECOLOGY

Althoughthe naturalcolonisation of disusedquarries has produceda

richdiverseflora,-alongdevelopmental periodis oftenrequired beforean
appreciablecoverof vegetationis established. The communityremainsopen
withmuchbareground. In sucha situation, the opportunityfor recruitment
to seedlingpopulationis likelyto be greaterthanthatin a closedcommunity
whererecruitmentdependson the occurrence of gapsin thevegetation cover.
However,seedlingmortalitymust be severeotherwise thisopencommunity
structurewouldnot be a persistentfeatureof the quarryfloorhabitat.
Ratesof mortalityamongjuvenileplantsare veryhigh:theyoung
seedlingis themost susceptible
phasein theontogenyof the individualand
mortalitygenerally
decreaseswithage. Thiscan be demonstratedin the
followingsimpleway.
1. If an individual'sontogenyis dividedintoa numberof discrete intervals
by separatingeitherdevelopmental stagesor age states(Figure 4A),it
is possibleto assignprobability values(Pi)to the chancesof passing
fromone stage/state to the next.
2. Multiplicationof the interval probabilities(Pi)givesthe overall
probabilitythatan individual reachesa certainstageof development
or age (Figure4B).
3. Probabilityhistograms for the Pi'sfor a generalised perennial and an
annualshowtwo alternatives (Figure 4C). In the first,a postulated
increasein interval probability valueswith age/development gradually
levelsoff. In the second,thepostulated increase continues untila
pointis reached,perhapscoincident with flowering, whenprobability
valuesdeclinesharply.
4. Considerthe shadedareasof thehistograms coveringseedling establishment
up to a limitof 60 days;the calculated probability of an individual
passingfromemergedseedlingto a sixty—day old seedling is of the order
0.18-0.75,depending on species, for seedlingson quarryfloors
(Figure4D). Similarwork in Americagivesa slightlylowerrangeof
0.08-0.55(Skaller 1977).
 33

B. pr(Emerged --o-S4x
Seedling 3) = P1 x P2 x P3

C. Probability
Histograms
-
perennial annual

60
1 4daylw 1-

0I 0-

D. Quarryfloorspecies:pr(Seedling---4.S+60
days)= 0.18-0.75

E.
Seed t Emergedseedling
Po

4 60 days)= 0.02-0.10
Quarryfloorspecies:pr(seed---4.seedling

Figure4 Survival
probabilities
in seedZings

5. However,if the intervalfrom seed to emergedseedlingis also included,

then the probability
thatan individual passesfrom seed to sixty-day
old seedlingfallsabruptlyto 0.02-0.10(Figure4E).
From thisit wouldappearthat themajor limitingperiodof natural
colonisationis the passagefrom seed to seedling.For the particular
quarry
studiedby Skaller,it was concludedthat seedinputwas not a limiting
factorin the colonisationprocessbut ratherthe eventsfollowingthe
arrivalof seedson the spoilsurface.
 34

COHORT VARIABILITY

The approachoutlinedabove,whileshowingthemagnitude of seed/seedling

loss,is an over-simplification and doesnot showone important aspectof
seedlingecology- thatof cohortvariability. Baskins& Baskins(1972)found
thatearlygerminating cohortsof Leavenworthia stylosasufferedhighmortality
but had a greaterprobability of flowering, whereaslatergerminatingcohorts
avoidedsuchsevereearly ,mortality but had a lowerprobabilityof flowering.
Thisgeneralrelationship can be foundin quarryseedling but
populations
an overallinfluence is exertedby theprevailingclimaticconditions.
Seedlingemergence and mortality froma permanent on a
quadratestablished
quarryfloor,was studiedintensively for twoyearsand seedling survivorship
curveswere constructed for different cohorts(Figure5). During1979,
germination was curtailedby thedry conditions occurringduringJune/July
and the relationshipbetweenearlyand latecohortsis not shown. However,
during1980thisrelationship can be clearlyseen.
One of themost noticeable aspectsof seedlings emergingin quarry
habitatsis the longperiodof timetheyspendat a smallsizeand at an
eirlystageof development (at the cotyledon,firstand secondleafstages).
Researchelsewherehas shownthatannualgrowthratesare smalland it is
commonto findannualsand perennials remainingas basalrosettes for extended
periodsof time (Raynal1979,Klemow& Raynal1981). Perhapsthe singlemost
importantcharacteristicof juvenile plantsthatinfluences theirfitnessis
theirsizeduringenvironmental stress.Mortality is stronglydependenton
sizewith smallerindividuals suffering greaterhazards.For manyplantsthe
totallengthof the juvenile periodwillbe a functionof size- itselfa
functionof growthrateswhichreflect, amongstotherthings,nutrient
availability.
CAUSES OF SEED AND SEEDLINGLOSSES

1. The most important causesof seedlossesduringthe earlystagesof

colonisation are probablyphysiographic processes suchas wind andwater
erosionand frostheave,leadingto seedburial. In an experiment on the
recoveryof dyed seedssownon to a barequarryfloor,lossesamountedto
20-50%after25 days,30-75%after50 daysand 60-90%after125days (which
includedan overwintering period).Seedswere foundto havemoveddown the
spoilprofileduringthe courseof theexperiment.
2. Probablythemost important factorcausingseedling mortality is
desiccation actingbothdirectly, duringperiodsof drought, and indirectly,
for exampleafterspoilmovementand exposure or roots. The spoilis freely
drainedand the surfacedriesrelatively quickly: duringlatespringand
summerthemoisturecontentof the spoilis frequently belowthepermanent
wiltingpoint(Figure5).
3. Failureof the radicleto penetrate the spoil,as a resultof the
formation of a cementation layeror becauseof waterlogging of the spoil.
4. Predation or grazingby invertebrates can causehighmortality in
seedlingpopulations and may be selective in nature.
5. High surfacetemperatures may causedeathdirectlyand are certainly
capableof stressthrougheffectson evapotranspiration.
6. In activequarries, theadverseeffectsof dustcan be important: dust
on leavesreducesphotosynthesis, interferes with transpirationand increases
the riskof disease. The overriding influence of nutrientdeficiency may
increasethe effectof thesemortality agents,for exampleplantsbecome
more susceptible to waterstress.
 35

1979
SPOIL MOISTURE

02 0

wilting point
0
A A S 0

100 1979
SEEDLING SURVIVAL
50

•—

100 • • ......

50

1980
V) SEEDLING SURVIVAL
cl<5

1980
SPOIL MOISTURE

220

.?10
.6
wilting point
0
A A

andseedling
Figure5 SpoiZmoisture survivorship for 1979and
curves
study.Survivorship
quadrat
1980;datafroma permanent
curves emerging
arefortotaZseedZings
 36

EXPERIMENTS ON ORIGANUMVULGARE

Someof thesepointsmay be illustrated to a particular

by reference
species,studiedin a quarryhabitat.Origanum magare(marjoram) is a common
constituentof quarryfloras:it is an aromatic herbwith a bushy
perennial
appearance.The seedsare small(6.4x 10-5g)and produceseedlings of only
a few millimetresin size. DuringFebruary1980,a numberof areasof quarry
floorwere artificiallyseeded,and seedlingemergence,survivaland
developmentweremonitoredintensively throughouttheyear. The quarryfloor
consistedof a shallowlayer(0-10cm) or spoiloverlying
of crushedlimestone
thebaserock: the spoilcontainedlittleor no organicmaterial,was freely
drainedand poor in bothmacro-andmicro-nutrients.
Germinationin the plotsbeganin Apriland was more or lesscomplete
by July;totalplotgermination was in the range18-51%and comparedwith a
laboratorygerminationvalueof over90%. The population fluxof one of the
seededareasis illustrated (Figure6A) and showscumulativegermination,
mortality
cumulative and thenumberof seedlings present.The seedling
populationpeaksin earlyMay and thendeclinesto a levelwhichremains
relativelystablethroughout July-October. Cohortsurvivorshipcurves
(Figure6B) clearlydemonstratethe relationshipbetweenearlyand late
cohortsremarkedupon earlier.However, duringthe followingwinter,the
was greatlyreducedby the effectsof frostheave,with only
population
1-3%of the seedlingssurvivingto March1981.
Duringthemonitoring of theplots,the seedlings presentwere classified
accordingto six stagesof development(Figure7). The histograms in this
figuregive the proportionalrepresentationof theseclassesat thedates
indicated.Histogram 8 (November) showsthatevenaftera considerable
periodof time (around180 days)most of the seedlingsare stillat an early
stageof development.Age-class frequencydistributions are
of the seedlings
also shownfor selecteddates(Figure8). The developmental stagesdo not
reflectage classes.The sizeof an individual, whichoftenreflectsits
stageof development, in determining
is more important itsbehaviour than
its chronologicalage (Gross1981, Werner1975).
0. vulgarewas alsousedin experiments designedto increase
recruitment and reducemortality.A sawdust mulchwas appliedto sownplots
of O. vulgare. Recruitment in plotsmulchedwith sawdustwas verymuch
higherthanin non-mulched plotsandmuch of the earlyseedling mortality
was avoided(Figure9). Themain effectof themulchwas to increasethe
spoilmoisture2-4%. Althoughrecruitment and seedlingsurvival were found
to be greaterin mulchedplots,the rateof seedling development was
unaffected and was similarto thatof non-mulchedplots.
CONCLUSIONS

Seedlingecologyformsan important partof the studyof a primary

successionsuchas thatoccurring in disusedquarries.The seedling
representsone of themost susceptible stagesduringthe
and vulnerable
ontogenyof a plant;rates of and
recruitment mortalityin seedling
populations therateat whichcolonisation
may inflUence proceeds.
ACKNOWLEDGEMENTS

Thisworkwas fundedby a NaturalEnvironment ResearchCouncil

Instituteaward. Dr B N K Davisand Mr R N Humphriesgavevaluablehelp
programme
in the planningof the research of the
and in thepreparation
manuscript.
 37
A
500

400

100

50

li
A M J i A
1980
Figure 6 Origanumvulgare:seedlingemergence
and mortality.A: popula-
tion flux diagram o--o cumulative
recruitment, v--vcumulative
mortality, e•—• seedlingspresent. B: cohortsurvivorship
curve.
 38

400
Seed•
0.5 6r 3c m

0.

IV .V1

-
-0
a)

'46

a)
_o

0
IL
A M J J A S 0
1980
stageanaZysis
Figure7 Origanumvulgare:Developmental of
(asa proportion
dates.
seedssown)at seZected

0.51 A

0.51 C

01
0 28 56 84 112 140 161
Age class (days)

age-class
Figure8 Origanumvulgare:SeedZing (asa proportion
distribution
of seedZings
present)at threedates.
 39
E 40 A.

100

B.

20

100

AA i i A
19801 -

Figure 9 Origanum Population

vulgare: andcohort
fluxdiagrams
results
curves(mean
survivorship fromthreerepZicates)
A: non-mulched
plots pZots
B: muZched
 40

REFERENCES

BASKINS,J.M. & BASKINS,C.C. 1972. Influence

of germination
dateon
survivaland seedproduction
in a naturalpopulation
of Laevenworthia
stylosa.Am.Midl.Nat,88, 318-333.
GROSS, K.L. 1981. Predictions=c;f
fatefromrosettesizein four 'biennial'
plantspecies:Verbascum thapsus, Oenotherabiennis, Daucuscarota
and Tragopogondubius.Oecologia, 48, 209-213.
KLEMOV, K.M. & RAYNAL, D.J. 1981. Population ecologyof MelilotusaZba
in a limestone quarry. J. Ecol.,69, 33-44.
RAYNAL, D.J. 1979. Population ecologyoE Hieracium floretinum (Compositae)
in a centralNew York limestone quarry.J. appl.Ecol,16, 287-298.
SKALLER,P. 1977. PZantcolonisation andsoildevelopment in TheJamesville
Quarry.PhD thesis,StateUniversity, Syracuse,New York.
WERNER, P.A. 1975. Predictions of fatefromrosettesizein Teasel
(Dipsacus fullonum L.). Oecologia, 20, 197-201.