BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Lichens of the Callahan Mine, a Copper- and Zinc-Enriched Superfund Site in Brooksville, Maine, U.S.A. Author(s): Nishanta Rajakaruna, Tanner B. Harris, Stephen R. Clayden, Alison C. Dibble, and Fred C. Olday Source: Rhodora, 113(953):1-31. 2011. Published By: The New England Botanical Club, Inc. URL: http://www.bioone.org/doi/full/10.3119/10-03.1 BioOne (www.bioone.org ) is an electronic aggregator of bioscience research content, and the online home to over 160 journals and books published by not-for-profit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use . Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.
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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors,nonprofit publishers, academic institutions, research libraries, and research funders in the common goal ofmaximizing access to critical research.
Lichens of the Callahan Mine, a Copper- andZinc-Enriched Superfund Site in Brooksville, Maine,U.S.A.Author(s): Nishanta Rajakaruna, Tanner B. Harris, Stephen R.Clayden, Alison C. Dibble, and Fred C. OldaySource: Rhodora, 113(953):1-31. 2011.Published By: The New England Botanical Club, Inc.URL: http://www.bioone.org/doi/full/10.3119/10-03.1
BioOne (www.bioone.org) is an electronic aggregator of bioscienceresearch content, and the online home to over 160 journals and bookspublished by not-for-profit societies, associations, museums, institutions,and presses.
Your use of this PDF, the BioOne Web site, and all posted and associatedcontent indicates your acceptance of BioOne’s Terms of Use, available atwww.bioone.org/page/terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, andnon-commercial use. Commercial inquiries or rights and permissionsrequests should be directed to the individual publisher as copyright holder.
tomentosum, Trapeliopsis granulosa, and Verrucaria muralis (Brodo
et al. 2001; Hinds and Hinds 2007; Smith et al. 2009; Appendix).
Species characteristic of nutrient-enriched substrata were also
well represented, although it is unclear whether the nutrient
enrichment was via organic matter, including bird and other animal
Table 1. Bioavailable Cu, Zn, Pb, and Cd content, given in ppm (mg/g drysoil), in soil samples collected along Waste Rock Pile 1 (WR), Goose Pond(GP), and Tailings Pond (TP). Means (6 SE) based on three composite samplescollected at beginning, mid, and end points of transects placed at the three sites.Significance (p value , 0.05) based on a one-way ANOVA.
Normal background levels of total Pb in soils from Maine range
from 10–50 ppm (Bruce Hoskins, Analytical Laboratory, Univ.
Maine, Orono, pers. comm.), whereas worldwide the upper limit
has been reported as 70–100 ppm (Kabata-Pendias 2001; Nash1989). The amounts of Pb extracted from chelators such as DTPA,
used in our analysis (4–7 ppm; Table 1), are much less than the total
content but give a better index of bioavailability (Cui et al. 2004).
Given that the ore and mine tailings contained an average of 5%
(50,000 ppm) and 0.06% (600 ppm) total Pb, respectively
(Environmental Protection Agency 2003), the bioavailable values
we report in Table 1 are much lower. However, the EPA studies
2011] Rajakaruna et al.—Lichens of Callahan Mine 11
document that all organisms tested in and around the Callahan Mine
(benthic, aquatic, and salt grass) accumulated significantly higherconcentrations of Pb than the same taxa collected from ‘unpolluted’
reference locations (Environmental Protection Agency 2009). For
example, salt grass at the Callahan Mine accumulated 14 times more
lead than the same taxon collected from the reference locations.
Our ongoing studies of lichens of metal-enriched substrates in
Maine point to an interesting trend. The Callahan Mine (this study)
and the Ni-enriched serpentine outcrop at Pine Hill shared in
common 22 species (29% of the total flora documented), includingBuellia ocellata, a species that we reported for the first time in New
England (Harris et al. 2007). Ecologically, the lichen flora at the
Callahan Mine showed a similar species composition to that of Pine
Hill, with a greater percentage of saxicolous species compared to
terricolous species, and a relatively higher percentage of micro-
lichens among the saxicolous species compared to a significantly
greater percentage of macrolichens among the terricolous species.
Favero-Longo et al. (2004), in a review of lichens of serpentinesubstrates worldwide, suggested that many species occupying Cr-
and Ni-enriched serpentine substrates can also be found on other
calcareous and siliceous substrates. Six of the 76 species we
documented (ca. 8%) were also collected from a calcium-rich
spring seep isolated on the granitic terrain of Mt. Katahdin,
Maine’s tallest mountain (Miller et al. 2005), whereas 35 species (ca.
46%) were shared in common with the subalpine and alpine lichen
floras of Katahdin (Dibble et al. 2009; Hinds et al. 2009). Severalspecies known to occur on nutrient-enriched bird nesting rocks were
also found at the Callahan Mine. They include Physcia dubia and P.
phaea (Hinds and Hinds 2007), Acarospora fuscata and Amandinea
punctata (Smith et al. 2009), and Xanthoria parietina (Rajakaruna et
al. 2009b). These observations suggest that chemically and
physically harsh substrates, or alpine climates, that prevent the
formation of dense vascular vegetation in exposed sites can provide
a competition-free refuge for various lichen species in the region.We hope that this and other studies we have conducted in the recent
past (Rajakaruna et al. 2009a) will generate additional field
exploratory work documenting the biodiversity of unusual habi-
tats—especially rock outcrops—across New England.
ACKNOWLEDGMENTS. The authors thank Laura Briscoe and
Anthony Naples for assistance with lichen collection and initial
12 Rhodora [Vol. 113
identification of specimens, James W. Hinds for verifications and
identification of the macrolichens, and Jose Perez Orozco for the
preparation of Figure 1. For permission to access the Callahan Mine,
we thank: Sally N. Mills (Hale & Hamlin, LLC), the Maine
Department of Environmental Protection, the Environmental
Protection Agency, and the Maine Department of Transportation.
Two anonymous reviewers provided useful comments. The study was
funded by grants from the Maine Space Grant Consortium to T.B.H.
and N.R. and from the Maine Sea Grant (DV-05-009) to N.R.
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16 Rhodora [Vol. 113
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,N
Z;
wid
esp
rea
da
cro
ssN
A;
on
sili
ceo
us
rock
s(N
BM
:T
H3
73-X
)
Asp
icil
iave
rru
cig
era
Hu
eW
RM
IR
ock
Wid
esp
rea
da
cro
ssN
A,
esp
ecia
lly
east
ern
NA
;o
nsi
lice
ou
sro
cks
(NB
M:
TH
42
4-1
)B
aeo
my
ces
rufu
s(H
ud
s.)
GP
MI
So
ilC
osm
op
oli
tan
bu
tm
ost
lyin
tem
per
ate
an
db
ore
al
reg
ion
s;w
ides
pre
ad
acr
oss
NN
A;
on
sha
ded
an
dd
am
pro
cks,
wo
od
,p
eaty
an
da
cid
soil
s,a
nd
ba
rko
fro
ots
(NB
M:
TH
36
5-Y
,T
H3
71-X
,T
H3
76,
TH
39
6,
LB
CM
16
01
7,
LB
CM
16
02
1)
Bu
elli
ao
cell
ata
(Flo
t.)
Ko
rb.
WR
MI
Ro
ckIn
EU
,M
N,
NA
,A
S,
AF
,a
nd
AU
;R
are
inN
E;
on
sili
ceo
us
rock
s,p
ebb
les,
an
dst
on
ewo
rk(N
BM
:T
H4
21)
Ca
lop
laca
ho
loca
rpa
(Ach
.)A
.E.
Wa
de
WR
GP
MI
Ro
ckW
ides
pre
ad
inN
A,
EU
,a
nd
wes
tern
AS
;m
ain
lyo
nsi
lice
ou
sro
cks
sub
ject
ton
utr
ien
ten
rich
men
t;a
lso
on
tree
ba
rk,
wo
od
,a
nd
calc
are
ou
sro
ck,
incl
ud
ing
con
cret
ea
nd
mo
rta
r
(NB
M:
TH
22
0-2
,T
H2
92-5
,T
H3
09-2
,T
H3
23
-X-2
,T
H3
65-X
-1,
TH
39
8-3
,T
H3
99-3
,T
H4
20-1
,T
H4
24-2
,T
H4
35-2
,T
H4
47-1
)
Ap
pen
dix
.C
on
tin
ued
.
18 Rhodora [Vol. 113
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Ca
nd
ela
riel
laa
ure
lla
(Ho
ffm
.)Z
ah
lbr.
WR
GP
MI
Ro
ckC
osm
op
oli
tan
;in
NA
,m
ost
lyin
wes
tern
reg
ion
sa
nd
NE
;o
nd
ust
imp
reg
na
ted
wo
od
an
db
ark
,co
ncr
ete
an
dm
an
-m
ad
eb
asi
csu
bst
rata
(NB
M:
TH
22
2-2
,T
H3
55,
TH
39
8-1
,T
H4
35-1
,T
H4
47-3
)
Ca
nd
ela
riel
lavi
tell
ina
(Eh
rh.)
Mu
ll.
Arg
.W
RG
PM
IR
ock
Co
smo
po
lita
n;
wid
esp
rea
da
cro
ssN
A;
on
sili
ceo
us
an
dca
lca
reo
us
rock
s,ru
stin
gir
on
,st
ain
edg
lass
,w
oo
d,
an
dra
rely
on
ba
rka
nd
soil
(NB
M:
TH
36
5-X
-2,
TH
44
7-2
,L
BC
M1
70
29-1
)C
lad
on
iaca
rio
sa(A
ch.)
Sp
ren
g.
GP
MA
So
ilC
ircu
mp
ola
r,a
rcti
cto
tem
per
ate
;E
U,
SS
A,
NA
,N
AF
,A
S;
com
mo
nin
NE
;o
nw
eak
lyto
stro
ng
lyca
lca
reo
us
soil
ino
pen
ha
bit
ats
;le
ssco
mm
on
on
org
an
icsu
bst
rata
or
on
rock
;in
EU
als
oo
nm
ine
spo
il-h
eap
s
(NB
M:
TH
23
8-2
,T
H4
02-1
)
Cla
do
nia
cerv
ico
rnis
sub
sp.
vert
icil
lata
(Ho
ffm
.)A
hti
GP
MA
So
ilB
roa
dg
lob
al
dis
trib
uti
on
incl
ud
ing
EU
,N
A,
AS
,A
U,
NZ
;co
mm
on
inN
E;
on
soil
ino
pen
ha
bit
ats
;le
ssco
mm
on
on
wo
od
or
mo
ssy
rock
;in
EU
,fr
equ
ent
on
min
esp
oil
-hea
ps
( HC
OA
:T
H4
03)
Cla
do
nia
chlo
rop
ha
ea(F
lork
eex
So
mm
erf.
)S
pre
ng
.W
RG
PM
AS
oil
Co
smo
po
lita
n,
bu
tm
ost
freq
uen
tin
tem
per
ate
an
db
ore
al
reg
ion
s;co
mm
on
inN
E;
on
hu
mu
s,p
eat,
log
s,tr
eeb
ase
s,ro
cks
wit
hth
inso
il( N
BM
:T
H2
39-1
,T
H3
23-Y
,T
H3
26,
TH
38
3,
TH
43
0-2
)
Ap
pen
dix
.C
on
tin
ued
.
2011] Rajakaruna et al.—Lichens of Callahan Mine 19
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Cla
do
nia
cris
tate
lla
Tu
ck.
GP
MA
So
ilE
nd
emic
toE
NA
;co
mm
on
inN
E;
on
log
s,st
um
ps,
tree
ba
ses
(NB
M:
TH
32
1-2
;H
CO
A:
TH
25
1,
TH
26
7,
TH
37
6,
TH
40
5)
Cla
do
nia
flo
erk
ean
a(F
r.)
Flo
rke
GP
MA
So
ilB
roa
dg
lob
al
dis
trib
uti
on
incl
ud
ing
EU
,M
N,
NA
,S
A,
AS
,S
AF
,A
U,
NZ
;co
mm
on
inN
E;
on
tree
stu
mp
s,lo
gs,
mo
ssy
scre
e,fe
nce
po
sts,
pea
t,a
nd
hu
mu
s
(HC
OA
:T
H3
21)
Cla
do
nia
furc
ata
(Hu
ds.
)S
chra
d.
GP
MA
So
ilB
roa
dg
lob
al
dis
trib
uti
on
incl
ud
ing
EU
,M
N,
NA
,S
A,
AS
,A
F,
AU
,N
Z,
sub
an
tarc
tic
isla
nd
s;co
mm
on
inN
E;
on
rock
sw
ith
thin
soil
or
mo
sses
,h
eath
lan
ds,
gra
ssla
nd
sa
nd
law
ns,
an
dli
tter
ov
erro
cks
(NB
M:
LB
CM
14
00
3;
HC
OA
:T
H2
31,
LB
CM
00
7)
Cla
do
nia
gra
cili
s(L
.)W
illd
.su
bsp
.g
raci
lis
GP
MA
So
ilW
ides
pre
ad
inb
ore
al
reg
ion
s;in
NA
,m
ain
lyin
east
ern
ha
lfo
fco
nti
nen
t;in
NE
,la
rgel
yre
stri
cted
ton
ort
hea
ster
nm
ost
are
as;
on
thin
soil
ov
erro
ckin
hea
thla
nd
so
ro
pen
fore
sts
(HC
OA
:T
H3
48)
Cla
do
nia
gra
cili
ssu
bsp
.tu
rbin
ata
(Ach
.)A
hti
GP
MA
So
ilB
roa
dg
lob
al
dis
trib
uti
on
;co
mm
on
inN
E;
on
rott
ing
wo
od
an
dh
um
us
ino
pen
wo
od
s,cl
eari
ng
s,h
eath
lan
ds,
or
on
rock
ou
tcro
ps
( HC
OA
:T
H3
49)
Ap
pen
dix
.C
on
tin
ued
.
20 Rhodora [Vol. 113
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Cla
do
nia
gra
yi
G.
Mer
r.ex
Sa
nd
st.
GP
MA
So
ilIn
EN
A,
EU
,Ja
pa
n;
un
com
mo
nin
NE
;o
nlo
gs,
stu
mp
s,tr
eeb
ase
s(N
BM
:T
H3
03-2
,L
BC
M1
70
28-1
)C
lad
on
iam
aci
len
taH
off
m.
GP
MA
So
ilB
roa
dte
mp
era
te-b
ore
al
dis
trib
uti
on
incl
ud
ing
EU
,M
N,
NA
,S
A,
AS
,A
F,
AU
,N
Z,
an
dsu
ba
nta
rcti
cis
lan
ds;
com
mo
nin
NE
;o
nlo
gs,
tree
ba
ses,
hu
mu
s,ro
cks
wit
hth
inso
il,
an
da
mo
ng
mo
sses
ina
cid
icw
oo
dla
nd
sa
nd
hea
thla
nd
s
(NB
M:
TH
32
1-1
)
Cla
do
nia
ma
xim
a(A
sah
ina
)A
hti
GP
MA
So
ilIn
com
ple
tely
circ
um
bo
rea
lin
oce
an
icre
gio
ns
incl
ud
ing
EU
,N
A,
an
dA
S;
inco
ast
al
an
dm
on
tan
ea
rea
sin
NE
;o
nh
um
us
an
dp
eat
inco
nif
ero
us
fore
sts
an
db
og
s
(NB
M:
LB
CM
14
01
0;
HC
OA
:T
H3
48)
Cla
do
nia
ph
yll
op
ho
raH
off
m.
WR
GP
MA
So
ilC
ircu
mp
ola
rin
Arc
tic,
bo
rea
l,n
ort
h-
tem
per
ate
reg
ion
s,in
clu
din
gE
U,
NA
,S
A,
AS
,su
ba
nta
rcti
cis
lan
ds;
fair
lyco
mm
on
inN
E;
on
mo
ssy
rock
sa
nd
inh
eath
lan
ds
(HC
OA
:T
H4
50)
Cla
do
nia
po
cill
um
(Ach
.)G
rog
no
tG
PM
AS
oil
Bro
ad
glo
ba
ld
istr
ibu
tio
nin
clu
din
gE
U,
NA
,S
A,
AS
,A
F,
NZ
,su
ba
nta
rcti
cis
lan
ds,
an
dA
N;
com
mo
nin
NE
,es
pec
iall
yM
ain
e,V
t.,
Co
nn
.;o
nca
lca
reo
us
soil
s
(NB
M:
TH
20
0-C
)
Ap
pen
dix
.C
on
tin
ued
.
2011] Rajakaruna et al.—Lichens of Callahan Mine 21
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Cla
do
nia
po
lyca
rpo
ides
Ny
l.G
PM
AS
oil
Bro
ad
glo
ba
ld
istr
ibu
tio
n,
incl
ud
ing
EN
A;
fair
lyco
mm
on
inN
E;
on
thin
soil
ov
erro
ck(N
BM
:T
H2
38-1
)
Cla
do
nia
py
xid
ata
(L.)
Ho
ffm
.W
RG
PM
AS
oil
Co
smo
po
lita
n;
com
mo
nin
NE
;o
nth
inso
ilo
ver
rock
,le
ssco
mm
on
lyo
nw
oo
do
rtr
eetr
un
ks;
oft
enin
rath
erd
ryh
ab
ita
ts
(NB
M:
TH
30
3-1
,T
H4
39-2
;H
CO
A:
TH
35
0,
TH
39
0,
TH
40
4)
Cla
do
nia
ran
gif
erin
a(L
.)F
.H.
Wig
g.
GP
MA
So
ilC
ircu
mp
ola
rin
NH
emis
ph
ere
incl
ud
ing
EU
,N
Aa
sfa
rso
uth
as
Fla
.,a
nd
inS
SA
;co
mm
on
inN
E;
on
rock
sw
ith
thin
soil
,m
oss
-lic
hen
hea
ths,
bo
gs,
aci
dic
op
enw
oo
dla
nd
s
(NB
M:
TH
40
6;
HC
OA
:T
H2
46)
Cla
do
nia
rei
Sch
aer
.W
RG
PM
AS
oil
Cir
cum
po
lar
bo
rea
lto
tem
per
ate
,in
clu
din
gE
U,
MN
,N
A,
EA
F,
AS
,A
U,
NZ
;fa
irly
com
mo
nin
NE
;o
nlo
gs
an
dro
cks
wit
hth
inso
il;
esp
ecia
lly
on
min
era
lso
ilin
no
tv
ery
aci
dic
wo
od
lan
ds,
hea
ths,
an
dw
ast
ela
nd
s
(NB
M:
TH
23
5,
TH
26
5,
TH
33
0-Y
,T
H3
42,
TH
34
7,
TH
42
6,
TH
43
0-1
)
Cla
do
nia
sca
bri
usc
ula
(Del
ise)
Ny
l.W
RG
PM
AS
oil
Bro
ad
glo
ba
ld
istr
ibu
tio
nin
clu
din
gE
U,
NA
,S
A,
AS
,N
AF
,A
U,
NZ
,a
nd
sub
an
tarc
tic
isla
nd
s;co
mm
on
inN
E;
on
thin
soil
ov
erro
ck,
on
mo
ssy
sub
stra
tain
fore
sts
an
dfo
rest
op
enin
gs;
als
oo
nh
eav
ym
eta
lm
ine
spo
il
(NB
M:
TH
22
4,
TH
34
6,
TH
35
0,
TH
43
0-3
)
Ap
pen
dix
.C
on
tin
ued
.
22 Rhodora [Vol. 113
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Cla
do
nia
sty
gia
(Fr.
)R
uo
ssG
PM
AS
oil
Bo
rea
la
nd
Arc
tic
reg
ion
so
fN
Hem
isp
her
ein
clu
din
gE
U,
NA
,a
nd
AS
;k
no
wn
fro
mM
ain
e,N
.H.,
Vt.
,a
nd
Ma
ss.;
ma
inly
inw
etb
og
s
(HC
OA
:T
H2
31-2
)
Dib
aei
sb
aeo
my
ces
(L.
f.)
Ra
mb
old
&H
erte
lG
PM
IS
oil
InE
U,
NA
,G
reen
lan
d,
AS
,A
F;
wid
esp
rea
din
EN
Aa
nd
NW
NA
;o
ncl
ay
eyo
rsa
nd
ya
cid
icm
iner
al
soil
alo
ng
roa
dsi
des
,in
hea
thla
nd
s,a
nd
ov
erro
cko
utc
rop
s
(NB
M:
TH
21
4,
TH
22
9,
TH
25
5,
TH
41
3)
Dip
losc
his
tes
mu
sco
rum
(Sco
p.)
R.
Sa
nt.
WR
GP
MI
So
ilC
osm
op
oli
tan
;w
ides
pre
ad
acr
oss
NA
;o
ver
gro
win
go
ther
lich
ens,
mo
sses
,a
nd
org
an
icd
etri
tus
on
the
gro
un
d(N
BM
:T
H2
00-B
,T
H3
73-Y
,T
H3
77,
TH
41
6,
TH
43
9-1
,T
H4
61,
LB
CM
14-0
01-2
)E
vern
iam
eso
mo
rph
aN
yl.
GP
MA
Ro
ckC
ircu
mb
ore
al;
com
mo
nin
NE
,es
pec
iall
yin
Na
nd
mo
un
tain
ou
sre
gio
ns;
ma
inly
on
bra
nch
esa
nd
tru
nk
so
fco
nif
ero
us
tree
s
(HC
OA
:T
H3
16,
TH
36
2)
Fu
scid
eap
usi
lla
Tø
nsb
erg
GP
MI
Ro
ckP
rob
ab
lyco
smo
po
lita
n;
ma
inly
on
ba
rko
rw
oo
do
fco
nif
ers
an
do
ther
aci
d-
ba
rked
tree
s(N
BM
:T
H3
89)
Ap
pen
dix
.C
on
tin
ued
.
2011] Rajakaruna et al.—Lichens of Callahan Mine 23
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Hy
po
gy
mn
iap
hy
sod
es(L
.)N
yl.
WR
GP
MA
Ro
ckA
rcti
c,b
ore
al,
an
dN
tem
per
ate
NA
an
dE
U,
EA
F,
AS
incl
ud
ing
inH
ima
lay
as,
MN
;co
mm
on
inN
E;
on
tree
sa
nd
wo
od
,le
sso
ften
on
rock
(HC
OA
:T
H2
15,
TH
23
3,
TH
29
3,
TH
30
7,
TH
31
5,
TH
32
8,
TH
36
8,
TH
42
9,
TH
46
0)
Lec
an
ora
dis
per
sa(P
ers.
)S
om
mer
f.W
RG
PM
IR
ock
Co
smo
po
lita
n;
wid
esp
rea
din
Ea
nd
CN
A;
on
calc
are
ou
sro
cks,
als
oo
nm
ort
ar
an
dco
ncr
ete
(NB
M:
TH
22-1
,T
H3
17,
TH
32
0-Y
-1,
TH
32
4,
TH
33
5,
TH
33
7-1
,T
H3
65-
X-5
,T
H3
98-2
,T
H4
47-5
,L
BC
M1
70
26)
Lec
an
ora
po
lytr
op
a(H
off
m.)
Ra
ben
h.
WR
GP
MI
Ro
ckC
osm
op
oli
tan
;w
ides
pre
ad
inN
an
dW
NA
;m
ain
lyo
nsi
lice
ou
sro
cks
(NB
M:
TH
25
9-2
,T
H2
83-1
,2
92-4
,T
H3
27,
TH
37
1-Y
-2,
TH
39
1-4
,T
H4
23-
1,
TH
43
8-4
,T
H4
48,
TH
45
3,
TH
46
7-2
,T
H4
68,
LB
CM
17
02
4,
LB
CM
17
02
5-2
)L
eim
on
iser
rati
ca(K
orb
.)R
.C.
Ha
rris
&L
end
emer
GP
MI
Ro
ckW
ides
pre
ad
inE
tem
per
ate
reg
ion
sin
CE
U,
NA
,A
U,
NZ
;o
np
ebb
les
an
dsm
all
sto
nes
ino
pen
are
as
(NB
M:
TH
25
9-4
)L
epto
giu
mim
bri
catu
mP
.M.
Jørg
.G
PM
AS
oil
Pro
ba
bly
circ
um
arc
tic
inN
Aa
nd
EU
,b
ut
dis
trib
uti
on
po
orl
yk
no
wn
;in
NE
,k
no
wn
fro
mo
ne
site
inW
ash
ing
ton
,C
o.,
Ma
ine;
on
calc
are
ou
sso
ila
nd
clif
fle
dg
es,
oft
ena
mo
ng
mo
ss
(NB
M:
TH
40
1)
Ap
pen
dix
.C
on
tin
ued
.
24 Rhodora [Vol. 113
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Lic
hen
oth
elia
con
vex
aH
enss
enG
P*
RP
rob
ab
lyco
smo
po
lita
n;
on
exp
ose
dsi
lice
ou
sro
cks
(NB
M:
TH
25
9-5
,T
H3
91-6
)M
ela
nel
ixia
sub
au
rife
ra(N
yl.
)O
.B
lan
co,
A.
Cre
spo
,D
iva
ka
r,E
ssl.
,D
.H
aw
ksw
.&
Lu
mb
sch
GP
MA
Ro
ckT
emp
era
tea
nd
bo
rea
lN
Aa
nd
EU
,Ic
ela
nd
,M
N,
CA
,N
an
dC
AF
,a
nd
AS
;co
mm
on
inN
E;
on
ba
rko
fb
ran
ches
an
dtw
igs,
esp
ecia
lly
aci
d-
ba
rked
tree
s(H
CO
A:
TH
24
7,
TH
33
8,
TH
35
1,
TH
37
9,
LB
CM
00
5)
Pa
rmel
iasq
ua
rro
saH
ale
GP
MA
Ro
ckC
om
mo
nin
EN
Aa
nd
EA
sia
,ra
rein
WE
uro
pe
an
dW
NA
;co
mm
on
inN
E;
on
tree
s(H
CO
A:
TH
36
2-2
)
Pa
rmel
iasu
lca
taT
ay
lor
GP
MA
Ro
ckC
osm
op
oli
tan
;co
mm
on
inN
E;
on
tree
sa
nd
sili
ceo
us
rock
s(N
BM
:L
BC
M1
70
28-2
;H
CO
A:
TH
23
4,
TH
24
8,
TH
27
6,
TH
30
8,
TH
31
8,
TH
33
1B
,T
H3
32,
LB
CM
00
6)
Pel
tig
era
rufe
scen
s(W
eiss
)H
um
b.
WR
GP
MA
Ro
ckC
osm
op
oli
tan
;fa
irly
com
mo
nin
NE
;o
nso
ilin
op
ena
rea
s(H
CO
A:
TH
38
2,
TH
45
5)
Ph
ysc
iaa
dsc
end
ens
(Fr.
)H
.O
liv
ier
GP
MA
Ro
ckC
osm
op
oli
tan
exce
pt
AN
;co
mm
on
inN
E;
on
ba
se-r
ich
tree
ba
rk,
sto
ne,
an
do
ther
nu
trie
nt-
rich
sub
stra
tain
clu
din
gm
arb
leg
rav
esto
nes
(NB
M:
TH
38
0;
HC
OA
:T
H3
13,
TH
33
1A
)
Ph
ysc
iaa
ipo
lia
(Eh
rh.
exH
um
b.)
Fu
rnr.
GP
MA
Ro
ckC
osm
op
oli
tan
exce
pt
AN
;co
mm
on
inN
E;
on
ba
se-r
ich
tree
ba
rk
(HC
OA
:T
H3
31
A-2
)
Ap
pen
dix
.C
on
tin
ued
.
2011] Rajakaruna et al.—Lichens of Callahan Mine 25
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Ph
ysc
iad
ub
ia(H
off
m.)
Let
tau
WR
MA
Ro
ckC
osm
op
oli
tan
exce
pt
AN
;co
mm
on
inN
E;
on
rock
sen
rich
edw
ith
bir
dg
ua
no
,tr
eeb
ase
s,a
nd
du
st-
imp
reg
na
ted
ba
rk
(HC
OA
:T
H3
93,
TH
42
5)
Ph
ysc
iap
ha
ea(T
uck
.)J.
W.
Th
om
son
GP
MA
Ro
ckT
emp
era
tea
nd
bo
rea
lN
Aa
nd
EU
;fa
irly
com
mo
nin
NE
;o
na
cid
rock
s,es
pec
iall
yth
ose
enri
ched
wit
hb
ird
gu
an
o
(HC
OA
:T
H3
36)
Ph
ysc
iasu
bti
lis
Deg
el.
GP
MA
Ro
ckE
nd
emic
tote
mp
era
teN
A;
fair
lyco
mm
on
inN
E;
on
aci
dic
rock
insh
ad
edo
ro
pen
,m
ois
to
rd
rysi
tes
(NB
M:
TH
28
1-B
)
Pla
cyn
thie
lla
icm
ale
a(A
ch.)
Co
pp
ins
&P
.Ja
mes
GP
MI
So
ilIn
EU
,M
N,
NA
,A
S,
AF
,a
nd
Ta
sma
nia
;in
EN
A,
on
wo
od
,ro
ttin
gb
ark
,w
oo
dy
deb
ris,
an
dh
um
us
(NB
M:
TH
40
2-3
)P
orp
idia
cru
stu
lata
(Ach
.)H
erte
l&
Kn
op
hW
RM
IR
ock
Inte
mp
era
tea
rea
so
fE
U,
MN
,A
F,
NA
,S
A,
AS
,A
U,
an
dN
Z;
on
sili
ceo
us
rock
s(N
BM
:T
H2
41-1
,T
H4
38-1
)P
orp
idia
ma
cro
carp
a(D
C.)
Her
tel
&A
.J.
Sch
wa
bG
PM
IR
ock
InE
U,
MN
,N
A,
CA
,S
A,
AS
,A
F,
AU
,a
nd
NZ
;o
nsi
lice
ou
sro
cks
(NB
M:
TH
37
2-Y
)P
un
ctel
iaru
dec
ta(A
ch.)
Kro
gG
PM
AR
ock
Wid
esp
rea
din
tem
per
ate
reg
ion
s,in
clu
din
gE
NA
,M
exic
o,
Arg
enti
na
,ea
ster
nA
S,
SA
F;
com
mo
nin
NE
;o
ntr
ees,
wo
od
,a
nd
rock
(HC
OA
:T
H3
60)
Ap
pen
dix
.C
on
tin
ued
.
26 Rhodora [Vol. 113
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Ra
ma
lin
ain
term
edia
(Del
ise
exN
yl.
)N
yl.
GP
MA
Ro
ckB
ore
al
an
dte
mp
era
teN
A,
Ru
ssia
;fa
irly
com
mo
nin
NE
;m
ain
lyo
nsi
lice
ou
scl
iffs
an
db
ou
lder
s,le
sso
ften
on
ba
rk(H
CO
A:
TH
25
0-2
,T
H3
10-2
)R
hiz
oca
rpo
nci
ner
eovi
ren
s(M
ull
.A
rg.)
Va
in.
GP
MI
Ro
ckC
om
mo
nin
Ea
nd
Sb
ore
al
reg
ion
sin
clu
din
gN
an
dC
EU
an
dN
A;
on
sili
ceo
us
rock
sa
nd
met
al-
rich
min
esp
oil
(NB
M:
LB
CM
14
00
9-1
)
Rh
izo
carp
on
gra
nd
e(F
lork
eex
Flo
t.)
Arn
old
GP
MI
Ro
ckW
ides
pre
ad
inb
ore
al,
tem
per
ate
reg
ion
s;o
nsi
lice
ou
sro
ckin
op
enh
ab
ita
ts(N
BM
:T
H2
92-1
,T
H4
67-1
)R
hiz
oca
rpo
nin
fern
ulu
m(N
yl.
)L
yn
ge
GP
MI
Ro
ckK
no
wn
fro
mE
U,
Ru
ssia
,G
reen
lan
d,
NA
,a
nd
Fa
lkla
nd
Isla
nd
s;o
nsi
lice
ou
sa
nd
met
al-
rich
rock
s(N
BM
:L
BC
M1
40
09-2
)
Rh
izo
carp
on
leca
no
rin
um
An
der
sW
RG
PM
IR
ock
Pa
n-t
emp
era
tein
NH
emis
ph
ere;
on
sili
ceo
us
rock
sin
op
enh
ab
ita
ts(N
BM
:T
H4
28,
TH
46
6)
Rh
izo
carp
on
red
uct
um
Th
.F
r.W
RG
PM
IR
ock
Co
smo
po
lita
n;
com
mo
nin
NE
;o
nsi
lice
ou
sro
ck(N
BM
:T
H2
26-X
,T
H2
41-X
,T
H2
59-
3,
TH
29
0,
TH
29
2-2
,T
H3
09-3
,T
H3
20-X
-2,
TH
32
3-X
-3,
TH
34
0,
TH
35
7-1
,T
H3
72-X
,T
H3
91-3
,T
H3
98-
5,
TH
39
9-1
,T
H4
33,
TH
43
8-3
,L
BC
M1
40
13,
LB
CM
14
01
4-2
,L
BC
M1
60
19,
LB
CM
21
03
5-1
)
Ap
pen
dix
.C
on
tin
ued
.
2011] Rajakaruna et al.—Lichens of Callahan Mine 27
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Rh
izo
carp
on
rub
esce
ns
Th
.F
r.W
RG
PM
IR
ock
Tem
per
ate
toso
uth
ern
bo
rea
lin
EN
Aa
nd
NW
EU
;o
nsi
lice
ou
sro
ckin
op
eno
rsh
ad
edh
ab
ita
ts(N
BM
:T
H3
39-2
,T
H3
57-2
,T
H4
38-2
,T
H4
58,
LB
CM
14
00
9-3
,L
B1
40
14-
1,
LB
CM
16
02
0-1
,L
BC
M1
90
30)
Rin
od
ina
gen
na
rii
Ba
gl.
GP
MI
Ro
ckW
ides
pre
ad
inN
an
dS
Hem
isp
her
es,
esp
ecia
lly
inco
ast
al
reg
ion
s;o
nn
utr
ien
t-en
rich
edsi
lice
ou
sro
cks,
mo
rta
r,co
ncr
ete,
etc.
,p
art
icu
larl
yin
dis
turb
edsi
tes;
som
etim
eso
nru
stin
gir
on
(NB
M:
TH
22
0-1
)
Sco
lici
osp
oru
mu
mb
rin
um
(Ach
.)A
rno
ldW
RG
PM
IR
ock
Co
smo
po
lita
n;
on
sili
ceo
us
rock
s,m
emo
ria
ls,
met
al-
rich
rock
s;u
nco
mm
on
on
oth
ersu
bst
rata
( NB
M:
TH
28
9-1
,T
H3
09-1
,T
H3
23-
X-1
,T
H3
37-2
,T
H3
39
-1,
TH
35
2,
TH
36
5-X
-3,
TH
37
8,
TH
39
1-5
,T
H4
20-2
,T
H4
23-2
,L
BC
M1
60
23,
LB
CM
17
02
9-2
)S
tere
oca
ulo
nd
act
ylo
ph
yll
um
Flo
rke
WR
GP
MA
Ro
ckD
isco
nti
nu
ou
sly
circ
um
po
lar
inte
mp
era
tea
nd
bo
rea
lre
gio
ns;
fair
lyco
mm
on
inM
ain
e,N
.H.,
Vt.
,a
nd
Ma
ss.;
on
sili
ceo
us
an
dm
eta
l-ri
chro
cks
(NB
M:
TH
22
8,
TH
37
5-X
,T
H4
36,
TH
44
4)
Ap
pen
dix
.C
on
tin
ued
.
28 Rhodora [Vol. 113
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Ste
reo
cau
lon
pil
eatu
mA
ch.
WR
GP
MA
Ro
ckD
isco
nti
nu
ou
sly
circ
um
po
lar
inte
mp
era
tea
nd
bo
rea
lre
gio
ns,
als
oin
mo
nta
ne
sub
tro
pic
s;fa
irly
com
mo
nin
Ma
ine,
N.H
.,V
t.,
Ma
ss.,
an
dN
WC
on
n.;
on
sili
ceo
us
rock
s,b
asa
lt,
min
esp
oil
hea
ps
(NB
M:
TH
25
3,
TH
30
6,
TH
39
7,
TH
40
9,
TH
41
2,
TH
41
9,
TH
44
2,
TH
45
6,
TH
45
9)
Ste
reo
cau
lon
sax
ati
leH
.M
ag
n.
GP
MA
Ro
ckA
rcti
cto
Nte
mp
era
tein
NH
emis
ph
ere;
com
mo
nin
NE
exce
pt
R.I
.;o
nsi
lice
ou
sro
cka
nd
gra
vel
lyso
il( N
BM
:T
H2
98,
TH
37
4)
Ste
reo
cau
lon
tom
ento
sum
Fr.
GP
MA
So
ilC
ircu
mp
ola
ra
rcti
c,b
ore
al,
an
dN
tem
per
ate
inN
Hem
isp
her
e;co
mm
on
inN
NE
,u
nco
mm
on
inS
NE
;o
ng
rav
elly
soil
or
soil
ov
erro
ckin
op
enh
ab
ita
ts
(NB
M:
TH
23
0,
LB
CM
14
01
1)
Sti
gm
idiu
msp
.G
P+
Ro
ckD
istr
ibu
tio
nu
nk
no
wn
;li
chen
ico
lou
so
nL
eca
no
rap
oly
tro
pa
( NB
M:
TH
28
3-2
)T
rap
elia
pla
cod
ioid
esC
op
pin
s&
P.
Jam
esG
PM
IR
ock
InE
U,
Azo
res,
NA
,a
nd
AS
;co
mm
on
inN
EN
A,
rare
else
wh
ere;
on
sili
ceo
us
rock
sa
nd
min
esp
oil
hea
ps
(NB
M:
TH
39
4)
Tra
pel
iop
sis
gra
nu
losa
(Ho
ffm
.)L
um
bsc
hW
RG
PM
IS
oil
Co
smo
po
lita
n;
wid
esp
rea
da
cro
ssN
A;
on
pea
tyso
ila
nd
com
pa
cto
rga
nic
det
ritu
s,ro
ttin
gw
oo
d,
rece
ntl
ych
arr
edw
oo
d(N
BM
:T
H4
32,
TH
43
7)
Ap
pen
dix
.C
on
tin
ued
.
2011] Rajakaruna et al.—Lichens of Callahan Mine 29
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Ver
ruca
ria
mu
rali
sA
ch.
GP
MI
Ro
ckW
ides
pre
ad
inE
U,
MN
,N
A–
SA
,A
S,
AF
,A
U,
an
dN
Z;
on
lim
esto
ne,
mo
rta
r,b
rick
,p
ebb
les,
lim
esto
ne
soil
,a
nd
som
etim
eso
nsi
lice
ou
sro
cks,
mo
stly
ind
istu
rbed
ha
bit
ats
(NB
M:
TH
28
7,
TH
30
1,
TH
32
0-Y
-2,
LB
CM
16
01
6,
LB
CM
17
02
7)
Xa
nth
op
arm
elia
con
sper
sa(E
hrh
.ex
Ach
.)H
ale
GP
MA
Ro
ckT
emp
era
teN
A,
SA
,E
U,
Jap
an
,a
nd
mo
nta
ne
tro
pic
al
are
as;
com
mo
nin
NE
;o
nsi
lice
ou
s,so
met
imes
slig
htl
yn
utr
ien
t-en
rich
edro
cks
(HC
OA
:T
H2
57,
TH
27
1,
TH
36
6)
Xa
nth
op
arm
elia
cum
ber
lan
dia
(Gy
eln
.)H
ale
WR
GP
MA
Ro
ckT
emp
era
teN
A,
SA
,S
AF
;co
mm
on
inN
E;
on
sili
ceo
us
rock
(HC
OA
:T
H2
25,
TH
25
2,
TH
25
6,
TH
26
4,
TH
27
0,
TH
29
9,
TH
31
4,
TH
31
9,
TH
35
4,
TH
34
4,
TH
44
9)
Xa
nth
op
arm
elia
pli
ttii
(Gy
eln
.)H
ale
GP
MA
Ro
ckT
emp
era
teN
A,
SA
,S
AF
;fa
irly
com
mo
nin
NE
;o
nsi
lice
ou
sro
ck(H
CO
A:
TH
32
7,
TH
36
9)
Xa
nth
op
arm
elia
viri
du
lou
mb
rin
a(G
yel
n.)
Len
dem
erG
PM
AR
ock
Tem
per
ate
EN
A;
com
mo
nin
NE
;o
nsi
lice
ou
sro
ck
(HC
OA
:T
H2
64-2
,T
H3
44-2
)X
an
tho
ria
eleg
an
s(L
ink
)T
h.
Fr.
WR
GP
MA
Ro
ckW
ides
pre
ad
inN
Hem
isp
her
ea
nd
pa
rts
of
SH
emis
ph
ere,
exte
nd
ing
far
into
Arc
tic
an
dA
nta
rcti
c;co
mm
on
inN
Eex
cep
tR
.I.;
on
inla
nd
calc
are
ou
sro
cks
an
dco
ast
al
sili
ceo
us
rock
s,in
clu
din
gb
ird
nes
tin
gro
cks;
als
oo
nm
an
-ma
de
sub
stra
tes
lik
eco
ncr
ete
(HC
OA
:T
H4
54,
TH
30
2,
TH
35
3,
LB
CM
03
1)
Ap
pen
dix
.C
on
tin
ued
.
30 Rhodora [Vol. 113
Ap
pen
dix
.C
on
tin
ued
.
Sp
ecie
sS
ite
Na
ture
Su
bst
rate
Ra
ng
e/F
req
uen
cy/S
ub
stra
teE
colo
gy
Xa
nth
ori
ap
ari
etin
a(L
.)T
h.
Fr.
GP
MA
Ro
ckC
osm
op
oli
tan
;in
NA
ma
inly
alo
ng
Atl
an
tic
an
dP
aci
fic
coa
sts
an
dG
ulf
of
Mex
ico
,b
ut
als
oex
ten
din
gin
lan
d;
com
mo
nin
coa
sta
lM
ain
e,N
.H.,
Ma
ss.,
R.I
.,a
nd
Co
nn
.;o
nsi
lice
ou
sa
nd
calc
are
ou
sro
cks,
nu
trie
nt-
enri
ched
tree
tru
nk
sa
nd
bra
nch
esa
nd
oth
ersu
bst
rata
such
as
fen
cep
ost
s,ro
of
tile
s,a
nd
old
bo
nes
(HC
OA
:T
H2
36,
TH
23
7,
TH
24
3,
TH
27
4,
TH
28
0,
TH
29
4,
TH
32
9,
LB
CM
03
2)
2011] Rajakaruna et al.—Lichens of Callahan Mine 31