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Soil Development in Relation to Vegetation and Surface Age at Glacier Bay, Alaska

Author(s): Robert L. Crocker and Jack Major


Source: Journal of Ecology, Vol. 43, No. 2 (Jul., 1955), pp. 427-448
Published by: British Ecological Society
Stable URL: http://www.jstor.org/stable/2257005 .
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[ 427 ]
SOIL DEVELOPMENT IN RELATION TO VEGETATION
SURFACE AGE AT GLACIER BAY, ALASKA*

AND

BY ROBERT L. CROCKER AND JACK MAJOR


University
of California
(WithPlates 4 and 5 and seventeen
Figuresin theText)
INTRODUCTION

Very few reliablefunctionsof the rate of developmentot soil propertiesare available.


The paucityofquantitativedata is due primarilyto thedifficulty
withany
ofestablishing,
of
as
the
degree precision,
great it is, is
age of particularland surfaces.This difficulty,
accompaniedby othersscarcelyless important,includingthe magnitude,sequence,and
ofvariationsin climate,parentmaterial,or biota whichmighthave occurred
significance
since the initiationof soil formation.The Glacier Bay regionin South-easternAlaska
althoughhavingits own peculiarlimitationspresentsthe opportunityof establishingthe
rate ofchange,over a shortperiod,of manysoil properties,withmostof the above difficultiesreducedto the minimum.This especiallyapplies to those propertieswhichrelate
directlyto vegetation. At GlacierBay a fairlydetailedknowledgeofthehistoryofrecent
deglaciationhas been established(Cooper, 1937; Field, 1947), and along with Cooper's
(1923, 1931, 1939) classic vegetationsuccessionstudiesit affordsan exceptionallygood
basis for this kind of investigation.In the presentpaper the rate of developmentof
some of the characteristics
of the forestfloor,and changesin the mineral-soilproperties
of reaction(pH), organiccarbon,total nitrogen,calcium carbonate,and bulk densityof
thefineearthare reported.These changesare relatedbothto age ofthesurfaceand to the
developingvegetation.
For purposes of general orientation,and because regionalclimate is an important
ecologicalfactor,available data forthe nearestthreemeteorologicalstationsto Glacier
Bay, namelyGustavus,Cape Spencerand Haines, are brieflysummarizedbelow (Fig. 1
and Table 1). Near GlacierBay the Alaskan coast is dissectedby severalfiordstrending
north-south.Cape Spencer,on the outercoast, fullyexposed to the PacificOcean, has
a verywetmaritimeclimate. GlacierBay lies in thenextfiordinland.The neareststation
to Glacier Bay is that at the Civil AeronauticAuthorityairport,at Gustavus. It is
located near the mouthof the fiordon Icy Straitand probablygivesa reliablesummary
of the climateof the lowerbay. Here it is drierand less maritimethan at Cape Spencer.
Inside the innermostfiord,leadingto Skagway,the Haines CAA stationis decidedlythe
most continentalof the three. The calculated 'potential evapo-transpiration'(after
Thornthwaite,
1948) at all stationsis verysimilar.The regionalclimateof GlacierBay is
probablysomewhereintermediatebetweenthat at Haines and Gustavus CAA, and in
termsof Thornthwaite'sclassificationof 1948, all the climateswould be perhumidand
mildmicrothermal.
* This paper resultedfroman invitationfromProfessorD. B. Lawrence,Botany Department,Universityof
Minnesotaforthe seniorauthor to participatein his expeditionto Glacier Bay in the summerof 1952. The
studieswereaided by a contractbetweenthe Officeof Naval Research,U.S. Departmentof the Navy and the
Universityof Minnesota(NR 160-183).

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Soil development
at GlacierBay

428
S~~50

?-

&
Gustavus
CAA

Cope Spencer

Precipitation

* Potential
evopotomnspirotion
/, MoWisture
deficiency

\
/ ;

Homes CAA

,,

o
C-

41+

J FMAMJ

J ASONDJ

FMAMJ

JASOND

J FMAMJ

J ASONDJ

Month
Fig. 1. A summaryof some meteorologicaldata forthe threestations
nearestGlacier Bay.

Table 1. Climaticdatafor threestationsin theneighbourhood


of Glacier Bay, Alaska

Av. annual

Period of
record(years)Temp.
Station
(Temp.) (Ppt.) (? C.)
16
16
5-8
Cape Spencer
Gustavus CAA
16
16
5-1
Haines CAA
26
28
4-7

Ppt.
(mm.)
2924
1419
1531

(a) Generaldata
Water
Water
Potential
Relative
deficit
evaposurplus
continent- (dry season) (wet season) Moisturetranspiration
factor
(cm.)
(cm.)
(cm.)
ality
448
0
6-7
239-1
53-3
0
177
51-2
90-6
13-8
51-1
205
20-2
7-0
109.0

(b) Mean monthlytemperature(? C.)


Jan. Feb. Mar. Apr. May June July Aug.
4-6
11-3
11-4
0-4
0-1 2-4
7*3
9-5
Cape Spencer
12-8
12-2
Gustavus CAA - 2-4 - 2-1 0-9
4-1
8-0
11-2
14-1
Haines CAA
- 4-7 -3-6 -0-1
4-1
12-9
941
13-3
Station

BROAD

FEATURES

OF

THE

ICE

RECESSION

AND

VEGETATION

Sept.
10-2
99
10 1

Oct. Nov.
7-1
3-4
6-0
1-3
5-3 - 0-3

Dec.
1.1
- 15
- 3-8

SUCCESSION

The recessionwhichfollowedthe post-Pleistoceneadvance ('Little Ice Age') of Alaskan

glaciers has been remarkably rapid in the Glacier Bay region. The rate and extent of

deglaciationis summarizedin Fig. 2. The ice has recededas much as 60 miles(96 km.),
and the old foresttrim-lineindicates that the vanished ice was as much as 2600 ft.
(792 m.) thickin places. A verydetailed account of recessionin the fiordtributaryarm
knownas MuirInlet is available (Field, 1947). Here the recordis good sincethe 1890's,
whenthe naturalist-explorer
JohnMuirbuilt a cabin near the ice front.It is especially
about
The
broad
1913.
good since
pictureof the rate and extentof glacial recessionat
GlacierBay as a whole has been providedby Cooper (1937) in an analysisof historical
data in relationto his own ecologicaland paleo-ecologicalresearches.He concludedthat
glacial retreat commenced approximately 150-200 years ago. More recent investigators
have confird
this conclusion but have tended to favour the older date (Field, 1947;
fit
in Lawrence & Elson, 1953).
1953,
Lawrence,
The general features of plant succession following deglaciation are well understood.
Cooper's studies commenced in 1916, and extended over three other expeditions (1921,

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ROBERT

429

L. CROCKER AND JACK MAJOR

1929 and 1935). Lawrence,his colleagueat the Universityof Minnesota,has continued


the workon fourmoreexpeditionsin 1941, 1949, 1950 and 1952. Broadly,threemajor
vegetationtypes can be recognized:the pioneercommunity,the willow-alderthicket,
and the spruce forest.These types representa physiognomicdevelopmentalsequence,
136?30'

137?00'

[..~,

... 9

/907

./7*..*

/949/8

ALASKA

135030'

13600'

4 p CANADA

.
ig4

GOOSE COVE

~/
?

Cooper(1931):~~TETOVE
~~~~~~~~~from
morerecent work).

The process of development which is responsiblefor the present aspect is brieflyas follows. The first
arrivals are the mosses Rhacomitriumcanescens Brid., and R. lanuginosum (Hedw.) Brid., Epilobium

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430

Soil developmentat Glacier Bay

latifoliumL. (broad-leaved willow herb), EquisetumvariegatumSchleich. (variegated horsetail) and Dryas


DrummondiiRich. The last is by farthe most importantbecause of its mat-forminghabit. To some extent
with these, and finallysurpassingin importanceall but Dryas, appear 3 species of prostratewillows,Salix
arctica Pall. being most important. Next comes a group of shrubby willows, Salix Barclayi And.,
S. sitchensisSans. and S. alexensis (And.) Coville. These usually begin lifein depressed or prostrateform,
later developingan erecthabit, thus constitutingthe transitionfrompioneerto thicketstage. S. sitchensis
and S. alexensis maintain themselves as important members of the latter community. Alder (Alnus
tenuifoliaNutt.*) gradually asserts itselfand becomes the most characteristicthicketdominant. Finally
Sitka spruce (Picea sitchensisCarr.) supersedes the shrubs, at firstforminga pure stand, and to this the
two hemlocks, Tsuga heterophylla
Sarg. and T. mertensianaCarr. gradually add themselves.

In the initialstages the vegetationis verysparse,but slowlyit beginsto fillin. The


significant
pioneerplants,as Coopernoted,all have verymobiledisseminules.The early
appearance of the colonizingvegetation,as shown in P1. 4, phot. 1, is due to the variabilityof the effectivedissemmulefactorand the variabilityof the soil parentmaterial
whenlaid downas freshmorainicdebris. Almostany ofthe major speciesofthe developmentalsequenceare physiologically
able to becomeestablishedearlyon thebare surfaces,
but neithersuch a surfacenor the distributionof disseminulesis uniform.The resulting
vegetationis consequentlyheterogeneous.
In the MuirInlet region,small thicketsof willowor alder,and isolated cottonwoods
areas largelyoccupiedby the Dryas
T. & G.), set amid intervening
(Populus trichocarpa
15-20
mat givea variable,but pleasing,appearanceafter
years. Bare groundis stillvery
frequentat this stage-indeed the coveris stillfarfromcomplete. It is not until35-40
yearshave passed that the alder coveris almostcontinuousand the vegetationtakes on
an appearanceof homogeneity.Even thenthe ascendingscatteredcottonwoodsovertop
the alder, and beforelong the spruce,heraldingthe suppressionand eliminationof the
shrubthickets,beginsto followsuit. The classificationof the changesmakingup this
sequence into stages is largelya conveniencewhichshould not mask the continuityand
of the developmentalprocesses. The stages are not entirelyarbitrary,
interdigitation
however,forthe pointsmarkingthe eliminationofthe prostratepioneersand herbswith
expansionof the alder is quite definiteat any particularsite. The point of elimination
of the tall shrubs,such as alder, in the developingspruceforestis equally specificand
final.
THE

SOIL PROPERTIES

AS FUNCTIONS

OF AGE OF LAND SURFACE

Field observations
In the earlieststages of deglaciationthe soils are merelydisorganizedaccumulationsof
morainicdebriswhosesignificant
propertiesshow no regularvariationwithdepth.With
the depositionofthestrandedglacialtillin a stableposition,climateand vegetationbegin
to modifythe soils,and to impartto themespecialcharacteristics.In thiswet maritime
climateleachingof solubleconstituentsand exchangeablemetalcationssoon modifythe
surfacehorizons.The processesappear to be greatlyacceleratedwiththe colonizationof
the newsurfacesby plants,and especiallyaftersurfacelitteraccumulationshave covered
the mineralsoil. It was noteworthy
in the fieldstudies,forexample,that the degreeof
*

Accordingto Hulten (1944) the commonalder in the Glacier Bay area is Alnus crispa (Ait.) Pursh,subspecies sinuata (Regel). We shall adopt the Hulten nomenclaturehere. WhereverA. crispa is mentionedit
will referto this subspecies.

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Vol. 43, Plate 4

JournalofEcology

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Alnus and
in
Goose Cove
The early
of plant
Phot. 2.
1. The
colonization, Goose
Phot.
stagesof
plant colonization,
region, Muir
Dryas in
Cove region,
MuirInlet.
Inlet. Alnts
and Dryas
early stages
foreground;Dryas, Populus, Salix and occasional Alnus in centre; Nunatak Nob and remnant of
Hulbert.
C. Hulbert.
L. by
in distance.
distance. Photograph
McBride Glacier
Glacier (right)
McBride
Photograph
by L. C.
(right) in

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Vol.43, Plate 5

JournalofEcology

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themossyfloor.Photograph
through
disappearanceofalder-onlythebuttsofstemsnowprotrude

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ROBERT

L. CROCKER AND JACK MAJOR

431

loweringof mineral-soilpH, forsurfacesof similarage and parentmaterial,was largely


dependentupon the type and age of vegetation.
Permanentlyfrozengrounddoes not occur in this part of Alaska, but the freezing,
effectsupon the earlybare, and partiallybare, surfaces,tend to
thawing,wetting-drying
a
produce stonypavementbeforethe shrubvegetationis establishedand prominent.This
stonypavementis moreor less continuousexcept wherepatches of finertill,like those
sampled in our investigation,occur. With the developmentof the alder thicketstage
organicresiduesbuild up and restupon the pavement.The littercoverappears to retard,
and soonto inhibitfurther
pavementdevelopment,but the pavementalreadyformedhas
ofthemineralsoiland organicresiduesintermixing,
a markedeffect
uponthepotentialities
and must greatlyinfluencethe rate and patternof translocationof materialsfromthe
forestfloorto the mineralsoil.
The establishmentof vegetationon the bare deglaciated areas initiatesgradientsin
manysoilproperties.Those mostobviouslyaffectedand observablein thefieldare organic
continuesuntil,in the oldest
matter,reaction,bulk density,colour.This differentiation
membersofthesequence,some5-8 in. (13-20 cm.)oforganicresidueshave accumulatedas
the forestfloor,and the colourof the mineralsoil itself,and its bulk density,show conand variationwithdepth.The early variabilityofthe mineral-soil
siderablemodification
to
the
related
irregularpatternof pioneervegetation,seems to have largely
properties,
foresthas developed. However,a statistical
the
time
the
spruce-hemlock
disappearedby
of the older soils to be more apparent
shown
the
have
greateruniformity
study might
than real, forthe forestfloorverydefinitelycontinuedto show large areal variations.
ofthefineearthattributableto the pedogenicprocesseswas
No texturaldifferentiation
discerniblein the fieldin any of the soils of the sequence. This is not unexpected,for
studies elsewheresuggestthat, at least forthose few reliable chronosequencesalready
studied the formationor illuviationof significantquantities of clay requires a much
longerperiodof timethan has been operativesince the beginningof the currentGlacier
Bay deglaciation.
A fieldobservationof pedological interestwas the fact that in relativelyrecently
deglaciated regionsthe surfacesdevoid of vegetation,or only sparsely covered, had
developed a markedvesicularstructurein the upper I- in. (6-12 mm.). This vesicular
layeralthoughthinappeared as quite definitewithin5-10 yearsof disappearanceof the
ice, and forparentmaterialsof equivalenttexturecontinuedto develop and increasein
thicknessuntil vegetationcover became fairlycomplete. By the time the alder-willow
forestfloorresiduesbegan to accumulatethelayerseemsto have completelydisappeared.
vesicular
The vesicularstructurewas identicalin appearance withthe little-understood
those
of
the
Intersoils-more
cold
of
desert
so
characteristic
especially
many
layer
was
vesicular
structure
the
in
mountainRegion the U.S.A. Although
veryfrequenton
not
form
on verysandy
thebare deglaciatedsurfacesit was obviousin thefieldthatit did
whether
parentmaterials. A seriesofsamplesweretakenwiththe idea ofdemonstrating
a changein bulk densityof the fineearth (weightof fineearth per unit volume of soil
in situ) accompaniedthe developmentof the vesicularlayer, and they indicated, surthatat least withthe degreeofaccuracypossible underthe methodofsampling,
prisingly,
fromthat ofthe finetil
the bulk densityofthe vesicularlayerdid not differ
significantly
of the initial surface. It seems very probable in the Glacier Bay region that the reorientationand groupingof discreteparticlesto produce the vesicularlayeris related in

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432

Soil developmentat Glacier Bay

some way,not onlyto texturebut to the alternationoffreezingand thawing.This might


well be a mechanismelsewherefor the layer is particularlyprominentin cold desert
regionsand apparentlymuchless so, or absent,in hot deserts.
The fieldinvestigations,duringwhich the foregoingobservationswere made, were
primarilydesignedto be thebasis forthecollectionofsamplesforlaboratoryexamination
which would characterize,in more definiteand quantitative terms the qualitative
ofthefield.The laboratoryanalyseswerealso
observationsor less accuratemeasurements
necessaryfor investigatingpropertieslike total nitrogennot measurablein the field.
time states of the deTherefore,a numberof soil samples representativeof differing
velopingecosystemweretaken. To avoid complexitiesintroducedby the colonizationlag
arm of the fiord,the samples were collectedmostlyalong the eastern
in the north-west
side of GlacierBay. The inherentcoarsenessand stoninessof glacial till makes it a very
difficult
parentmaterialupon whichto base eco-pedologicalinvestigationsof this type.
As a resultthegenerallinesofinvestigation
plannedin advance,foundedupon a statistical
and area-basisapproach,had to be abandoned,and a moreelastic,moreselective,method
employed. Althoughthe coarsenessof the glacial till made the selectionof sample sites
and laborious,by generalstandardsthe till is apparentlyexceptionally
both frustrating,
fine. For example,Ahlmann(1952,personalcommunication)has indicatedthat the most
strikingfeaturesof the Alaskan glacial recessionsare not onlythe extentand magnitude
finenessof the morainictill! The main diffiof deglaciation,but also the extraordinary
was
to
find
finetill to permittaking density
in
site
selection
sufficiently
culty sample
of
the
minorrelieffactorsand parent
samples, at the same time keeping variability
betweenthe
materialto a minimum,and in maintainingthe desireduniformrelationships
and
site positionand the immediatevegetationunits.This latterwas especiallydifficult
to
the conditionswereonly partlymet. Too muchweightshould not be giventherefore
the finerdetail of the time functions. However, we are confidentthat these inherent
do not mask the generaltrendsand the main characteristics.
difficulties
The mineral-soil
sampleswereall takenfrompitsat sitesselectedforsimilarityofrelief
the differences
and parent material. As effectiveclimate did not vary significantly,
to
the
time
could
be
attributed
betweenthe soil samples
states, and in the
reasonably
in the kinds of plants present.The zones sampled varied
early stages to differences
somewhatin depth,but onlyrarelywerethe soils sampledto morethan 24 in. (61 cm.):
on the youngersurfacesthe profileswere rarely sampled below 12 in. (30 cm.). The
mineral-soilsamplesweretaken fromimmediatelybelow the locationof the forestfloor
bulk densityof the fineearthwere
samples. Whereverpossiblesamplesfordetermining
taken. This was done by means of a cylinderof knownvolume (140 c.c. approximately).
Stones and gravel were includedin the bulk mineralsamples along withthe fineearth
and werelaterseparatedin thelaboratory.In the fewsubsoilswherestoninessprevented
for
effective
densitysampling,values wereassumedon the basis of thosedeterminations
otherhorizonsof the profileand thefieldnotes. The forestfloorsweresampledas a single
stratumin the earlierstages of the succession,but as separatelitter(L), fermentation
(F)
and humus(H) horizons,on the olderforests.The livingmoss on the floorsof the spruce
forestswas includedin the L-layer. Althoughthese separationswere ratherarbitrary,
theanalysesbothforthe GlacierBay material,and thatfromtheHerbertand Mendenhall
recessional moraines (to be reportedelsewhere),show that the forestfloorsampling
divisionswere meaningfuland consistent.

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ROBERT

L. CROCKER AND JACK MAJOR

433

Laboratoryanalysis
The soil samples representativeof the different
profilesand theirhorizonswere airdried and sieved througha 2 mm. screen. The relative amounts of the fineearth and
> 2 mm.fractionwererecorded,and moisturedeterminations
made on the fineearthby
at
The
105?
F.
floor
forest
materials
were
air-dried,
oven-drying
weighedand groundto
a
40-60
in
a
mesh
sieve
Mill.
pass
Wiley
Standard methodsof analyses were employedto determinereaction (pH), calcium
carbonate,nitrogenand organiccarbon. Reaction was determinedon the soil paste by
the glass electrode,and calciumcarbonateby Williams'(1948) modification
ofthe manometricmethod.Total nitrogenwas determinedby a Kjeldhal methodand organiccarbon
by the chromicacid oxidationmethodof Walkeyand Black, both as describedby Piper
(1942). The Walkey and Black percentageswere convertedto total carbon by use of
separate factors(mineralsoil x 1.29, and forestfloorx 1.04), obtained by determining
total carbon upon a number of samples by dry combustiontechniques. The weight
constituents
wereall calculatedon theoven-drybasis.
percentagesoftheabove-mentioned
For quantitativeexpressionthe weightpercentageswere convertedto weightper unit
volume of in situ soil, by using the percentageof fineearth,and the soil bulk density
values.
Bulk densitiesof the fineearth materialwere determinedfromthe separate samples
taken by the fieldmethodsalready described,and are expressedas weightof fineearth
per unitvolumeof fineearth+ pores. This value was derivedby sievingthe special field
sample to separate the >2 mm. fraction,weighingthe oven-dryfineearth, and subtractingthe volumeof > 2 mm.fraction(determinedby a waterdisplacementtechnique)
fromthe total volumeofthe wholesample. As the special densitysamplesweretakenin
positionswhichavoided coarsedebris,the proportionoffineearthin themwas somewhat
greaterthan in the largersoil columnof that sampled for generallaboratoryanalyses.
The bulk densitiesdeterminedfromthe densitysampleshad to be modified,*therefore,
wheredata wereto be expressedon the basis of a soil columnofunitcross-section,
as for
in
in.
example g./sq.m./18 (46 cm.) profile.
THE RESULTS

The analyticalresultsare presentedlargelyin the formof graphs.The graphs,in their


somewhatfreelydrawnand smoothedlines,illustratethe major effectsand moregeneral
conclusions.In the authors'opinionthe limitationsimposedby bothfieldand laboratory
of the
methods,the subjectivityof samplingnecessitatedby the inherentdifficulties
of
and
the
small
number
do
not
warrant
the
of
studied,
system
samples,
fitting exact
curvesto the data, nor to moredetailed considerationsof any but the broaderchanges
with time. The values fromwhich the curves are drawn, however,are shown on the
graphsand thispermitsalternativeinterpretations.
The data are most convenientlypresentedas theyrelate to (i) the early stages of the
plant succession,and, (ii) the chronosequenceas a whole.
* This modificationwas made as follows:

F.d,.dr
Bulk densityfineearth,generalsample=F d
fR+ R. .
F. d,
df

Where, F=% fine earth (general sample), R=% >2 mm. fraction(general sample), dr= density>2 mm.
fraction,d! =bulk densityfineearth correctedfor> 2 mm. fraction(special densitysample).

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434

Soil development
at GlacierBay

(i) The earlystagesofthesuccession


Alongthe easternside of GlacierBay the pioneerand thicketstageshave persistedfor
approximately80 years. The variable patternof plant colonizationduringthe earliest
phases presentedan opportunityfor makingsome directcomparisonsof the effectof
different
plant species upon the rate of change of soil properties. Sites were carefully
isolated fromindividualsof
selected wherea particularkind of plant was sufficiently
of
to
interaction
to
or
otherspecies reducelikelihood
slight, even negligibleproportions.
it was quite easy on terraindeglaciated30 yearsor less to findareas which
Furthermore
apparentlyhave always been devoid of macroscopicvegetation.

c0

8.0
I

6.0-

w 5.0-

4.0-L

,,

10

20

,
30

,
40

,
50

60

, 70

AGE OF SURFACE (years)

Fig. 3. Rate of changeof reactionof the surfacesoil (0-2 in.) duringthe


earlydecades followingdeglaciation.

Soil reaction(pH)

The reactionof the soil parentmaterial,the initialfinemorainicdebris,is quite high,


pH 8-0-8-4.This is largelybecause the till contains,along with granitic,schistoseand
gneissicmaterial,a considerableproportionofmarble,indeedup to 7-10 % carbonatesby
analysis.The pH ofsurfacesoilsfallsrapidlywiththe onsetof vegetationestablishment.
In Fig. 3 the reactionsofall the surface(0-2 in.) (0-5 cm.) soils analysed,whichrelateto
thepurelypioneerand shrubstagesofthesuccessionare plottedas a functionofestimated
age ofsurfaceexposure(yearssinceglacialrecession).WhiletheloweringofpH withtime
is veryapparentfromthesedata, it is also to be notedthat,especiallyin the earlystages,
surfacesofany particularage thereis a considerablerangein reaction.These
fordifferent
resultsbecome much more understandableand the variabilityis reduced if they are
consideredin relationto the incidenceand age of the colonizingvegetation,and this
arrangementis shownin Fig. 4. In this figurethe age of the Alnus, Populus and Salix
is that determinedby directgrowthincrement(ring)analysis*of the individualshrub
* We are gratefulforthe tree and shrubsamples taken forus by D. B. Lawrence and Roland Schoenicke,
and later analysed by D. B. Lawrence.

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ROBERT

435

L. CROCKER AND JACK MAJOR

but the estimateis


under which the soil was sampled. The age of Dryas is indefinite,
considereda reasonableapproximation.
A significant
conclusionto be drawnfromFig. 4 is that the type of plant markedly
affectsthe rate ofchangeofpH ofthe mineralsoil. The bare surfaces,that is the surfaces
whichhave apparentlyalways been devoid of macroscopicplants showedpracticallyno
reductionin pH over this 30-yearperiod.This means that,in the absence of vegetation,
the leachingeffectper se is veryslight. On the otherhand all surfaceswithvegetation,
eventhoughit be verysparseas withthePopulus and Salix sites,showgreaterchangesin
the data are not as completeas we wouldlike exceptin the case
reaction. Unfortunately,
of findingSalix or
of alder (Alnus crispa). This is largelybecause of the great difficulty
Populus olderthan about 20 yearswhichby that timeis not closelyassociated withthe
morerapidlyspreadingalder.
x

8.0-

CT

0c
*a.
0
|
70-

^K

*/

/
7.0

./

-Bare surface
-1Populus
trichocarpa
^
^Dryas spp.
^
Slaix Barclayi

>
r
C

1.0 2
o

.^-2.0

-6.0 I
01

40-

40

AGE

IN YEARS

Fig. 4. Rate ofchangeofreaction0-2 in. horizonrelativeto typeofvegetativecover; rate ofchange


in calciumcarbonatecontentunderAlnus.

The most strikingfeatureof Fig. 4 is the evidenceof the remarkableacidifyingeffect


of the alder. Under alder the uppermostsoil horizonsare reducedfroma pH of 80, or
more, to pH50, within35-50 years. This is certainlya remarkableeffect. Its early
stagesare wellillustratedby a sequenceofsamplestakenfroma 29-year-oldsurfacenear
Goose Cove in a transectfromthe oldest (initial)memberof an expandingalder thicket
to the bare groundbeyondthe thicket-a distanceof approximately10 yards.
Cover
Reaction (pH)

Bare
7-9

9-year
alder
7-2

12-year
alder
7-1

18-year
alder
6-5

Alderhas an abundantleaf fall,and the shrubhabit of growthand highsociabilityof


the alder plants make them effectivein trappingand holdingsome of the leaf fall of
otherspecies like Salix and Populus. In the early stages the litterunder alder has a
reactionofpH 5-6-6-1. No doubt atmosphericdust additionsin the vicinityoflargebare

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436

Soil developmentat Glacier Bay

areas tend to reduce the acidity. At all events the decomposingalder littertends to
becomemoreacid withtime,and in the finalstages of alder dominancereactionvalues
of 4-2-4-6are usual.
The leachingofcalciumcarbonateunderthe Alnus is veryrapid as would be expected
fromthe reactiontime-curves.The values forcalcium carbonate,however,appear to lag
a littlebehindthe pH values, whichsuggeststhat in the leachingprocesssmall 'loci' of
calcium carbonatemay not be completelydissolvedby the descendingacidic solutions,
so that whilethe bulk of the fineearthmay be slightlyacid, relictloci of marblemay be
preserved. In any case under 18-year-oldalder, growingon a surfacewhose age is
estimatedat 31 years,calcium carbonatehas fallenfrominitial values of about 5 % to
0-3%. Adjacent areas of similarage devoid of macroscopicvegetation,containedsome
3-8% calciumcarbonatein the equivalentsurfacehorizon. In anotherregion,bare ofice
forapproximately20 years the percentageof CaCO3 in the fineearthhad been reduced
fromoriginalvalues of 7-9 % in the surface2 in. to 6, 5 and 3-4 %, underSalix, Populus
and Dryas respectively.Evidentlyotherspecieshave muchless affectthan alder on rate
of leachingof CaCO3.
6.0
?
J

5.0-

o, 4.03.00
t:

2.0-

w
I--- 1.0-

20

30

4,

50

AGE OF ALDER (years)


Fig. 5. The accumulationof organicresidueson the mineral
soil underdevelopingalder thickets.

organicresidues
Above-surface
Alderextendsin theearlystagesas rapidlyexpandingand denseisolatedthicketswhich
soonerorlaterincreasein numberand finallycoalesceas a continuousalderthicket.These
thicketsare excellenttrapsforthe annual litterfall,and above-surfaceaccumulationsof
organicresiduesbuild up fairlyrapidly.The rate of accumulation,expressedin termsof
kg./sq.m.and plottedagainst age of alder plants is shownin Fig. 5. Within40-50 years
5-6 kg./sq.m.of above-surfaceorganicresidueshave accumulatedin a mantlewhichis
approximately6-7 cm. deep. The curve does not pass throughzero because a periodof
severalyearsmustelapse afterestablishment
beforean alderbush accumulateslitter.The

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ROBERT

L. CROCKER AND JACK MAJOR

437

curve is remarkablylinear until at least 40 years of alder age, with an annual rate of
accumulationofsome 150 g. litter/sq.m.As pointedout already,thislinearincreasedoes
not continueindefinitelysince, by normal plant succession,spruce (Picea sitchensis)
eliminatesthe alder.
Dryas mat amountsto 0-9-1-4kg./sq.m.in 20 years. NeitherSalix nor Populus show
accumulationsof thismagnitudein the fewsituationswe sampled.

360320E

280-

240E
CD200-

w 160-

Forest Floor

120z

_
MinerolSoil (0-18 )

4016

32

AGE OF Alnus

48

crispo

(years)

Fig. 6. Accumulationof nitrogenin the mineralsoil and forest


floorunderAlnus crispa.

The carbon and nitrogenprofiles


The accumulation of total nitrogen and organic carbon in the mineral soil and the litter
residues under alder is summarized in Figs. 6 and 7. In both cases the rate of accumulation relative to age of alder is almost linear. There is, however, the suggestion of a
decrease in rate of accumulation of both carbon and nitrogen in the mineral soil by
48 years; this suggestion is considerably stronger for the carbon and nitrogen of the
forestfloor.The total carbon and nitrogen accumulation can be interpretedas the sum of
two linear functions to 40 years: while the mineral soil accumulates carbon and nitrogen
from a very small but definiteamount in the initial state (see later discussion), the forest
floordoes not begin to accumulate for several years after establishment of the alder. The
annual rates of accumulation of carbon and nitrogen under alder are approximately as
follows (g./sq.m./year):
Carbon
Nitrogen

Mineral
soil
35
2-6

Forest
floor
50
4-2

Almost 4 kg. of carbon and 0-3 kg. of nitrogen per square metre have accumulated under
50-year-old alder. This represents some 34,800 lb. carbon and 2760 lb. nitrogen per acre.
J. Ecol. 43

28

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438

Soil developmentat Glacier Bay

It is not a measureof the nitrogenand carbonin the ecosystemof course,but onlythat


portionof it in the soil and forestfloor-the livingplants and animals are neglected.
Nevertheless,the mean annual rate of net accumulation(gainsless losses) to thispart of
theecosystem(soil+ forestfloor)for50 years,of700 lb. organiccarbonand 55 lb. nitrogen
per acre is highlysignificant.
After40 years approximately50-60% of the carbon and nitrogenwas accumulated
in the forestfloor.The distributionof nitrogenwithdepth forrepresentativeprofilesis
shownlater (Fig. 13). The patternof accumulationof organiccarbonis very similarto
that fornitrogenexceptforthe different
magnitudeof the quantities. Both showinitial
build-upin the surfacehorizonsfollowedby gradual extensiondownwards.

4-

E
Ito

<

"/

forest
foor underAlns cripa.

6
32
AGE OF Alnus crispa

48
(years)

Fig. 7. Accumulationof organiccarbonin the mineralsoil and

The Dryas mat is very strikingin the early stages of the Glacier Bay colonization,
although the succession is destined to pass through the alder thicket stage to final spruce-

hemlockdominanceirrespectiveofDryas havingbeen in the sequence. Nevertheless,


the
accumulationof nitrogenand carbon underand withinthe Dryas mat is of interestand
some analysesare givenin Table 2. Unfortunately
the age of the mat at any particular
site cannot be reliablydeterminedand we can only estimatethe general age of the
surface. For comparisonthe data fromwithinan alder thicketadjacent to the Dryas
samplingsitesin the Hugh Millerarea, and froma bare surface,approximatelyzero age,
near the dead ice remnantbehindNunatak Nob, in AnchorageCove, are included.
It is to be noted that data forthe Dryas mat refersto both the livingDryas plus any
organicresiduesupon the mineral-soilsurface.In the case of alder the surfacelitterwas

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ROBERT

L. CROCKER AND JACK MAJOR

439

sampled and analysed and, except unavoidably,no living alder was included. It is
obvious fromthe table that the accumulationof nitrogenin the Dryas-soilsystemis
considerablewhen comparedwiththe initialtill,but alder is neverthelessa much more
effectiveagent. Not onlyis theremorelitterunderalder than the total Dryas mat, but
its percentagenitrogenis twice as high.
Table 2. Accumulationoforganiccarbonand nitrogen
undervaryingconditions
Cover: locality
Bare except sparse moss:
Goose Cove
Bare surface:AnchorageCove
Bare except algal crust:
Goose Cove
Dryas: Goose Cove (1)
Dryas: Goose Cove (2)

Surface
age
(years)
31
24
24

Horizon
depth
(in.)
0-2
3-6
0-6
0-0-4

24-25

Mat
0-2
Mat
0-2
Mat
0-1
1-3
3-6
Mat
0-1
1-3
3-6
0-2
3-6
0-2
3-6
Litter
0-2
3-6

24-25

Dryas: Hugh MillerInlet (1)


-

Dryas: Hugh MillerInlet (2)


Populus trichocarpa:Goose
Cove (19 years old)
Salix Barclayi: Goose Cove
(20 years old)
Alder: Hugh MillerInlet
(19-20 years old)

24-25
24-25
-

Nitrogen
x
g./sq.m.
%
0-015
7-7
5-4
0-006
3-3
0-0075
0-093
1-313
0-019
1-330
0-021
1-036
0-025
0-0065
0-002
1-075
0-030
0-010
0-007
0-018
0-007
0-027
0-004
2-076
0-063
0-005

12-20
9-46
18-6
12-3
20-50
5-96
3-39
2-29
20-3
7.3
6-8
7-4
12-4
7-0
19-3
4-0
46-0
28-4
5-4

Organiccarbon
-.g./sq.m.
%
0-17
87
0-04
40
43
0-02
189
32-3
0-32
33-8
0-37
32-5
0-54
0-15
0-08
30-6
0-60
0-16
0-20
0-34
0.09
0-49
0.09
28-4
1-08
0-08

300
160
473
217
640
132
81
79
570
146
115
209
234
97
345
96
630
489
83

(ii) The complete


chronosequence
The vegetationdevelopmentalsequencesfromthe mostrecentlydeglaciatedregionsto
the terminalmorainepass throughthe stages alreadymentioned-bare surface,pioneer
(transition
stage (Salix, Dryas, Epilobium,etc.) alder thicket,spruce-cottonwood-alder
As
in
the
and
forest.
mentioned
earlier,
spruce
studying temporalchangesin soil
stage),
a
occur.
this
of difficulties
which
number
sequence
properties
accompany
One problemis the uncertaintyof the absolute surfaceages in the lowerBay. From
1880 onwardsthe rate ofrecession,and positionofthe ice frontat varioustimeshas been
veryaccuratelyrecorded. But littledirectevidenceis available about thepositionsofthe
ice at particulartimes duringthe recessionfromthe terminalmorainein the Bartlett
Cove area to the 1880 position. Soil samples were taken along the easternside of the
lower Bay representativeof the spruce-cottonwood-alder
stage (Sandy Cove) and the
the
exact
Cove
and
Bartlett
but
forest
Track
Cove),
(Bear
ages of the surfacesare
spruce
somewhatdoubtful.We have consideredthat our oldest site, on the innerside of the
terminalmoraineat BartlettCove, has been a stable surfaceand ice-freefor182 years.
This is in reasonableagreementwith Cooper's estimate(167-217 years),but it is based
moreparticularlyupon tentativeconclusionsofLawrence(personalcommunication).The
ages ofthemorainesurfacesat Bear TrackCove and Sandy Cove have been consideredby
28-2

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at GlacierBay
Soil development

440

Lawrence,afteranalysingall the evidenceand a studyof recentaerial photographs,as


122 and 95 years. Undoubtedlythereare errorsin these approximations,but they are
not likely to be great. In any case the developmentof soil propertiesrelative to the
successionalstage in the vegetationis the significant
thing.
The variabilityof the patternof vegetationdistributionduringthe pre-alderthicket
stage also causes problemswhenconsideringthe sequenceas a wholebecause,as we have
demonstrated,it is reflectedvery clearlyin soil variability.This variabilityhas been
resolvedherewhenplottingthe data forthe overallchronosequence,by summingall the
data available fora particularpre-alderthicketsurfaceand derivinga mean value. While
thisresolutionis expedient,and the onlyone whichappearedreasonable,it is nevertheless
probablyonly a roughapproximationof what a statisticalstudy,had it been possible,
would have produced. Here again the erroris not likelyto be great enough to mask
generaltrendsof the type withwhichour paper is concerned.

o
IcJ 1.5

i 1.2

iI

1.0-

^^

0.8-

0.8
0.7 _~--.e___
100

200

ESTIMATED SURFACE AGE (years)


Fig. 8. Bulk densityof the fineearthof subsurfacesoil as a function
of time since deglaciation.

Bulk densityoffineearth
of bulk densityare not onlydifficult
but
In a mediumlike glacial till, determinations
involve a large experimentalerror. It is likelythat in the earlieststages thereare some
settlingeffectswith consequentincreasesin bulk density. From the outset,however,
on the bare surfaces,
influencesare significant
freezingand thawingand even solifluction
ofthe alder-willow
thickets. Over
but theirimportanceis reducedwiththe establishment
the chronosequenceas a wholethereappears to be a continualdecreasein bulk density
of the fineearth.This is moreor less confinedto the uppermosthorizonswhererootsare
heavily concentrated.The values forthe surfacehorizondecreasefrom1-4g./c.c.in the
stages. The generaleffectsare shownin
early stagesto 0-8g./c.c.in the spruce-dominant
Figs. 8 and 9. They are rathersimilarto the declinein bulk densityreportedin the Shasta

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ROBERT

L. CROCKER

441

AND JACK MAJOR

chronosequencein California(Dickson & Crocker,1953). Their close parallelismto the


organicprofileleaves littledoubt of a geneticlinkagebetweenthe two properties.
Reaction(pH)
The rapid declinein soil reactionceases afterthe alder thicketstage. The sprucelitter
is no moreacid than the late alderlitterand thereappears to be littlefurthersignificant
changein reactionof eithermineralsoil or forestfloorduringthe periodof sprucedominance investigatedhere except a slightdownwardextensionof the acidity. These facts
are well illustratedin Figs. 10 and 11.
BULK DENSITY FINE EARTH (gms.FE/(vol.FE. + pores))
08

0.6

1.0

1.2

22
S^^yri.'

Is

1.4

\,

6-

a-

II

wj 12-

ages.
Fig. 9. Bulk densitydepth functionsforprofilesof different
8.0-

7.G

z?

0
F

6.0 -

\5s xrJ
L----Forest
x

4.0

50

100

Floor

150

200

AGE OF SURFACE (years)

Fig. 10. Reaction of litterresiduesand surfacehorizonsof the


mineralsoil as a functionof age of surface.

Organiccarbonand nitrogen
The organiccarbon and nitrogentime functionsforthe sequence as a whole do not
presentsuch a regularpictureas duringthe earlysuccessionalstages. This is due partly
to the difficulty
experiencedin the oldersitesin samplingat a standardsituationrelative
to the previousand presentvegetationunits,and to the small numberof samples.There

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at GlacierBay
Soil development

442

whichmakeforvariabilityincluding
are also a numberofotherinvestigationaldifficulties
the fact that owingto the variable amountof > 2 mm. fractionin different
profilesthe
amountsof fineearthare not strictlycomparable. Always a source of variability,this
factorbecomesmoreimportantin the oldersoilssampledwherethe percentagesof coarse
REACTION (pH)
4.0
180

5.0

6.0

70

8.0
28yrs
yrs.

,3\\570yrs.yrs.

yrs.

122yrs

m) 6 -

i"

\\\

w 12

18in the reactionchangeswithdepthin


Fig. 11. Differences
soils of different
ages.
Pioneer

Alder

stages prc

f ote

5.013

4.0

/
X/

X /

01
3.0
0

10

100

150

20-

1.0

50

ESTIMATEDSURFACEAGE (years)
Fig. 12. Organiccarbonaccumulationwithinthe mineral
soil and forestfloor.

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200

ROBERT

L. CROCKER AND JACK MAJOR

443

materialare higher. Despite this a usefulpictureof the rate of accumulationof organic


carbonand nitrogenrelativeto time (vegetationsuccessionalstages) is obtained.
Overthe periodofthe wholechronosequenceorganiccarbonaccumulatesin the upper
18 in. (46 cm.) of mineralsoil at a fairlyregularrate of about 15 g./sq.m.per annum
(Fig. 12).
ORGANIC CARBON(Kgs./metre/inchcolumn)
0.1

Oy

LOyrs.

024
r

20
I -

/
Ig
Fig. 13. The distributionof organiccarbonwithdepthin soils
of different
ages.
Pioneer -

es

Alder sta.

250
0)
?

200 -

E
z

//~?

I -I
0g

50

100

150

200

ESTIMATEDSURFACEAGE (years)
total
in
the
nitrogencontentof soils on surfacesof varyingages.
Fig. 14. Differences

Thereis an extensionin depthof the organiccarbonprofilewithtimebut even in the


organiccarbonresidesin theupperoldestprofileof thesequence60 % of themineral-soil
most 6 in. (15 cm.). The patternof distributionof organiccarbon with depth,and the
downwardextensionas soil developmentproceeds,is shownin Fig. 13.
Total nitrogenfollowsmuchthe same accumulationpatternas organiccarbon(Fig. 14)

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Soil developmentat Glacier Bay

444

forestthereappears to be not onlya reductionin the


exceptthat in the spruce-dominant
rate ofaccumulationbut an actual declinein total amounts.This is moreapparentin the
forestfloor(see p. 445) than the mineralsoil.
profile
Duringthe vigorousalderthicketstage nitrogenaccumulatesin themineral-soil
at theaveragerate*of 1.5 g./sq.m.per annum,and in theforestfloor+ mineral-soil
system
at about 4-9 g./sq.m.per annum. Duringthe alder-sprucetransitionperiodthereis very
little increase in the wider system(mineralsoil+floor), but the total nitrogenin the
mineralsoil appears to continueto increaseslightlyas translocationfromthe forestfloor
organicresiduescontinues. However,the depth functionsfornitrogen(Fig. 15), which
NITROGEN(gms./metre/Il
inchcolumn)
4.0
TPyrs
-C

8.0
28yrsy

16

12.0

20.0

sys.

15

II

Fig. 15. The distributionof total nitrogenwithdepth.


0
(r

ALDER
34

30-4

o ioJ
.,

a:

-+

yr.

.0

5
5

1
lb
15,

THICKET
older

STAGE
48

20
20

I
0

SPRUCE

older
yr.

i 1 i
l105

I2

20

FOREST

Beor rrockCove

05

Bartlett Cove

b
20

J
0

5 ,0

15
'5

DEPTH (inches)
ratiosand theirvariationwithdepthat
Fig. 16. Carbon-nitrogen
the alder thicketand spruce-dominant
foreststages.

contrastverymarkedlyforthe oldersoils withthosefororganiccarbon,suggesta major


changein the patternof nitrogenwithinthe profilewiththe developmentof the spruce
forest. In contrastto organiccarbon thereis a fall in the amountsof nitrogenin the
upper parts of the profileduringthe earlysprucedominance.This is verywell reflected
withinthe carbon-nitrogen
ratios,a few of whichare presentedin Fig. 16. While the
ratiosforboth mineralsoil and litterresiduesat the alderthicketstage are mostlyin the
range13-15,withthe declineofalderand the developmentofthe spruceforest,theforest
floorand uppermosthorizonof the mineralsoil have ratiosof 30 or more.
* This average includes all
analyses for surfaceswhere thicket stage exists irrespectiveof the actual
cover.

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445

ROBERT L. CROCKER AND JACK MAJOR

Theforestfloor
As mentionedearliera stonypavementis usuallydevelopedbeforetheshrubvegetation
is establishedand prominent.Withthe alderthicketstage organicresiduesbuild up, and
rest upon this pavement,whichmust greatlyinfluencethe rate and patternof translocationof materialsfromthe forestfloorto the mineralsoil. The forestfloorbuilds up
rapidlyunderalderthicket,at a rate ofabout 85 g. drymatter/sq.m.
perannum,and this
rate is not greatlyreduced with spruce dominance.The organicresiduescontinueto
increase throughoutthe sequence until the oldest sites sampled are some 5 in. thick
(13 cm.), and weigh(oven dry) about 9-10 kg./sq.m.(Fig. 17).

8.0-

W 6.0D

4.0w

.I

,
50

100

.I

150

200

ESTIMATED SURFACE AGE (yeors)


Fig. 17. The rate of accumulationof above-surfaceorganicresidues.

One ofthe strikingfactsabout theforestfloorin the fieldwas thelargenumberofroots


usuallypresentin its lowerpart,and in the transitionzone to the mineralsoil. This was
fromthe well-developedalderthicketstagethroughto the earlystagesofthe
noteworthy
forest
whichis as faras our studygoes. The conclusionseemsinescapablethat the
spruce
draw
most
oftheirnutrientsfromthisregion.This concentration
ofrootsis not so
plants
total
when
one
the
functions
for
considers
surprising
depth
nitrogen.
The reactionoftheforestfloorresiduesdo notvarygreatlyfromthe alderthicketstage
onwards. For the most part they fall between pH 4-0 and 4-6, and, where sampled
separately,the mostcompletelydecomposedlowerzone (H) is moreacid than the fresh
litter(L) and fermentation
(F) zones.
the
data on theforestfloorsare thosefromthe organiccarbon
most
Perhaps
interesting
and nitrogendeterminations
because theyappear to indicateclearlythe mainreasonsfor
the changein rates of nitrogenand carbonaccumulationwhichfollowedthe elimination
ofalderand the emergenceofthe spruceforest,and whichweresuggestedby the mineral
soil analyses. Whilecarboncontinuesto increaseat a reducedratein thefloorofthespruce
forest(Fig. 12) thereis a markedfallin absolutenitrogenlevel-that is, thereis a period
of negativeaccumulation,an actual net loss fromthe forestfloor(Figs. 14, 15). The C/N

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446

Soil developmentat Glacier Bay

ratiosforthe forestfloormaterialincreasefromapproximately13 at the alder stage,to


33-36 fordifferent
partsofthe sprucefloors.The ratiosin theupperlitterhorizonsappear
is small and may
to be higher(35-36) than in the lowerzones (33-34), but the difference
not be significant.
DISCUSSION

Amongstthe many featuresof the relationshipbetweenthe developmentof vegetation


thantherateofaccumuand soilgenesisin the GlacierBay regionnoneis moreinteresting
lation ofnitrogen.The presentstudydoes not providedirectevidenceforthe mechanism
ofnitrogenfixationin the soil-vegetation
portionofthisecosystem,but it fitsproperly,as
an exploratoryinvestigation,into a studyof that problembeingpursuedas a long-term
projectby ProfessorD. B. Lawrence.
The mean annual net accumulationrate fornitrogenof 55 lb. per acre (62 kg./ha.)per
annum duringthe alder thicketestablishmentis veryimpressive.Amongstthe species
otherthan alder and willowprominentduringthe earlystages ofplant successionon the
Richards)and Shepherdia
deglaciatedsurfacesare Dryas spp. (especiallyD. drummondii
canadensis(L.) Nutt. The rootsof both of these,like alder,have mycorrhizaewhichare
believedcapable offixingnitrogenfromthe atmosphere(see Lawrence,1953). It is very
evidentin thefieldthatAlnus,Dryasand Shepherdiaare all capable ofgrowingvigorously
at the verylow nitrogenlevels ofthe initialglacial debris.The data fornitrogenaccumuthanDryas.
lationundertheDryas mat (Table 2) indicatesthatthe alderis moreeffective
A comparisonofthenitrogenpercentagesin theveryyoungtillwitholderbare surfaces
(up to 31 years)indicatesan accretionof nitrogenin the surfacehorizonswithage. That
is, thereappears to be evidenceof a low level ofnitrogenfixationor accumulationin the
absence of macroscopicvegetationaltogether.The relativehighpercentageof nitrogen
immediatelyunderthe remnantsof an algal crust,forexample,is suggestivethat these
associatedwiththemare capable offixation.On theotherhand,
algae, ormicro-organisms
it mightwell be that this relativelylow orderof nitrogenfixationis purelya selective
surfaceadsorptionphenomenaof the type suggestedby Ingham (1950).
At all eventsit is clear that, duringthe initialstages,the rate of developmentof the
soil nitrogenprofile,like manyothersoil properties,forexamplereaction,dependsupon
which
Thereis an areal patternof soil profileformation
the patternof plant distribution.
at
establishment.
This
is
first
of
the
areal
reflects
largely
pattern pioneerplant
partially
relatedto factorsoutsideoursystemdependingupon suchthingsas thelocationand state
of neighbouring
ecosystems,the preadaptationof the disseminules,and the accidentsof
dispersal.
The decline in the rate of nitrogenaccumulationfollowingeliminationof alder and
dominanceof spruce,and the subsequentnet loss of nitrogenfromthe mineralsoilforestfloorsubsystemsuggestedby the grossanalysisforthe mineralsoil,is obviouslya
real effect.Althoughthe values forcarbonand nitrogenin theoldestsoil seemlowerthan
expected and one mightsuspect some variationinherentin the methodof study,the
evidenceforloss ofnitrogenis not affectedhowever.When all the data are consideredas
a whole,takinginto account compositionof the forestfloor,and especiallythe carbonnitrogenratios and the way theyvary both in .thefloor,and withdepth in the mineral
soil,therecan be littledoubt that the developmentalvegetationchangeshave influenced
very directlyboth the patternof nitrogendistribution,and the absolute amounts of
nitrogenin the mineralsoil and mineralsoil-forestfloorsystems.

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ROBERT L. CROCKER AND JACK MAJOR

447

thealderthicketstage,and thesharpreducThe rapid build-upof nitrogenthroughout


tionin nitrogenin thefloorand uppermineralsoil oftheyoungspruceforestfollowingthe
noduleson
eliminationof alder,is verystrongevidenceforthe factthat the mycorrhizal
the rootsof alder do have the capacityfornitrogenfixation. Followingthe elimination
of the alder one can readilyvisualize the spruceforestdevelopingat the expense of the
nitrogenaccumulatedin earlierstages. Mostofthisnitrogenmay wellbe retainedwithin
the widervegetation-soilsystem,and little or none of it may be lost altogether. It is
logical,too, in view of the root concentrationin the forestfloorand upper mineralsoil
thatthereductionofnitrogenshouldoccurfirst,and be greatesthere.The loss ofnitrogen
fromthe mineralsoil is not to be lookedupon as a permanenttrend,forfurther(developmental) changes in structureand compositionof the forestdo occur although the
is beyondthe scope of this inquiry.
investigationof theirnatureand significance
SUMMARY

The developmentofcertainsoil propertiesin relationto thebroaderfeaturesofvegetation


dynamicsand as a functionoftimehas been studiedin the extensiverecentlydeglaciated
areasoftheeasternshoresofGlacierBay, Alaska. The physicalnatureofglacialtillmakes
mediumforstudiesof this type but it has been possibleto demonstratea
it a difficult
effectsof plants on soil formation.
numberof interesting
In the pioneerstagesofsuccession,as the rate of changein soil propertiesis dependent
ofplant colonization,the accidentsand factorsofdispersal
upon the actual micro-pattern
and establishmentare highlysignificant.The areal patternof soil propertiesis very
variableat thisstage. The moreor less continuousdevelopmentofthe alderthicketleads
to greateruniformity.
Under alder the reaction of uppermosthorizonsof the glacial till is reduced from
pH 8-0to less than pH 5-0 within35-50 years. Duringthe same periodcalciumcarbonate in the fineearth is reduced frominitial values of the order of 5% to negligible
quantities. Under otherspecies,the rate of changeis muchless, and on areas devoid of
vegetationit is still slower.Within40-50 years 5-6 kg./sq.m.of above-surfaceorganic
residues,6-7 cm. deep,withpH 4-2-4-6accumulateunderAlnus. The 18 in. (45 cm.) deep
mineral-soil
profileand forestfloorcombinedhave accumulatedalmost4-0kg. of organic
carbonand 0-3kg. of nitrogenper sq.m. beneath50-year-oldalder.
The Dryas mat is also capable ofaccumulatingconsiderablequantitiesofnitrogen,but
as alder. Significant
nitrogenaccretionsalso occurin the
Dryas is not nearlyas effective
absence of macroscopicvegetation.
The chronosequenceas a wholecoversthe mainphases in vegetationchangesfromthe
ofthespruce(Picea sitchensis)
initialcolonizationofthebare surfacesto theestablishment
dominatedforest.The changesin the mineral-soilpropertiesof bulk densityof the fine
earth,reaction,organiccarbon, calcium carbonate and total nitrogenare traced, and
relatedto broad vegetationchange.The amountofforestfloormaterial,its reaction,and
its carbonand nitrogencontentswerealso determined.An absoluteloss ofnitrogenfrom
the mineralsoil-forestfloorsystemis recordedwith the eliminationof alder and the
ratios forthe upper mineralsoil and
emergenceof the spruce forest. Carbon-nitrogen
forestfloorincreasefromthe orderof 12-16 to morethan 30 over this stage.
Organiccarbon increasesthroughoutthe whole sequence. Soil reaction(pH) declines

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448

Soil developmentat Glacier Bay

rapidlyat firstbut apart froma downwardextensionof acidity does not change much
afterthe alder thicketstage. The bulk densityof the fineearthfractionof the surface
horizonsappears to fallthroughout,but is moresignificant
duringthe earlystages.
REFERENCES
COOPER, W. S. (1923). The recent ecological historyof Glacier Bay, Alaska. Ecology,6, 197.

COOPER,W. S. (1931). A thirdexpedition to Glacier Bay, Alaska. Ecology, 12 (1), 61-95.


COOPER,W. S. (1937). The problem of Glacier Bay, Alaska: a study of glacier variations. Geogr.Rev.
27 (1), 37-62.
COOPER, W. S. (1939). A fourthexpedition to Glacier Bay, Alaska. Ecology,20 (2), 130-59.
DICKSON, B. A. & CROCKER,R. L. (1953). A chronosequence of soils and vegetation near Mt. Shasta,
California. Part III. J. Soil Sci. (in the Press).
FIELD, W. O., JR. (1947). Glacier recession in Muir Inlet, Glacier Bay, Alaska. Geogr. Rev. 37 (3),
369-99.
E. (1941-50). Flora of Alaska and Yukon. Lunds UniversitetsArsskrift.
HULTEN,
INGHAM, G. (1950). The mineral content of air and rain and its importance to agriculture. J. agric.
Sci. 40, 55-61.
LAWRENCE, D. B. (1953). Development of vegetation and soil in south-eastern Alaska, with special
referenceto the accumulation of nitrogen. Final Report ONR, Project NR 160-183.
LAWRENCE, D. B. & ELSON, J. A. (1953). Periodicity of deglaciation in North America since the late
Wisconsin maximum. Geogr.Ann., Stockh.35 (2), 83-104.
PIPER, C. S. (1942). Soil and Plant Analysis. Adelaide (Australia): The Hassell Press.
THORNTHWAITE, C. W. (1948). An approach toward a rational classificationof climate. Geogr.Rev. 38
(1), 55-94.
WILLIAMS, D. E. (1948). A rapid manometricmethod forthe determinationof carbonate in soils. Proc.
Soil Sci. Soc. Amer. 13, 127-9.

(Received2 April 1954)

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