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OIKOS 26: 264-275. Copenhagen 1975
W. A. REINERS
R. H. MARKS
P. M. VITOUSEK
deposition rates. This study describes the concentrations and total quantities of
Pb concentrations are high for a remote area, and comparable with many heavily-
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.
BJIHSHHS 3arps3HHTeJeil.
? OIKOS
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and aerosols, including heavy metals. If montane eco-
1. Introduction
The long range transport of heavy metals from centers of heavy metals, we might then hypothesize that heavy
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,
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
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
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
for deposition.
2. Methods
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
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-
bulk precipitation does not take into account aerosols to approximately 760 m (Bormann et al. 1970). The next
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
Mt Moosilauke. Results of that study suggested that Pb to the density of interceptive surfaces of the evergreen
New England mountains may be regarded as interception Fern. and mountain ash Pyrus americana (Marsch) DC.
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Wet digestion was tested in comparison with the
(Ait.) Raf., Cornus canadensis L., Dryopteris spinulosa
3. Results
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
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.
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
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
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LITTER LAYER Fig. 1. Concentrations of Pb and Zn
A40- ,LX LEAD O ZINC shire. Bars around points are + one
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-
I 0 -spruce -fir
~2220- o-fir
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
180 180
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
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0 0~0~~Ro &0
V666 6666 ~~~~~~6 666i 666 ; C (666; 6 666 < 6666; 5-4C 6 666 C 6666
o0
o OS
- ~~~~~~~~~~~~
ti..4 W) '0 I(2 0 t ~0O t \0 e 'f2t -00' 0 NU N~ 002 I rat N WI or-'r . t'- C'r en r o
o c6 0 - -- c c - i i - e4i- - - - -
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bol0' OD CN en C0O 42~ -r-- 0-000000 000 0000o I 7"~ 00 aD0 00 00 q O0000 enfnWr-o e
Cl C
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to ~ o
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N N - N N 00a e 0r 0 t- a 0-- 00
- ~ -- o -~ - -- ' t I O I - t 0 N - I c
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o co q Q t0o or 0 ooo
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C, (0 00 0 C- ~t O, tn C1 tn X O. r- t NC) 00 tn 00 _<5c
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t0 -, 00r t tn - r- ??o o cq N c1 en (= n dc en t1 r- (= m d-
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C " _ N It _ _ _
- 1o??m td e Fn 2 i
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10000 Fig. 2. Total weight of organic matter,
Pb and Zn per m2 for five ecosystem
2 7000- > ,. 223 t. ,2g8/7/o// rors for fir, krummholz and tundra
2 W j~sL represent intersite variation.
o ~~~~~~~~~~~SPRUCE-FIR
6000 1291 2 9 0
Iii NORTHERN
HARDWOODS
1256 CA
240 I 2(9
3000-
2000 3?3
FIR
B ~~~~~~~~~~~~~ALPINE
L KRUMMHOLZ TUNO'RA
CQ .0 0
o ~~~~~~~OTENSPRUCE-FIR
0.50 ?0.17
-j
H~~~~~~~~~~~~~~~~
NO007HERS
HAR060005
1.49 tO0?3
0.0 ALPINE
1.15? 0.44
FIR .
0.64 ~~~~~~~~~~~~~~~~0.03
CD . 0.14?..,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.
N 0.5 F ?0
- 0~~~~~~~~~~.12?004 00
.04 #~~~~~~~~~~~~0 /2
500 600 700 800 900 1000 1100 1 200 1300 1400 1500
ELEVATION, METERS
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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
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
factors behind the irregular and ambiguous distribution Mexico (Fig. 2C). His total of 0.77 g m-2 is nearly half of
ional process.
cause differences in recyclable Zn but present evidence
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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
4. Discussion
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
would expect that decomposition rates would also Compared with the data in Tab. 2, Pb levels in the
view, the ash-free weights in Fig. 2A should be used. high by any other standards.
accession rates.
Our higher Pb concentrations exceed those reported
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Tab. 2. Comparisons of lead and zinc concentrations measured in this study with other data in the literature for areas
This study
spruce-fir ........................... 79 53
Littleton formation
Inorganic soils
Organic layers
Heath, Sweden (above ground litter) ...... 66, 80 53, 68 Tyler et al. 1973
Spruce needle litter, Sweden ...... ....... 12, 62 - Rifhling and Tyler 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.
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
medium (Hartman 1974). Based on the distribution pat- technological change, sometimes instituted in the interest
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HAGEN, A. and LANGELAND, A. 1973. Polluted snow in
the first environments that might build up detectable
45-58.
as testing sites for effects of such elevated exposures. HARTMAN, L. M. 1974. Fungal flora of the soil as condi-
194.
systems for understanding the pattern of pollutant
of these pollutants.
184: 989-992.
Geoderma 5: 197-207.
of New England. - Science 177: 514-516.
No. 18.
Academic Press, N.Y.
577-580.
147: 502-503.
1179.
22: 366-372.
769-784.
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