Lichens, a unique forage resource threatened by air pollution

Paper presented at T h e F i r s t A r c t i c U n g u l a t e C o n f e r e n c e ,
N u u k , G r e e n l a n d , 3 - 8 . September, 1991.
Lichens, a unique forage resource threatened by air pollution
David R. Klein a n d Tatyana J. Vlasova
1
1
2
U . S . F i s h a n d W i l d l i f e Service, A l a s k a C o o p e r a t i v e W i l d l i f e Research U n i t , U n i v e r s i t y of A l a s k a F a i r b a n k s ,
F a i r b a n k s , A K . 99775, U . S . A .
2
E x t r e m e N o r t h Institute f o r A g r i c u l t u r a l Research, K o m s o m o l s k a y a Str. 2, 663302 N o r i l s k , Russia.
Abstract: L i c h e n s are the p r i m a r y w i n t e r forage f o r most m a i n l a n d c a r i b o u a n d reindeer herds i n N o r t h A m e rica a n d f o r the m a j o r i t y o f domestic a n d w i l d reindeer i n Siberia and n o r t h e r n E u r o p e , c o l l e c t i v e l y t o t a l i n g
i n excess of 5 m i l l i o n animals. L i c h e n s represent a unique forage resource t h r o u g h o u t m u c h o f the c i r c u m p o lar N o r t h that cannot effectively be replaced b y vascular plants. L i c h e n s are p a r t i c u l a r l y sensitive t o the effects of air p o l l u t i o n . T h e increased pace o f e x p l o i t a t i o n a n d processing o f minerals and p e t r o l e u m
resources
t h r o u g h o u t the c i r c u m p o l a r N o r t h , w i t h associated i n t r o d u c t i o n of p o l l u t i o n products i n t o the atmosphere
has already resulted i n losses of lichens a n d their reduced p r o d u c t i v i t y i n extensive areas adjacent t o large metallurgical complexes i n the T a i m y r of Siberia, o n the K o l a P e n i n s u l a , a n d i n adjacent parts of F i n l a n d . Losses
of terricolous lichens i n the T a i m y r f r o m p o l l u t i o n generated b y the N o r i l s k metallurgical c o m p l e x have been
nearly complete w i t h i n a 300,000 h a area closest t o the p o l l u t i o n source a n d damage and reduced g r o w t h extends over an area i n excess of 600,000 h a . T h e A r c t i c also is a sink f o r atmospheric p o l l u t i o n generated i n
the h e a v i l y i n d u s t r i a l i z e d n o r t h temperate regions of the w o r l d . Assessment of the effects o n lichens of this
global scale increase i n air p o l l u t i o n is d i f f i c u l t because of the lack of representative
Keywords: lichens,
controls.
p o l l u t i o n , sulfur d i o x i d e , c a r i b o u , reindeer, Soviet U n i o n , A r c t i c pastyres,
winter
Rangifer, 12 (1): 21-27
Introduction
is available
Lichens are a p r i m a r y w i n t e r f o o d f o r over 5
though
lichens
million
pounds
that
c a r i b o u and reindeer (both Rangifer
ta-
as w i n t e r forage ( K l e i n 1982). A l act
as
unique
chemical
most
of
reindeer are w e l l adapted to use m a n y
A m e r i c a , the
tundra
taiga
Eurasia,
(Andreev
and
their
adjacent
Klein
1982). L i c h e n - d o m i n a t e d plant c o m m u n i -
ties o c c u p y vast regions
areas
of
of the
northern
1977,
por-
(Rundel
acidic
defenses
randus) that o c c u p y the n o r t h e r n boreal forest
North
herbivores
contain
1978),
comagainst
caribou
and
arboreal
and terricolous l i c h e n species as forage. T h e p r i m a r y value of lichens for reindeer and c a r i b o u
is
in
their
high
content
of
available
energy
tions of N o r t h A m e r i c a and Eurasia ( A n d r e e v
w h i c h is the major dietary requirement of these
1977,
northern
K e r s h a w 1977)
(Figure
1). L i c h e n s are a
ungulates
i n winter. Thus
the
abun-
unique forage that, u n l i k e vascular plants, accu-
dance and availability of lichens at h i g h n o r t h -
mulate l i v i n g plant tissue over several years that
ern latitudes enables successful o v e r w i n t e r i n g of
Rangifer, 12 (1), 1992
21
F i g . 1.
large herds of migratory caribou and domestic
reindeer. Suitable areas w i t h high lichen biom¬
ass for w i n t e r grazing b y reindeer or caribou
occupy 3 0 - 4 0 % of the land area of the n o r t h ern regiaons of N o r t h A m e r i c a and Eurasia
(Andreev 1977, K e r s h a w 1977).
T h e uniqueness of lichens, i n contrast to vascular forage plants, derives f r o m their structure,
physiology, and ecology as a f u n c t i o n of the
symbiotic u n i o n of fungi and blue green algae.
T h e structure of lichens is f o r m e d b y the fungal
component, w h i c h provides an interstitial environment favorable for blue green algae. W h e n
sufficient moisture and light are available, the
22
algae are able to photosynthesize and draw o n
nutrients f r o m the atmosphere, i n c l u d i n g trace
nutrients, that become sequestered i n the fungal
tissues. T h e photosynthates that are produced
are then available for g r o w t h of b o t h the algal
and fungal components as a s y m b i o t i c "organ i s m . " Lichens have n o roots and hence derive
v i r t u a l l y all of their requirements f o r g r o w t h
f r o m the atmosphere; their porous structure
also results i n their capture of particulate material falling f r o m the atmosphere ( H a w k s w o r t h
1971). Because of this characteristic lichens also
accumulate pollutants f r o m the atmosphere.
Rangifer, 12 (1), 1992
F i g . 1. M a j o r p o i n t sources of atmospheric p o l l u t i o n i n and adjacent to the boreal and taiga forests (Boreal and taiga forest zone s h o w n as crosshatching f r o m A h t i et al. 1968; p o l l u t i o n sources f r o m
N o r w e g i a n Institute f o r A i r Research 1984 and C a n a d a D e p a r t m e n t E n e r g y , M i n e s and Resources 1988).
N o r t h America
1. O i l field c o m p l e x , P r u d h o e B a y , A l a s k a
2. P e t r o c h e m i c a l industries* K e n a i , A l a s k a
3. C o p p e r - z i n c smelting, F l i n F l o n ,
4.
5.
6.
7.
Saskatchewan
N i c k e l smelting, T h o m p s o n , M a n i t o b a
I r o n sintering plant, W a w a , O n t a r i o
N i c k e l - c o p p e r smelting, S u d b u r y , O n t a r i o
Z i n c - c o p p e r - c a d m i u m smelting, T i m m i n s , O n t a r i o
8. I r o n sintering plant, K i r k l a n d L a k e , O n t a r i o
9. C o p p e r smelting, N o r a n d a , Q u e b e c
10. I r o n - t i t a n i u m agglomerate plant, P o r t C a r t i e r , Q u e b e c
11. C o p p e r smelting, M u r d o c k v i l l e , Q u e b e c
12. Lead-silver-copper smelting, Belledune, N e w B r u n s w i c k
13. I r o n sintering plant, L a b r a d o r C i t y , L a b r a d o r
Eurasia
1. I r o n smelting, Sydvaranger, N o r w a y
2. C o p p e r - n i c k e l smelting, N i k e l - Z a p o l y a r n i , K o l a Peninsula, U . S . S . R .
3. C o p p e r - n i c k e l smelting, M o n c h e g o r s k , K o l a Peninsula, U . S . S . R .
4. I r o n smelting, K o s t o m u k s h a ,
U.S.S.R.
5. C o a l - f i r e d p u l p and paper p r o d u c t i o n , Svetogorsk, K o l a P e n i n s u l a , U . S . S . R .
6. I r o n smelting and steel p r o d u c t i o n , Cherepovets,
U.S.S.R.
7. C o a l - f i r e d p o w e r and paper p r o d u c t i o n , A r k a n g e l s k , U . S . S . R .
8. C o a l - f i r e d p o w e r and paper p r o d u c t i o n , I k h t a ,
U.S.S.R.
9. C o a l - f i r e d p o w e r and paper p r o d u c t i o n , V o r k u t a , U . S . S . R .
10. C o p p e r - n i c k e l smelting, Sverdlovsk, U . S . S . R .
11. Steel p r o d u c t i o n , N i s h n i y T a g i l , U . S . S . R .
12. Steel p r o d u c t i o n , Serov, U . S . S . R .
13. Oil-gas field c o m p l e x , S a m o t l o r , U . S . S . R .
14. W o o d products and paper p r o d u c t i o n , Igarka,
U.S.S.R.
15. N i c k e l - c o p p e r smelting, N o r i l s k , U . S . S . R .
16. C o a l - f i r e d p o w e r and paper p r o d u c t i o n , Y a k u t s k ,
Response of lichens to air pollution
Erasmus D a r w i n , the grandfather of Charles
D a r w i n , observed i n the 19th century i n E n g land that lichens were sensitive to air p o l l u t i o n ,
and deterioration of the lichen flora around
L o n d o n d u r i n g the industrial r e v o l u t i o n was
noted b y several E n g l i s h botanists ( H a w k s w o r t h and M c M a n u s 1989). S i m i l a r l y , the l i chen flora of southern Sweden, w h i c h suffers
f r o m acid precipitation f r o m heavy industrial
activity i n n o r t h e r n E u r o p e , declined m a r k e d l y
i n species diversity and biomass over several decades ( A r u p et al. 1989). In Canada, atmospheric p o l l u t i o n f r o m the large n i c k e l smelting
complex i n , the Sudbury district of O n t a r i o has
had a l o n g history of negative effects o n vegetat i o n o n the region, most lichens were eliminated w i t h i n several kilometers d o w n w i n d of the
Rangifer, 12 (1), 1992
U.S.S.R.
p o l l u t i o n sources (LeBlanc and R a o 1966). Sulfur, i n v a r y i n g forms, is the p r i m a r y pollutant
responsible for the decline of the lichen flora
around Sudbury, and laboratory studies confirmed the sensitivity of lichens to sulfur dioxide
and its derivatives i n acid precipitation ( H a w k s w o r t h 1971, Pucket et al 1973).
P o i n t sources of p o l l u t i o n f r o m industrial development have increased m a r k e d l y i n n o r t h e r n
regions i n the latter half of this century (Figure
1). Little effort has been expanded o n evaluat i o n of the direct effects of atmospheric p o l l u t i o n o n lichens as a forage for reindeer and carib o u . In addition to the economic and socially
important role of reindeer husbandry i n the
past several decades i n the Soviet U n i o n , F i n land, Sweden, and N o r w a y , caribou and w i l d
reindeer have been equally important i n the
23
subsistence e c o n o m y of n o r t h e r n peoples i n the
Soviet U n i o n , G r e e n l a n d , Canada, and A l a s k a
( K l e i n 1989). O n l y d u r i n g the past t w o decades,
as international concern over the deterioration
of the w o r l d ' s environment m o u n t e d , have efforts been expanded at assessing losses of lichen
species diversity and biomass as forage for reindeer and caribou i n areas of heavy atmospheric
pollution.
A i r p o l l u t i o n f r o m smelting of n i c k e l and
copper at M o n c h e g o r s k and N i c k e l o n the K o l a
Peninsula, of U . S . S . R . resulted i n sulfur fallout
measured b y H e l l e et al. (1990) i n adjacent areas
of n o r t h e r n F i n l a n d of 1.0-1.2 g / m , and to 0.4
g / m i n western L a p l a n d . In spruce (Picea abies)
stands at 6 7 ° 5 0 ' N and 6 5 ° 4 4 ' N , where these
authors began studies of aboreal lichens i n
1976, lichen biomass had been reduced b y 50%
up to 1988 at the more southern site and b y
75% at northern site under sulfure fallout levels
of about 0.6 g / m .
2
2
2
Pollution from the Norilsk complex
In the T a i m y r region of n o r t h central Siberia,
p o l l u t i o n f r o m the huge metallurgical complex
of N o r i l s k has had serious consequences for l i chens i n an area that has been heavily used b y
b o t h domestic and w i l d reindeer (Vlasova et al.
1991). N o r i l s k , l y i n g i n the A r c t i c at nearly
7 0 ° N is a city of 260,000 people. F o u n d e d i n
the mid-1930 s i n association w i t h development
of the w o r l d class copper-nickel sulfide deposits
i n the region, N o r i l s k supplies about t w o thirds
of the total Soviet p r o d u c t i o n of n i c k e l and is
/
the largest n i c k e l producer i n the w o r l d . T h e
source of p o w e r for m i n i n g , smelting, the city
of N o r i l s k , and satellite m i n i n g centers was originally coal f r o m local mines. In the late I960' s
hydroelectric development of the K h a n t a y k a
and K u r e y k a rivers lessened dependence o n
coal. C o n s t r u c t i o n of natural gas pipelines to
the area during 1969-1973 further reduced the
need for coal, but also enabled construction of
a second smelter. T h e discontinued dependence
o n coal resulted i n some reduction of atmospheric p o l l u t i o n . G r o w t h i n the industrial c o m plex, however, increase i n sulfur content of the
ore processed, and i n t r o d u c t i o n of an autogenous fusion processing m e t h o d increased the
p r o p o r t i o n of sulfur component i n discharge
gasses f r o m 5 % i n 1981 to about 50% b y 1989
(Vlasova et al. 1991). T h e N o r i l s k complex n o w
accounts for r o u g h l y 4.5 m i l l i o n metric tons of
atmospheric p o l l u t i o n annually (Saunders 1990).
T h i s includes nitrogenous and anhydrous sulfur
components that readily convert to acids i n
contact w i t h rain or surface water and substantial amounts of heavy metals (Kovalev and F i l i p c h u c k 1990). Sulfur dioxide emissions alone
equal about 2.2 m i l l i o n metric tons per year, an
amount about equal to the total of all c o m b i ned Canadian output (Saunders 1990).
Studies of p o l l u t i o n effects f r o m the N o r i l s k
metallurgical c o m p l e x o n vegetation of the region were initiated i n 1976 b y Soviet scientists
f r o m the Far N o r t h Institutes for A g r i c u l t u r a l
Research i n N o r i l s k j o i n t l y w i t h specialists
f r o m the Forest Resources Section of the
Table 1. Effects of atmospheric p o l l u t i o n f r o m the N o r i l s k metallurgical c o m p l e x o n vitality, species c o m p o s i t i o n , percent g r o u n d cover, and biomass of lichens i n the region (data f r o m V l a s o v a et al. 1991; p o l l u t i o n zones are referenced in F i g . 2).
Vitality
of lichens
Degree of
Pollution
Number
of species
Percent
cover
Biomass
(100 kg/ha)
Maximum
Absent, or o n l y fragments remaining
0-4
<
<1
Severe
H e a v i l y damaged, m o r p h o l o g i c a l changes
of up to one half of the thalli
5-20
<20
1-5
Moderate
Stressed, less than average size, changes i n
color of thalli
21-34
>20-40
>5-10
Slight
N o r m a l , average size
>34
>40-70
>10-15
Relatively
clean
G o o d condition
>34
(up to 96)
>70-90
>15
24
Rangifer, 12 (1), 1992
20
-
40
i
60 80 k m
i
70°N
-68°N
DEGREE OF POLLUTION
M ax imum
Severe
Y77A Moderate
V///X Slight
Relatively Clean
F i g . 2. Degree of p o l l u t i o n i n zone r a d i a t i n g f r o m the N o r i l s k metallurgical c o m p l e x as reflected i n c o n d i t i o n
of lichens ( f r o m data i n T a b l e 1).
U n i o n Scientific Research Institutes i n M o s c o w
and B r y a n s k (Vlasova et al. 1991). In the zone
of fallout of atmospheric p o l l u t i o n d u r i n g summer, vegetation over an area of 565,000 ha show e d various levels of damage and deterioration
w h e n surveyed i n 1989 (Vlasova et al. 1991) (Figure 2). T h i s compares to 457,000 ha i n this category i n 1986 and 323,000 ha i n 1976, thus the
Rangifer, 12 (1), 1992
annual increase i n area affected changed f r o m
13,400 ha/yr f r o m 1976 t h r o u g h 1986 to 36,200
ha/yr i n the f o l l o w i n g 3 years. Forests, dominated b y Larix sibirica, and most lichens have
been k i l l e d i n an area of over 300,000 ha closest
to the p o l l u t i o n source and extending o b l i q u e l y
over 50 k m to the southeast (Figure 2) (Table
1).
25
undergo a l o n g sequence of recovery, including
T h e pattern of the response b y lichens to p o l the establishment of pioneering forms that are
l u t i o n is consistent w i t h distribution of the plulittle used b y caribou and reindeer, w h i c h ultime of summer p o l l u t i o n f r o m the N o r i l s k memately leads to a climax o r terminal lichen
tallurgical complex, w h i c h i n t u r n is the procomplex dominated b y species of most i m p o r duct of seasonal w i n d patterns. T o p o g r a p h i c
tance as forage (Kershaw 1977). T h i s entire sefeatures, however, also have an important influquence requires a m i n i m u m of 20 to 25 years
ence o n distribution of pollutants. T h e Putoraf o l l o w i n g fire (Andreev 1977), and m u c h longer
na M o u n t a i n s l y i n g east and southeast of N o if n o l i v i n g lichens remain i n the area o r vascurilsk are a partial barrier to penetration of airlar plant succession becomes stagnated i n a grass
borne pollutants although major river valleys
or sedge stage ( K l e i n 1982, 1987).
and associated large lake basins serve as chanWidespread deterioration and death of lichens
nels f o r movement of p o l l u t e d air into l o w e r
i n the T a i m y r as a consequence of p o l l u t i o n
l y i n g areas i n this m o u n t a i n range.
f r o m the N o r i l s k metallurgical c o m p l e x is a caT h e extensive area over w h i c h lichens have
tastrophic loss of a major forage resource, and
been eliminated o r their biomass greatly reduhas m a r k e d l y l i m i t e d p r o d u c t i v i t y of the region
ced as a consequence of p o l l u t i o n f r o m the N o for reindeer. Sensitivity of lichens to air p o l l u rilsk metallurgical complex has been an i m p o r tant w i n t e r i n g area f o r domestic and w^ild rein- t i o n has been w e l l demonstrated i n the T a i m y r
at the cost of loss of biological diversity, disrupdeer. W i l d reindeer f r o m the T a i m y r herd,
t i o n of ecosystems, and reduced p r o d u c t i v i t y of
n u m b e r i n g close to 600,000, winter i n the P u t o the land. Increased exploitation o f mineral rerana M o u n t a i n s mostly east of the p o l l u t e d
sources i n the c i r c u m p o l a r N o r t h and associaarea, however, tens of thousands of animals
ted industrial development, expansion of h u m a n
f r o m this herd extend their winter grazing into
populations, p o l l u t i o n f r o m p o i n t sources, and
slightly t o moderately p o l l u t e d areas. D o m e s t i c
global increases i n atmospheric p o l l u t i o n pose a
reindeer husbandry has been m u c h more severemajor threat to lichens and their use as f o o d f o r
l y affected b y the loss of lichens. W i n t e r grathe
w o r l d ' s p o p u l a t i o n of reindeer and caribou.
zing of domestic reindeer i n areas of m a x i m u m
and severe p o l l u t i o n was discontinued and herd
numbers have declined accordingly since the
References
mid-1970's. Because the polluted areas are exA h t i , T . , H a m e t - A h t i , L . and Jalas, J . 1968. Vegetapanding, the available lichen forage continues to
t i o n zones a n d their section i n n o r t h w e s t e r n E u r o decline t h r o u g h death of lichens and decreased
pa. - Annates Botanici Fennici 5: 169-211.
lichen g r o w t h rates. T h e future of reindeer hus- Andreev, V . N . 1977. Reindeer pastures i n the subbandry and of the w i l d reindeer p o p u l a t i o n i n
arctic territories o f the U . S . S . R . - I n : W . Krause
the T a i m y r is therefore seriously threatened.
(ed.) Handbook of vegetation science. Part III. W .
J u n k Publishers. T h e H a g u e , N e t h e r l a n d s , p p . 275¬
313.
Concluding remarks
W i l l the current political and economic reforms
i n progress i n the Soviet U n i o n lead t o realistic
efforts t o reduce p o l l u t i o n emissions f r o m the
N o r i l s k metallurgical complex?
Technology
exits i n the West to i m p r o v e efficiency of the
smelting process at N o r i l s k . Recently initiated
exchanges of representatives f r o m the N o r i l s k
complex and C a n a d a ' s Inco n i c k e l p r o d u c t i o n
facilities at Sudbury, O n t a r i o and T h o m p s o n ,
M a n i t o b a , m a y ultimately lead to assistance i n
the adoption of n e w technology at N o r i l s k
(Saunders 1990). E v e n i f emissions of sulfur
components are reduced i n the future, recovery
of vegetation i n severely damaged areas w i l l require a l o n g time. Lichens, i n particular, must
26
A r u p , U . , E k m a n , S., Froberg, L . , K n u t s o n , T . a n d
Mattson, J . E . 1989. Changes i n the l i c h e n f l o r a o n
R o m e l e k l i n t . - Graphis Scripta 2: 148-155.
Canadian Department of Energy Mines and Re-
sources. 1988. Map 900A, Principal mineral areas.
Hawksworth, D . L . 1971. L i c h e n s as l i t m u s f o r air
p o l l u t i o n : A h i s t o r i c a l r e v i e w . - International Jour-
nal Environmental Studies 1: 281-296.
Hawksworth, D . a n d M c M a n u s , P. 1989. L i c h e n s
that tell a tale. - Country Life 183: 144-145.
Helle, T . , N o r o k o r p i , Y . a n d Saastamoinen, O .
1990. R e d u c t i o n o f arboreal lichens i n t w o spruce
stands i n n o r t h e r n F i n l a n d between 1976 a n d 1988.
- I n : K . K i n n u n e n a n d M . V a r m o l a (eds.) Effects
of air pollutants a n d acidification i n c o m b i n a t i o n
w i t h climatic factors o n forest, soils, and waters i n
n o r t h e r n Fennescandia. Nordic Council of Ministers,
Nord 1900: 20.
Rangifer, 12 (1), 1992
Kershaw, K . A . 1977. Studies o n l i c h e n - d o m i n a t e d
systems. X X . A n e x a m i n a t i o n o f some aspects o f
the n o r t h e r n boreal lichen w o o d l a n d s i n C a n a d a . Canadian Journal Botany 55: 393-410.
Klein, D . R. 1982. F i r e , lichens, a n d c a r i b o u . - Journal Range Management 35: 390-395.
Klein, D . R . 1987. Vegetation recovery patterns f o l l o w i n g o v e r g r a z i n g b y reindeer, i n St. M a t t h e w Island. - Journal Range Management 40: 336-338.
Klein, D . R. 1989. N o r t h e r n subsistence h u n t i n g
economies. - I n : R . J . H u d s o n , K . R . D r e w a n d L .
M . B a s k i n (eds.) Wildlife production systems: Economic utilization of wild ungulates. C a m b r i d g e U n i versity Press, C a m b r i d g e p p . 9 6 - 1 1 1 .
Kovalev, B. I. a n d Filipchuk, A . N . 1990. Forest
c o n d i t i o n s i n the zone o f p o l l u t i o n influence f r o m
the N o r i l s k M i n i n g a n d M e t a l u r g i c a l c o m p l e x . Forest Management 5. (In Russian).
LeBlanc, F . a n d Rao, D . N . 1966. R e a c t i o n dequelques lichens et mousses epiphytiques a V a n h y d r i d e
sulfureaux dans l a region de S u d b u r y , O n t a r i o . Bryologist 69: 338-346.
Rangifer, 12 (1), 1992
Norwegian Institute for A i r Pollution. 1984. Emission sources in the Soviet Union. Lillestrøm, N o r w a y . 29p.
Pucket, K . J., Nieboer, E . , Flora, W . P. a n d R i chardson, D . H . S. 1973. S u l p h u r d i o x i d e : its effects o n p h o t o s y n t h e t i c 1 4 C f i x a t i o n i n lichens a n d
suggested mechanisms o f p h y t o t o x i c i t y . - New Phytology, 72: 141-154.
Rundel, P. W . 1978. T h e ecological role o f secondar y substances. - Biochemical Systematics and Ecology
6: 157-170.
Saunders, A . 1990. P o i s o n i n g the arctic skies. Arctic Circle 1: 2 2 - 3 1 .
Vlasova, T . J., Kovalev, B . I. a n d Filipchuck, A . N .
1991 (In press). Effects o f p o i n t source atmospheric
p o l l u t i o n o n boreal forest vegetation o f northwestern Siberia. - I n : G . W e l l e r (ed.) Proceedings International Conference on the Role of Polar regions in
Global Change. F a i r b a n k s , A l a s k a . J u n e 1990.
Manuscript accepted 10 January, 1992
27