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Nordic Society Oikos

Heavy Metals in Subalpine and Alpine Soils of New Hampshire


Author(s): W. A. Reiners, R. H. Marks and P. M. Vitousek
Source: Oikos, Vol. 26, No. 3 (1975), pp. 264-275
Published by: Wiley on behalf of Nordic Society Oikos
Stable URL: http://www.jstor.org/stable/3543497
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OIKOS 26: 264-275. Copenhagen 1975

Heavy metals in subalpine and alpine soils of New Hampshire

W. A. REINERS

R. H. MARKS

P. M. VITOUSEK

Department of Biological Sciences, Dartmouth College

Reiners, W. A., Marks, R. H. and Vitousek, P. M. 1975. Heavy metals in subal-

pine and alpine soils of New Hampshire. - Oikos 26: 264-275.

Northern New England is subject to deposition of remotely derived air-borne

pollutants. Mountain ecosystems of the region are particularly exposed because

orographic precipitation and windy, cloudy conditions contribute to higher

deposition rates. This study describes the concentrations and total quantities of

Pb and Zn in organic horizons of soils distributed over an elevational gradient.

Pb concentrations are high for a remote area, and comparable with many heavily-

travelled roadsides. Zn concentrations fell within a broad range of levels pre-

sumed to represent natural conditions. The vertical distribution of Pb suggests

that levels are still rising; the elevational distribution suggests that deposition

rates are highest in subalpine forests where there is a combination of high winds

and ample interceptive plant surfaces. High elevation ecosystems may serve a

valuable role as amplifying systems for the detection of certain pollutants, and as

convenient systems for understanding the pattern of deposition and the ecological

effects of pollutants.

W. A. Reiners and R. H. Marks, Dept of Biological Sciences, Dartmouth College,

Hanover, New Hampshire 03755, USA. P. M. Vitousek, Dept of Zoology, Indiana

University, Bloomington, Indiana 47401, USA.

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Accepted 10 January 1975

? OIKOS

264 OIKOS 26:3 (1975)

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and aerosols, including heavy metals. If montane eco-

1. Introduction

systems of this region receive unusually high depositions

The long range transport of heavy metals from centers of heavy metals, we might then hypothesize that heavy

metal concentrations will be high in whichever compart-


of human activity to the biosphere at large is now a

ments of these ecosystems have the greatest affinity for


fairly well-recognized phenomenon (Elgmork et al. 1973,

Anon. 1972, Murozumi et al. 1969, Riihling and Tyler these elements.

1968, Weiss et al. 1971, Zoller et al. 1974). The transport Soils have a very high affinity for heavy metals,

especially clay or organic colloidal components (John


of heavy metals as small aerosols (Lee et al. 1968,

Natusch et al. 1974) together with their potential for et al. 1972, Lagerwerff 1967, Stevenson 1972). The soils

interaction with biota (Rdhling and Tyler 1968, Hagen of high elevation balsam fir forests ( > 1272 m) and fir

and Langeland 1973) provides an analogy with radio- krummholz of the White Mountains are variable, ranging

nuclides derived from nuclear devices, or with volatile from heavy mats of organic matter over rock and some

pesticides. weathered rubble, to shallow podzols with thin B

Deposition of air-borne heavy metals is related to horizons over rock. Substrates of alpine tundra range

several factors. These include distance and direction from scree slopes to deep bog soils (Bliss 1963). In all

from sources with respect to prevailing winds (Lazrus of these general ecosystem types, soil organic matter is

et al. 1970, Little and Martin 1972, Nieboer et al. 1972); probably the principal pool for retention of heavy

intensity and frequency of washout by precipitation metals. Following this reasoning, this study was designed

(Junge 1963, Ter Haar et al. 1967) and aerosol capture to test these hypotheses:

by intercepting surfaces (White and Turner 1970). A 1) Heavy metal concentrations and contents of organic

consideration of these factors suggested that deposition horizons would increase with elevation through the fir

of heavy metals might be particularly high on the upper zone in response to higher exposure with altitude. Above

slopes of New England mountains. The region as a whole the fir zone, concentrations would then decline in fir

is subject to remotely derived air-borne pollutants in krummholz and alpine tundra due to a decline in inter-

the form of nitric and sulfuric acids (Johnson et al. ceptive vegetative surfaces.

1972, Likens and Bormann 1974). Since precipitation 2) Concentrations would increase with depth in the

tends to increase with altitude, total deposition might organic horizons due to the normal turnover patterns of

be expected to increase at higher elevations, although carbon in organic layers versus the conservative behavior

concentration of precipitation may not be different. of the metals.

Furthermore, these high elevations are very windy and 3) Heavy metal concentrations and total soil contents

frequently cloudy. Such conditions increase the effective- would be quite high for a remote area, possibly in the

ness of aerosol and water droplet capture through range of those found in roadsides or areas immediately

horizontal interception, further increasing the potential downwind of smelters.

for deposition.

Schlesinger et al. (1974) measured bulk precipitation

2. Methods

(Whitehead and Feth 1964) and its heavy metal content

on Mt Moosilauke in the White Mountains of New 16 sites were sampled on the west-facing slopes of Mt

Hampshire for four months in 1971. Extrapolating from Moosilauke (710 50'W, 440 1N) in the White Mountains

these data to 12 months, we estimated that the Pb of New Hampshire. The sites were selected on the

deposition rate was approximately 20 mg m-2 yr-1. This


basis of representativeness of the general elevation,

figure is considerably lower than deposition calculated sufficient size to permit sampling, and absence of recent

for urban areas (346 mg m-2 yr-1, Chow and Earl 1970)
disturbance, either man-caused or natural. The lowest

but well above deposition rates in other remote areas site at 692 m was selected to characterize the northern-

(1.6 mg m-2 yr-1, Chow and Johnstone 1965). Since


hardwoods ecosystem type found on the lower slopes up

bulk precipitation does not take into account aerosols to approximately 760 m (Bormann et al. 1970). The next

intercepted by the vegetation, this was considered to be a


site at 1036 m characterized spruce-fir ecosystem types

lower limit for actual deposition.


found between 760 and 1280 m.

In order to account for interception, Schlesinger Sampling was more intensive between 1280 and 1400 m

and Reiners (1974) measured deposition in buckets where we expected maximum heavy metal deposition

bearing artificial conifer foliage at a 1372 m station on


due to the cloudy, windy climatic conditions, and also

Mt Moosilauke. Results of that study suggested that Pb to the density of interceptive surfaces of the evergreen

deposition might be higher than 135 mg m-2 yr-1, a rate


vegetation. Nine sites were sampled in the fir forest and

comparable with those found in cities.


fir krummholz zones, ranging from 1359 to 1393 m

While this is a gross estimate, the deposition rate


elevation. Dominant woody species of this zone are bal-

for Pb, and possibly for other heavy metals, is clearly


sam fir Abies balsamea (L.) Mill, paper birch Betula

high for non-urban environments. The upper zones of


papyrifera Marsh and B. papyrifera var. cordifolia (Regel)

New England mountains may be regarded as interception Fern. and mountain ash Pyrus americana (Marsch) DC.

points in a high velocity air stream laden with moisture


Minor species include Carex sp., Clintonia borealis

OIKOS 26:3 (1975) 265

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Wet digestion was tested in comparison with the
(Ait.) Raf., Cornus canadensis L., Dryopteris spinulosa

dry-ash process used here. Contrary to other reported


(0. F. Muell.) Watt, Lycopodium annotinum L., L. luci-

results, wet digestion with concentrated HNO3 produced


dulum Michx., Oxalis montana Raf., Ribes glandulosum

consistently, and often significantly, lower yields of Pb


Grauer and Solidago macrophylla Pursch.

and Zn than did the dry-ash process.


In addition, five alpine tundra sites were sampled

ranging in elevation from 1436 to 1460 m. All of these

alpine sites were on northern, western, or southern

3. Results

exposures and were dominated by a variety of tundra

Averages and standard errors for some soil characterist-


species. Common dominants included Carex bigelowii

ics are listed in Tab. 1. The numbers of replicates for


Torr., Juncus trifidus L., Potentilla tridentata Ait.,

each layer vary depending on rock obstruction encount-


Vaccinium vitis-idaea L. and Cetraria islandica (L.)Ach.

ered for all or part of each sampling point. A full com-


A 20 m transect line was laid out in each site. Samples

plement of ten samples is available for four of the sites


were taken within each 2 m interval along this line, the

due to an error in sampling procedure as described in


exact location of each point being determined with a

the methods section. This differential in sample size does


random number table. At each point, two round cores

not affect concentration values except to affect the


5.6 cm in diameter were collected with a stainless steel

variation around the estimates. It tends to bias total


coring tube and bulked as a single sample. The depth

contents upwards at these sites, however, as averages are


of each core was limited by obstructing rock or the lower

always calculated by dividing by 10 to account for rock


limit of organic layers. If no core could be taken at any

cover. This bias will be discussed later.


point due to obstructing rock, the sample was counted

It should be noted that averages for the hardwoods


as zero for purposes of accounting for rockiness in

and spruce-fir sites shown in Fig. 2 are derived from


estimating average organic horizon weight for the site.

the sums of averages of each layer from all 10 sample


Four exceptions to this procedure were the sites at 1364,

points. Thus, for these ecosystem types, standard errors


1390, 1387 and 1396 m at which we continued to de-

represent intrasite variation. Averages for the fir forest


termine random sample points until 10 samples had been

krummholz and tundra sites, on the other hand, are


collected.

derived from sums of averages of layers from the various


Core samples from the forested sites were divided

into L, F, or H layers (L = 01, F + H = 02 horizons, sites. Thus, standard errors for these types represent

intersite variation.
Anon. 1960) in the field and transported separately in

plastic bags. Tundra soil cores were transported entire The emphasis in this study was on high elevation

ecosystems where sampling was widely distributed and


and divided into two layers, L and F + H in the

laboratory. The material was always handled with in aggregate, generously replicated. For example, aver-

disposable plastic gloves. ages in Fig. 2 for alpine tundra are derived from 42 com-

Samples were removed from the plastic bags and dried posite soil cores. Sampling in the northern hardwoods

for 24 h in a forced-draft oven at 105'C. The samples spruce-fir sites is of much lower intensity and values

were weighed to 0.1 g precision and then ground in a presented for these ecosystem types are primarily for

porcelain mortar and pestle. From these ground samples, tentative comparative purposes.

representative subsamples of approximately 1 g were

3.1. Heavy metal concentrations

placed in fused quartz crucibles, dried for 4-6 h at 80'C

in a gravity draft oven, cooled in desiccators, and Pb concentrations vary exactly as predicted by our first

weighed to 0.1 mg precision. These subsamples were then hypothesis in all three layers (Fig. 1A, B, C). Concen-

ashed in a muffle furnace at 550'C for 6 h. After cooling trations increase with elevation up through fir forest

in a desiccator, the resulting ash was weighed to deter- sites, and then decline in krummholz and alpine tundra

mine organic content by difference. where the vegetation is more streamlined.

The ash was dissolved by adding 5 ml distilled water The relationship of Zn concentrations with elevation

and 5 ml concentrated HNO3 to the crucible and heating does not fit this hypothesis as well, due to smaller

on a sand bath. After evaporating to 5 ml, the resulting difference in concentrations between sites, and to the

solution was filtered through acid-washed Whatman 541 especially high concentrations in the lowest site repre-

filter paper into 100 ml volumetric flasks. The flasks senting the northern hardwoods zone. We attribute this

were brought up to volume with 2.5 N HNO3 and the latter phenomenon to the high levels of Zn in the leaves of

solution analyzed on a Perkin Elmer model 303 atomic our deciduous trees (Likens and Bormann 1970, Gosz

absorption spectrophotometer for Pb, Zn, and initially, et al. 1972). Zn is an essential micronutrient and is con-

Cd. Cd analyses suffered from interference difficulties centrated in angiosperms at about six times the levels

and were eventually discontinued. found in gymnosperms (Bowen 1966). If this is true in

All glassware was hot-acid washed, rinsed with tap our area, then specific biological influences are in this

water five times, and with glass-distilled water five case swamping out the pattern of variation with eleva-

times. Two empty crucibles were run as blanks through tion expected on the basis of exposure and aerosol cap-

the entire process for each set of samples. ture efficiency. This angiosperm recycling effect is not

266 OIKOS 26:3 (1975)

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LITTER LAYER Fig. 1. Concentrations of Pb and Zn

for the organic layers of five ecosystem

340- LEADZINCT types of Mt Moosilauke, New Hamp-

A40- ,LX LEAD O ZINC shire. Bars around points are + one

320- 1 standard error of the mean.

T~ - northern hardwoods

300- j 0 - spruce-fir

280. o-f ir

- ! o - krummholz

I 260 0

o 240

- 220-

B 200

z 1808

j 60 -160

140 -140

W 120 - 120

Cl 140- O100

I 80 80

60 60

40 40

20 20

o' 0

FERMENTATION LAYER

280-

260 B LEAD ZINC

j240 0-northern hardwoods

I 0 -spruce -fir

~2220- o-fir

3: 200. 0 - krummholz I-I200

A_&-alpine tundra

tr 180- -f~180

z 160 160

r 140 { 140

0i 1200 120

a80- 80

60 60

a. 40? 40

20 20

o0 _j~~~~~~~~~~1

HUMUS LAYER

~200 C LEAD ZINC E200

180 180

>. 160. 160

O140- 140

Z 120 , 120

-j 100~ too

80 80

w 60 60

40 st440

20 r 20

a.~ 600 800 0'00 120'0 1400 600 800 1000 1200 1400 0

ELEVATION, METERS

18 OIKOS 26:3 (1975) 267

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10000 Fig. 2. Total weight of organic matter,
Pb and Zn per m2 for five ecosystem

sooo- A 829S* KRUMMHOLZ ALPINE types on Mt Moosilauke, New Hamp-


9000 /4 647 82350 TUNDRA shire. Measures of variation are stand-
2508 806528 ard errors of the means.Standard errors

8000 'for the northern hardwoods and spruce-

<J 1252lx 'x,. // fir sites represent intrasite variation, er-

2 7000- > ,. 223 t. ,2g8/7/o// rors for fir, krummholz and tundra
2 W j~sL represent intersite variation.

o ~~~~~~~~~~~SPRUCE-FIR

5546? 2l901 '19,30?-

6000 1291 2 9 0

Iii NORTHERN

HARDWOODS

5000- 4180? 321

1256 CA

240 I 2(9

z ~~~~~~~3 594R1 478

3000-

2000 3?3

1000 H cr~~ ~~~~~~~~~~~~~~ 0.99t O.2 2 6 +.2

FIR

1.5- 1.45 ?0.22

B ~~~~~~~~~~~~~ALPINE

L KRUMMHOLZ TUNO'RA

0.99? 002 1.002?0.20

CQ .0 0

0U.R L4' 0.0?

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1.15? 0.44

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NORTHERN 0.90 ?0.090.3,

1.0 HARDWOODS KRUMMHOLZ k

N- HUBBARD 0.78 ?0.20

2 BROOK .3 0.1 6?,

0.64 ~~~~~~~~~~~~~~~~0.03

CD . 0.14?..,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.

0 ~~~~~~~~~~ ~ ~~~~~~~~~SPRUCE-FIR 0.29?"

z 0.44?t0.1 1 ~ 0.04 020 .7

N 0.5 F ?0

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.04 #~~~~~~~~~~~~0 /2

500 600 700 800 900 1000 1100 1 200 1300 1400 1500

ELEVATION, METERS

270 OIKOS 26:3 (1975)

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obvious for the alpine tundra sites, possibly due to Ash-free dry weights are considerably less variable

lesser quantity of plant turnover in the tundra. and easier to interpret (Fig. 2A). On this basis the mass

Contrary to our second hypothesis, Pb and Zn con- of potential heavy metal-binding organic matter falls

centrations do not increase with depth. In fact, the into two classes: relatively lower weights for northern

reverse is true for Pb. There is a nearly consistent trend hardwoods and spruce-fir sites, and relatively higher

for concentrations to decrease with depth - on a dry- weights for fir forest, fir krummholz, and alpine tundra

weight or even an ash-free dry-weight basis (Tab. 1). sites.

Ash-free dry weights are included to permit assessment It is in this context that the sampling bias becomes

of concentrations on an organic-matter basis. Such most serious. Three of the four sites biased are in fir

values are particularly useful for comparing soils, such forest and one is in krummholz (Tab. 1). If total weights

as tundra soils, undergoing different amounts of mixing in the three affected fir forest sites are reduced by 20%,

with inorganic horizons. the total for this site type will decrease to approximately

For Zn, this trend of decreasing concentration with 7560 g m-2. A similar reduction in the krummholz will

depth is less marked on a dry weight basis, and is decrease the average for that type to approximately 7580

completely obscured on an ash-free basis. Thus it would g m-2. With these revised figures, organic matter appears

seem that different processes are controlling vertical to increase slightly, but not significantly, with elevation

distribution of these two metals. along this particular portion of the gradient.

We suggest that the explanation for the distribution Total Pb contents in Fig. 2B are a subdued reflection

of Pb is partly historical. If we assume that little Pb of concentration variations with elevation. Totals in-

is translocated to shoots in our systems and thence re- crease with elevation up to the fir forest, then decline in

cycled, and that it is strongly bound to organic matter krummholz as initially predicted. The sampling bias

(Anon. 1972), then under steady state deposition and problem can be treated as described for organic weights.

litter decomposition conditions, Pb should concentrate Doing this, the average for fir forest sites is 1.24 g m-2

in the lower, more finely divided humus layer. Such and for krummholz sites is 1.15 g m-2. This treatment

behavior is indicated for spruce needles in Sweden by leaves the general rank order with elevation unchanged.

Nilsson (1972). If the first two assumptions are true, The relationship of Zn content with elevation bears

then perhaps deviation from this expected distribution little relationship to either that of Pb or our hypothesis

is due to the fact that deposition rates are not historically for elevational distribution (Fig. 2C). While the general

constant. In fact, the widespread dissemination of Pb pattern of exposure and deposition may hold for the

aerosols is fairly recent and its rate is accelerating (Muro- coniferous forest and krummholz sites, other factors

zumi et al. 1969). With slow organic matter turnover are overriding for the northern hardwoods and alpine

under these cold environmental conditions, the normal tundra sites. The same bias corrections applied for Pb

redistribution process may not have kept pace with results in Zn totals of 0.78 and 0.71 g m-2 in the fir

accelerating deposition rates. Implicit in this possibility forest and kruinmholz sites respectively.

is the likelihood that Pb concentrations in these soils Comparable data from a northern hardwoods type at

are increasing.
nearby Hubbard Brook Experimental Forest have

At the present time, we can only offer some possible


been contributed by Dr James Gosz, University of New

factors behind the irregular and ambiguous distribution Mexico (Fig. 2C). His total of 0.77 g m-2 is nearly half of

pattern for Zn concentrations. It may bind less strongly


our value and inasmuch as his datum is an average of

than does Pb to organic matter at these levels of


many more samples than ours, it may be more typical.

acidity, the history of deposition may be longer, or show


Several factors might contribute to such a disparity

less increase in rate, and recycling of native and anthro-


between presumably equivalent situations.

pogenic Zn by roots may be the dominating distribut-


We considered that differences in rock type might

ional process.
cause differences in recyclable Zn but present evidence

does not support this (Dethier and Schlesinger 1973). A

3.2. Heavy metal contents

second possible factor is that the Moosilauke site is

Heavy metal contents in the organic horizons are the


situated at a higher and generally windward, west-facing

products of concentrations and organic matter mass.


slope compared with the less exposed position of the

Dry weights of these layers vary widely and unsystemat-


Hubbard Brook site. While this could be a contributory

ically, depending on the depth of organic horizons, and


factor, it is doubtful that it is sufficient to explain a

on the amount of incorporated inorganic matter (Tab. 1).


difference of this magnitude. We believe that the princi-

The range of variation in site types increases to a large


pal cause behind this difference lies in the tree composi-

extent from fir forests to alpine tundra. There is a


tion of our site which is dominated by yellow birch. The

striking difference in the weight of the L-layer of the


Hubbard Brook site bears a more typical forest composi-

northern hardwoods site compared with coniferous


tion of sugar maple Acer saccharum Marsh., beech Fagus

sites, a manifestation, partially, of differences in climate,


grandifolia Ehrh. and yellow birch Betula lutea Michx.

but more importantly, in the decomposability of the two


(Bormann et al. 1970). Gosz et al. (1972) showed that the

types of fresh litter.


senescent leaves of yellow birch have about ten times

OIKOS 26:3 (1975) 271

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metal present per unit area or volume of soil. Highly
two co-dominants. This may be an illustration of how a

species can influence that spatial distribution of micro- organic soils such as ours have a high affinity for heavy

nutrient levels in soils in general. It also suggests the metals but a low bulk density. A low density soil will

yellow birch may be actively keeping Cd, an element obviously have a much lower metal content per unit

geochemically similar to Zn, at higher levels of avail- area or volume than a high density soil of equal metal

ability than would be found in other parts of the forest. concentration. To facilitate future comparisons with

other data we have provided percentage organic content

and total Pb and Zn contents on a unit area basis (Tab.

4. Discussion

1). For our present purposes, however, we can only

4.1. Weights of organic layers

make comparisons with extant concentration data that

The amount of organic matter accumulated in surface are difficult to evaluate in terms of total metal contents.

layers of soils is dependent on the rate of input from To partially compensate for this problem, data from the

aerial and subterranean plant organs balanced against literature have been divided into inorganic soils, and

the decomposition rate, and on the time since the last organic layers, or organic soils (Tab. 2).

disturbance. In all the sites examined in this study, there In addition to soil data, Tab. 2 also contains bedrock

have been no known disturbances of the forest floors by data for Mt Moosilauke. Since we are attributing our

man or other agencies so that a steady state balance is elevated concentrations to deposition of airborne pollu-

probably approximated in terms of forest floor mass. tants, it is necessary to check the metal contents of the

Inasmuch as plant biomass, and presumably, primary local bedrock. The values for the predominating bedrock

production decrease up the elevational gradient repres- of the study area (a metamorphosed shale) are somewhat

ented in Tab. 1 and Fig. 2A, it seems likely that detritus high for shales with respect to Pb but normal for Zn

inputs also decrease proportionately. Similarly, we (Bowen 1966).

would expect that decomposition rates would also Compared with the data in Tab. 2, Pb levels in the

decrease. It is virtually impossible to determine from the


spruce-fir, fir forest, and fir krummholz are higher than

dry weight data (Tab. 1) whether the net change in


would be expected for a remote area. Fir forest and

balance represented by weights of organic layers varies in


krummholz in particular have levels exceeded only by the

any direction because degrees of mixing with mineral


Pennsylvania data which probably represent regional

matter profoundly affect the values. For a more accurate


contamination levels for the area that are abnormally

view, the ash-free weights in Fig. 2A should be used. high by any other standards.

These may be amended further to correct for sampling


The comparisons of Zn data in Tab. 2 lead to the

bias as described in the previous section. The amended


conclusion that no outstanding enrichment is represented

sequence from lowest to highest sites is 4180, 5550,


by the Mt Moosilauke samples. It should be pointed

7560, 7580, and 8060 g m-2. This series suggests that


out, however, that all of the examples used in Tab. 2

accumulated organic matter increases by about 60 g


are from areas receiving general regional pollution and

m-2 for every 100 m increase in elevation. Evidently


our values may be significantly higher than those of

the climatic or biological factors associated with eleva-


areas upwind of human activity - areas unrepresented in

tional change depress decomposition rates more than the table.

accession rates.
Our higher Pb concentrations exceed those reported

Comparisons of total dry weight values with ash-free


for a contaminated salt marsh (Siccama and Porter 1972)

data from this sequence of sites is illustrative of the


or the vicinity of a fossil fuel power plant (Klein and

caution required in comparing data with those in the


Russell 1973). They are generally much lower than near-

literature. With this caveat, we note that our values


smelter values (Motto et al. 1970, Little and Martin

are considerably higher than those presented by Rodin


1972, Ruhling and Tyler 1973, Buchauer 1973), but

and Bazilevich (1967, Tab. 53), as forest floors for


comparable with some areas contaminated by alluvium

representative vegetation types. Our total for organic


derived from mine tailings or other mining-related

matter correspond quite well, however, with the 6200 g


contamination (Alloway and Davies 1971, Davies 1971,

m-2 for a Minnesota oak forest sampled and analyzed


Davies and Lewin 1974). Our higher levels for Pb and

in exactly the same manner as in this study by Reiners


Zn fall within the range of levels given for roadside

and Reiners (1970).


soils as reported by Lagerwerf and Specht (1970),

Lagerwerf (1971), Smith (1972), Gish et al. (1973), Ward

et al. (1974) and Hemphill et al. (1974).

4.2. Heavy metal concentrations

These concentrations of Pb or Zn are below those

The third initial hypothesis of this study was that Zn


known to have direct effects on the ecosystems such as

and Pb concentrations would be high for a remote area,


toxic effects on higher plants (Buchauer 1973), Davis

possibly comparable with concentrations in directly


and Barnes (1973), on decomposition (Bhuiya and Corn-

contaminated environments. In making comparisons


field 1972, Ruhling and Tyler 1973), on nitrification

between our data and data in the literature, it is clear that


(Tyler et al. 1974) or on cation replacement (Ruhling

concentrations do not adequately express the amount of


and Tyler 1973). However, if these ecosystems are not in

272 OIKOS 26:3 (1975)

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Tab. 2. Comparisons of lead and zinc concentrations measured in this study with other data in the literature for areas

presumed to not represent local contamination.

Area Pb (ppm) Zn (ppm) Source

This study

minimum - all sites, layers ............... 11 19

maximum - all sites, layers ...... ........ 336 169

weighted average, all layers

hardwoods ............. ............ 35 100

spruce-fir ........................... 79 53

fir forest ............................ 145 90

fir krummholz ........... ............ 120 73

alpine tundra ........................ 38 29

Littleton formation

Mt Moosilauke bedrock ....... ......... 46 90 Dethier and Schlesinger 1973

Mean for soils

world average (presumably inorganic) ..... 10 50 Bowen 1966

Inorganic soils

Connecticut, U.S.A ..................... 10-15 - Smith 1972

New Zealand .......................... 15 - Ward et al. 1974

Missouri, U.S.A ........................ 17 - Hemphill et al. 1974

Michigan, U.S.A 26 Klein and Russell 1973

West England ......................... 42 64 Davies 1971

Wales ................................ 42 129 Davies and Lewins 1974

Wales ................................ 45 80 Alloway and Davies 1971

Organic layers

Salt marsh, Conn ....................... 38 - Siccama and Porter 1972

Heath, Sweden (above ground litter) ...... 66, 80 53, 68 Tyler et al. 1973

Heath, Sweden (below ground litter) ...... 40, 45 72, 80

Spruce needle litter, Sweden ...... ....... 12, 62 - Rifhling and Tyler 1973

Spruce needle litter, Sweden Nilsson 1972

least decomposed ........ ............ 27, 34, 78 60, 65, 86

intermediate ............ ............ 49, 61, 102 72, 90, 91

most decomposed .................... 66, 105 78, 126

02, Pennsylvania ....................... 240-355 311-346 Buchauer 1973

steady states as the distribution data for Pb suggest, then England mountains are exposed to high winds, precipita-

accumulation may continue and such levels might even- tion, and cloudiness. The combination of high bulk

tually be reached - perhaps sooner for Cd than for Pb precipitation plus horizontal interception at high eleva-

or Zn (John 1973).
tions contributes to higher rates of aerosol deposition

We would not expect much biological amplification of than are likely to be occurring elsewhere in the region.

Pb in grazing food chains of forests as limited plant


Thus montane ecosystems are exposed to higher fluxes

uptake seems to disrupt this process. The alpine tundra


of all air-borne pollutants than are other similarly re-

may represent a different situation than the forests, mote environments in this region. In the case of hydro-

however, as shrub and graminoid plant surfaces that gen, sulfate and nitrate ions associated with acid rain

intercept aerosols are eaten directly by small mammals (Likens and Bormann 1974), this means that the flux

and insects. Hirao and Patterson (1974) have demon-


of these ions at high elevations is more than double the

strated such a phenomenon for a Sierran meadow eco-


flux at low elevations. Thus, any effects resulting from

system and we have detected Pb as high as 70 ppm on the


acid rain could be potentially more serious at high eleva-

leaves of one of the tundra plants of Mt Moosilauke. Am-


tions. Similarly, these high elevation ecosystems are

plification through detritus food chains should be exam-


exposed to high deposition rates of heavy metals, pollu-

ined, as fungal mycelia have been shown to concentrate


tants which appear to be firmly retained in the soils.

Pb by two orders of magnitude over levels in their culture


Likens and Bormann (1974) have suggested that

medium (Hartman 1974). Based on the distribution pat- technological change, sometimes instituted in the interest

terns of Zn, movement of Cd into both types of food


of environmental quality, has converted local air pollu-

chains could be significant.


tion problems to regional problems. If this is a general

trend, then it is likely that the hitherto protected high

4.3. Mountain ecosystems as environmental sensors

elevation environments will become the most exposed

The significance of concentrations reported here is


sites to certain pollutants. For this reason such ecosys-

probably not in their indication of potential effects,


tems serve a special purpose as exposure monitors. In a

but in their indication of ecological processes. New


sense, they are amplifying warning systems. They are

OIKOS 26:3 (1975) 273

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HAGEN, A. and LANGELAND, A. 1973. Polluted snow in
the first environments that might build up detectable

southern Norway and the effect of meltwater on fresh-

levels of certain new, exotic pollutants; they provide

water and aquatic organisms. - Environ. Pollut. 5:

particularly sensitive base line sites. Further, they serve

45-58.

as testing sites for effects of such elevated exposures. HARTMAN, L. M. 1974. Fungal flora of the soil as condi-

tioned by varying concentrations of heavy metals. -


This use is further enhanced by the fact that such eco-

Amer. J. Bot. 61: Suppl. 23 (abstract).

systems are usually simple and relatively free from

HEMPHILL, D. D., MARIENFELD, C. J., REDDY, R. S. and

other disturbances. We suggest the subalpine and alpine

PIERCE, J. 0. 1974. Roadside lead contamination in the

ecosystems should be examined as particularly useful


Missouri lead belt. - Arch. Environ. Health 28: 190-

194.
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HIRAO, Y. and PATTERSON, C. C. 1974. Lead aerosol

deposition into the biosphere and the ecological effects

pollution in the high Sierra overrides natural mechan-

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JOHN, M. K. 1973. Cadmium uptake by eight food crops

Acknowledgement - This study was supported by the Ri-

as influenced by various soil levels of cadmium. - En-

chard King Mellon Foundation, Pittsburgh, Pennsylvania.

viron. Pollut. 4: 7-15.

- , CHUAH, H. H. and VANHAERHOVEN, C. J. 1972. Cad-

mium contamination of soil and its uptake by oats. -


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