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Caloplaca stantonii sp. nov. and Its Relationship to Caloplaca bolacina and Other Lobate to
Squamulose Species in North America
Author(s): Ulf Arup
Source:
The Bryologist,
Vol. 95, No. 4 (Winter, 1992), pp. 449-457
Published by: American Bryological and Lichenological Society
Stable URL: http://www.jstor.org/stable/3243571 .
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The
Bryologist
95(4),
1992,
pp.
449-457
Copyright
?
1992
by
the American
Bryological
and
Lichenological Society,
Inc.
Caloplaca
stantonii
sp.
nov. and Its
Relationship
to
Caloplaca
bolacina
and Other Lobate
to
Squamulose Species
in North
America
ULF
ARUP
Department
of
Systematic Botany,
University
of
Lund,
Ostra
Vallgatan
18,
S-223
61
Lund,
Sweden
Abstract.
Caloplaca
bolacina
(Tuck.)
Herre and
C.
stantonii
sp.
nov.
belong
to
a
group
of
species
in
North America with more
or
less
lobate,
subsquamulose
to
squamulose
thalli. The
two
species
occur
mainly
on
rocks
in the coastal
mountains and on maritime
rocks in western
North
America.
Caloplaca
bolacina is here
lectotypified
and
provided
with a
thorough
description.
Caloplaca
stan-
tonii,
sometimes
confused
with C.
bolacina,
was
in
1989 distributed in
Lich. Exs.
Colo
No. 672
by
W. A.
Weber but
is
described
for
the
first
time in this
paper.
The
present
study
was carried out
to
clarify
the
taxonomy
of
C. bolacina
and C. stantonii and
their
relationship
to a
group
of
similar
species.
This
group,
here
referred
to
as the C.
squamosa
group,
comprises
many
species,
both
European
andAmerican.
A
key
to
the treated taxa
is
included.
The
taxonomy
within
the C.
squamosa
group
is not dealt with.
The results show that
C.
bolacina,
C.
stantonii,
and the
C.
squamosa
group
are
separated
from
each other
principally
in several anatomical
characters
as well as in
gross
morphology.
Caloplaca
bolacina
(Tuck.)
Herre
belongs
to a
group
of
species
with
yellow
to
orange
and
crustose,
lobate
to
squamulose
thalli. This
species
grows
mainly
on
rocks,
from sea level to
higher
altitudes
in
the coastal mountains
in
western
North America.
It
was described from California
by
Tuckerman
(1866)
and also included
in
keys
in
Hasse
(1913)
and
Fink
(1935).
It
has since then
been
a
species
fairly
well-known
by lichenologists
in
North Amer-
ica,
at
least
in
the
western
parts. However,
the ecol-
ogy
and
geographical
distribution
of
C.
bolacina
as
well as the
delimitation
of it from
other lobate
to
squamulose
species
has not
previously
been
fully
understood
(Noble
1982).
Also
Brodo
et al.
(1987)
pointed
out the need of further studies
ofC.
bolacina
and
similar
species.
In 1989
a
new
species
was distributed
by
Dr. W.
A.
Weber
in
Lich. Exs.
Colo
No. 672.
A
proper
description
was
to
be
published
later.
However,
since
in
this
investigation
I studied the
new
species
close-
ly,
both
in
the
field
and
in
the
herbarium,
Dr.
Weber
was
kind
enough
to
allow me to describe
Caloplaca
stantonii
in
this
paper. Caloplaca
stantonii is also
somewhat
squamulose
and has on some occasions
been confused
with
C.
bolacina,
which it
may
re-
semble.
The
purpose
of this
study
was to
clarify
the tax-
onomy
of
C.
bolacina
and
C.
stantonii and
their
relationship
to
other
lobate or
squamulose
species.
These
species
constitute
a
group
of
taxa,
including,
e.g.,
C.
squamosa
(B.
de
Lesd.)
Zahlbr.,
C.
modesta
(Zahlbr.)
Fink,
C.
sipeana
Magn.,
C.
intermedia
(B.
de
Lesd.)
Zahlbr.,
C.
subsoluta
(Nyl.)
Zahlbr.,
and
C.
irrubescens
(Nyl.)
Blom. The
taxonomy
within
this
group (in
this
paper
referred to as
the
C.
squa-
mosa
group)
is
still
to
be clarified and more
species
may
be involved than listed
here.
However,
further
studies
will
probably
prove
that
there
are
many
more
names
than
acceptable species.
The
nomenclature
though,
is
very
hard
to
settle because
of
difficulties
in
locating
the
type
material.
MATERIALS AND METHODS
The
study
is
based
on
material collected on a field
trip
to North
America
in 1989
and material
from
the
following
institutions:
ASU,
Bratt
(priv.
herb.),
CANL,
COLO,
FH, LD,
MIN, NY, UBC, us,
wwB.
A
total
of
ca.
130
specimens
of
Caloplaca
bolacina
and 16
specimens
of
C. stantonii
are
included
in
the
study.
My
own
material
is
deposited
at
the Botanical museum
in
Lund
(LD).
The
material was
examined
by
interference and
light
microscopy.
Measurements
of
morphological
characters
were
carried out on
dry
material,
using
a
dissecting
mi-
croscope.
Sections of the
apothecia
were
made both
by
hand
and
with a
freezing
microtome.
All
measurements
of anatomical
features
were
carried out on sections mount-
ed
in
water.
The
thickness of the
hymenium
was
recorded
by measuring
its
thickest
part.
However,
the variation
in
this feature is
very
little
within each
apothecium,
often
within
5
utm.
The
spores
were
released
from
the asci and
measured
in
water.
Since
only
small
differences between
old
and
fresh
spores
could be
recognized,
no
treatment
according
to
Steiner
and
Peveling (1984)
was carried
out.
Only
mature
and
normal-shaped spores
were
measured.
The
chemistry
was examined
by high
performance
thin-
0007-2745/92/449-457$1.05/0
450
THE
BRYOLOGIST
[VOL.
95
layer
chromatography (HPTLC)
as
described
by
Arup
et
al.
(1992),
a somewhat different
procedure
than that de-
veloped
by
Culberson
(1972, 1974).
RESULTS
Caloplaca
bolacina and
C.
stantonii
differ con-
siderably
from
the
C.
squamosa
group
in
structure
of the
thallus and the
excipulum.
Caloplaca
bola-
cina and
C.
stantonii
both have
prosoplectenchy-
matous,
anticlinally
to
irregularly
arranged
hyphae
with
oblong
cells.
Caloplaca
bolacina
has rather nar-
row
hyphae
with cells
much
longer
than wide and
with
lumina
regularly
tube-shaped
(Fig.
1-2),
while
C.
stantonii
has
hyphae
with cells
comparatively
shorter and wider and with
irregular-shaped
lumina
(Fig. 3-4).
The
C.
squamosa
group,
on the other
hand,
has a
paraplectenchymatous
structure of rath-
er thin-walled
hyphae
with rounded to
very
short
rectangular
cells
(Fig.
5-6).
These features
may
in-
dicate that the
three taxa are not
very closely
related
to
each other. Since the
taxonomy
of the
C.
squa-
mosa
group
is not
yet completely
understood,
the
differences discussed
below should be considered
somewhat
tentative. The
apothecia
in
the
C.
squa-
mosa
group
are
usually very
small,
0.2-0.5
mm
across,
but
may
sometimes exceed
1
mm. The
length
of
the
spores
mostly
varies between
9
and 13
Am
in
the
C.
squamosa
group
and between 13 and
17
Am
in
C.
bolacina,
but there is an
overlap
in
the
range
between
12
and 15
Am.
The
spores
of
C.
stan-
tonii,
however,
are
always longer
than
14
Am.
The
lobes and
areoles
in
the
C.
squamosa
group
are usu-
ally
more
or less flat while
in
C.
bolacina and
C.
stantonii
they
are
convex or inflated.
Furthermore,
one or
maybe
two taxa
in
the
C.
squamosa
group
possess
a black
prothallus,
lacking
in
both
C.
bo-
lacina and
C.
stantonii.
In
gross
morphology
at least
one,
maybe
undescribed,
species
(Fig.
10)
in
the
C.
squamosa
group may
be
very
similar to
C.
bolacina,
but
it
can
always
be
separated
by
its anatomical
structures.
Measurements of
morphological
and anatomical
features
in
C. bolacina
and
C. stantonii are
sum-
marized
in
Table
1.
TAXONOMIC TREATMENT
1.
Thallus cortex and
excipulum
paraplectenchyma-
tous,
consisting
of rather
thin-walled,
rounded
to
very
short
rectangular
cells;
apothecia
usually
less
than
0.8 mm
wide;
areoles and lobes
+
flat; spores
mostly
shorter than 13
m
............
C.
SQUAMOSA
group
1.
Thallus cortex
and
excipulum
consisting
of
anti-
clinally
to
irregularly arranged hyphae
composed
of
oblong,
thick-walled
cells;
apothecia
usually
more
than 0.8 mm
wide;
lobes
convex to
inflated; spores
mostly longer
than 13 m
....
........................................
2
2.
Thallus areoles and
squamules
?
inflated,
in-
cised
at
base,
tawny-yellow
to
reddish-orange;
apothecia
tawny-yellow
to
reddish;
medulla
without
crystals; hypothecium
occasionally
with
scattered oil
droplets; spore septum
longer
than
3 um
........................................
C.
BOLACINA
2.
Thallus areoles convex but not
inflated,
mostly
verruculose,
dirty-yellow
to
brownish-yellow;
apothecia
ferruginous
to
brownish;
medulla with
abundant
crystals;
hypothecium
and lower
part
of
hymenium heavily
inspersed
with oil
drop-
lets; spore septum
shorter than 2.5
um
.....................
......................................................................................................
C .
ST A N T O N II
CALOPLACA BOLACINA
(Tuck.)
Herre,
Proc.
Wash.
Acad. Sci. 12:
233. 1910.
(FIG. 1-2, 7-9)
Placodium
bolacinum
Tuck.,
Lich.
Calif.
18.
1866.
TYPE:
U.S.A.
CALIFORNIA.
Oakland
Hills,
Bolander
121
(lec-
totype
selected
here, FH!;
isotype,
us!).
Thallus to 4 cm
broad,
crustose
(Fig.
7),
lobate
to
squamulose
(Fig.
8)
or sometimes
subfruticose,
of
scattered to clustered
turgid
areoles or
imbricate
squamules,
occasionally
almost absent
(Fig.
9)
and
present
only
as
small
lobes at
base of
apothecia;
areoles
up
to 2.0
x
1.8
mm
wide,
0.25-0.90
mm
thick,
initially
more or less round but often becom-
ing irregular,
lobate or
squamulose,
in
all
stages
more
or less
inflated and incised at
base;
in
cross
section
showing
thick-walled
hyphae
with
oblong
cells
(Fig.
1), tightly
packed
in
vertical bundles
breaking
the
algae
into
clumps,
becoming irregular
and more loose
in
upper cortex;
cortex 45-80
Am
thick,
with
up-
permost
15-25
Am
pigmented,
often with
unpig-
mented
amorphous epinecral
layer
above; squa-
mules with or without
medulla,
sometimes with
lower cortex of
irregular
thick-walled
hyphae;
thal-
lus
color
varying
from clear
citrine-yellow,
buff,
tawny-yellow,
yellow-orange, orange
to
deep
red-
dish-orange, mostly very
smooth
and somewhat
shiny,
occasionally heavily
white-pruinose;
pro-
thallus
absent.
Apothecia
(Fig.
7-9)
rarely
lacking, mostly
fairly
abundant to
abundant,
lecanorine,
scattered to
crowded,
most often sessile to
raised but
occasion-
ally shortly
stalked or
immersed,
0.8-1.7(-2.7)
mm,
FIGURES
1-6.
Caloplaca
bolacina,
C.
stantonii,
and the
C.
squamosa
group.
-
1-2.
C.
bolacina.
-
1. Cross
section
of thallus
cortex; California,
Nash
8029
(ASU).
-
2. Cross section of
upper part
of
excipulum; California,
Anacapa
Island,
Fard,
1889
(FH).
-
3-4.
C.
stantonii.
-
3. Cross section of thallus
cortex; California,
Arup
L89224
(LD).
-
4.
Cross
section of
excipulum;
Oregon,
Coos
Co.,
Sunset
Bay
State
Park,
Arup
L89222
(LD).
-
5-6.
C.
squamosa
group.
Cross section of thallus cortex and
excipulum;
Arizona,
Lich.
Exc.
Colo
414
(LD).
Bars
=
10
utm.
1992]
ARUP: CALOPLACA
451
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-
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Y
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-
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Ik,
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,,.
a,,.
.,
*
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ppp
1
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,,
00
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I
f
C.
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r'
C
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4c
oo
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7
-
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-4
.
.
L. ?I
L
\
4L.0
fIoe
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7
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t
~ lI
,
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-_
,
;.'•:
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-
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.:
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,,,,,
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r'l.-
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C- (C-
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q
it
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,
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tl,
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v.
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452
THE BRYOLOGIST
[VOL.
95
.16
,
Ii
'
?1L
,3
,i
c
•r-
.
"
-kg
r,~~
r
.?t
",.
FIGURES
7-10.
Caloplaca
bolacina
and the
C.
squamosa
group.
-
7-9.
C.
bolacina.
-
7.
A
rather
typical
form;
crustose
to somewhat
lobate
inflated
areoles;
Mexico,
Parish 8814
(FH).
-
8.
An
extreme
form;
thallus
of imbricate
squamules
or almost
subfruticose;
California, Arup
L89242
(LD).
-
9.
Typical
maritime
form;
California,
Arup
L89236
(LD).
-
10.
C.
squamosa group.
A
taxon
in
the
group
which
may
resemble
C.
bolacina
very
much;
Arizona,
Lich. Exc.
Colo
414
(LD).
Bars
=
1
mm.
initially
more
or
less round
but
soon
irregular,
disk
concave
to
convex
but
mostly
more
or less
flexuose,
smooth
or
finely
granulate,
sometimes
cracked
in
very
large
apothecia,
tawny-yellow,
yellow-orange,
brownish-orange,
bright-orange
to
deep-orange
or
red,
always
darker
than
thallus
and
margins;
proper
margin
mostly
rather
thick
to
very
thick,
(25-)75-
150(-200)
tm,
smooth
or
with
weak
radiating
ridg-
es,
in most
stages
conspicuously
raised
but
with
age
sometimes
level
with
disk, consisting
of thick-walled
hyphae
with
oblong
cells
(Fig.
2),
rather loose and
unoriented
to
tightly
packed
and
more or less ra-
diating;
thalline
margin
thin to
very
thick,
some-
times
almost
excluded,
to 250
,m,
even to
flexuose,
often confluent
with
proper
margin,
in
well-devel-
oped
thalli
often
cracked,
concolorous
with
thallus;
hymenium
(70-)75-90
gm
high, hyaline,
epihy-
menium sordid
brownish-orange, granular
in-
spersed;
hypothecium
hyaline,
(90-)120-190(-230)
,gm
thick,
sometimes
with
oil
droplets up
to 5
im
broad;
paraphyses
simple
or
occasionally
branched,
1-2
gm
broad,
the
tips
mostly
capitate,
4-7
gm
1992]
ARUP: CALOPLACA
453
TABLE
1.
Mean,
standard
deviation,
and
range
for
quantitative
character
measurements
on
Caloplaca
bolacina and
C.
stantonii.
n = number of
specimens
studied. The
figures
on
spore
dimensions are calculated for n
=
154
(C.
bolacina)
and
n = 65
(C.
stantonii).
C.
bolacina
(n
=
30)
C.
stantonii
(n
=
12)
Thickness of thallus
(mm)
0.38
(0.07),
0.25-0.90
0.6
(0.06),
0.15-1.20
Maximum width
of
apothecia
(mm)
1.56
(0.40),
0.8-2.7
1.13
(0.25),
0.8-1.5
Minimum
width of
proper
margin
(mm)
90
(25),
25-150
48
(27),
0-100
Maximum width
of
proper
margin (mm)
141
(27),
75-200
166
(27),
125-200
Thickness of
hymenium (Am)
82
(6),
70-90
91
(10),
75-110
Thickness of
hypothecium (Lm)
165
(44),
90-270
167
(45),
115-270
Length
of
spores
(Lm)
14.88
(1.83),
10.5-19.0
17.22
(1.65),
14.0-20.6
Width of
spores
(Lm)
7.02
(1.03),
5.0-10.3
6.78
(0.77),
5.0-8.8
Width of
septum
(Am)
4.04
(0.83),
2.5-5.7
1.88
(0.40),
0.8-3.0
Ratio
of
spore length
to
spore
width 2.15
(0.32),
1.46-3.22
2.56
(0.33),
2.04-3.44
Ratio
of
septum
width to
spore
length
0.27
(0.05),
0.16-0.40
0.11
(0.02),
0.05-0.15
broad;
asci
cylindrical,
48-77
x
12-20
,m,
with 8
spores; spores polaribilocular, ellipsoid,
(10.5-)12.4-
17.5(-18.5)
x
(5.0-)5.5-8.5(-10.3)
gm,
septum
(2.5-)3.0-5.2(-5.7)
#m
(extreme
values within
pa-
rentheses
=
ca.
7%
of the
variation); pycnidia mostly
present
and
common,
appearing
as small
depres-
sions
or
rarely
somewhat
raised,
darker
than
thallus;
conidia
bacilliform,
2.5-3.5
x
1.0-1.5
im.
Chemistry.--Parietin
and
fallacinal as
main
pig-
ments and
smaller amounts
of
emodin, xanthorin,
and one unknown
anthraquinone.
Cortex
and
apo-
thecia
K+
purple, C-,
Pd-,
I-;
medulla
K-,
C-
to
C+
slowly
rose to
red,
Pd- and
I-.
Ecology.
-
Caloplaca
bolacina
is
mainly
saxico-
lous,
growing
on all
kinds
of
rocks,
both
acidic and
calciferous,
but
may
also be found
on
earth and
according
to Noble
(1982)
even on
driftwood.
The
species
seems to
prefer sun-exposed
rocks and boul-
ders
in the Coast
Range
and occurs at least
up
to
1,600
m. On
these
sites it is
mostly
well
developed
and
richly
fertile but
in
more
protected
habitats,
as
in
shaded
valleys
and
canyons,
it
is
often sterile and
built
up
of scattered areoles.
Caloplaca
bolacina
also
occurs
on
seashore
rocks where
it--together
with
strictly
littoral
species
of the
genus-forms
the
"or-
ange
belt."
The
vertical
range
on
the shore
is
rather
wide,
reaching
down
close
to the
supralittoral
fringe
(zonation
according
to
Stephenson
&
Stephenson
1949).
It
is,
however,
more
frequent
in
the littoral
zone but
prefers
growing
along
fissures
in
the
rock
rather than
on
exposed
surfaces.
The
species
is
mostly
fertile,
even on
the
seashore,
but becomes less so
when
growing
really
close to the sea.
Distribution.
-In the Coast
Range
in
northern
Mexico and southern California C. bolacina
seems
to
be
rather
common
(Fig.
11).
Further
north it
is
more rare and
according
to available
herbarium ma-
terial
restricted to
seashore rocks.
Caloplaca
bola-
cina
is
probably
a
temperate
to
subtropical
species,
avoiding
localities
with
cold
winters,
preferring
the
dry
and
hot
climate
in
Baja
California.
There are some
specimens
determined
to
C.
bo-
lacina
in
herbaria,
collected
in
and
east
of the
Rocky
Mountains.
They
all
seem to
be
misidentified and
so
far I
have not seen
any correctly
determined
ma-
terial
from
east
of
the
Rockies. The
misidentified
specimens
all
belong
to the
C.
squamosa
group.
Remarks.-
The
thallus
in
C.
bolacina
varies
very
much,
and
it is
easy
to
believe that
different
extreme
forms
belong
to
separate
species.
However,
much
of the variation
may
be
found
in
only
one
specimen.
Furthermore,
the variation is continuous
and
does
not allow
the
separation
of more
than
one
species.
Weber
(label
data
in
C.
bolacina,
Lich. Exs.
Colo
No.
657)
stated
that the extreme subfruticose forms
may
appear
as the result
of a
process
in
which the
interstices between the
stipes
or
squamules
are
filled
with soil
washing
in
from
above;
as
the thalli
be-
come
larger
the
stipes
shrink,
the soil washes
away
leaving
the
subfruticose
thalli.
I
do
not want to con-
firm
or
deny
the
possibility
of the described
process
but
I
do believe that
in
C. bolacina
both the crustose
and the subfruticose
forms
belong
to
only
one
spe-
cies.
In
Ryan (1988),
one
specimen
referred to
as
Cal-
oplaca sp.
4
from
Fidalgo
Island,
Washington,
be-
longs
to
C.
bolacina.
Other
specimens,
referred
to
by Ryan
as
Caloplaca sp.
4,
belong
to
C.
rosei Hasse
and to C. marina
(Wedd.)
Zahlbr.
subsp.
americana
Arup
(Arup 1992).
Recently,
Laundon
(1992)
pointed
out
the value
of
using
C
(bleach)
in
taxonomic
work
in the
genus
Caloplaca.
But
since the reaction to
the
spot
test
often varies within
species,
one must be careful when
evaluating
it.
During
the
past
years
I
have made
the
same
observations
as
Laundon and
I
agree
with
his
remarks.
Laundon
also
suggests
that
the
varying
reaction
to C
may
be due to
varying
amounts
of
emodin
in
the
lichen.
However,
both
emodin and
parietin,
as
well as
several
other
anthraquinones
respond
to
C,
becoming
rose, red,
or violet.
Selected
specimens.--CANADA.
BRITISH
COLUMBIA.
Vancouver Island:
Ashley Bay,
Arup
L89250
(LD).
454
THE BRYOLOGIST
[VOL.
95
400
Sa
g
00
i'
'-
'~
(
I
\
40
0
FIGURE
11. The
known distribution
of
Caloplaca
bolacina.
MEXICO.
BAJA
CALIFORNIA.
3.1
mi. south
of
Colonet,
Nash 4483
(ASU);
31 km E
of Rosario
near El
Aguajito,
Wetmore 64055
(MIN);
Coronado
Island,
Parish
8814
(FH).
BAJA
CALIFORNIA
DEL NORTE.
1
mile
E
of ocean near Bahia
de San
Quintin,
Nash 8732
(ASU,
CANL,
MIN, NY,
UBC);
Cerro
Kentin,
Ryan
21241
(ASU).
BAJA
CALIFORNIA
SUR.
W
of San
Ignacio,
Nash 8940
(ASU,
CANL,
MIN).
U.S.A.
CALIFORNIA.
Anacapa
Island,
Fard,
1889
(FH).
LOS ANGELES
CO.:
San
Clemente
Island,
Weber &
Santesson
L-42616
(coLO);
Santa
Catalina
Island,
Nuttall
488
(FH),
Bratt
4185
(Bratt).
MARImN
CO.:
Mill
Valley,
Herre,
1930
(NY).
MENDOCNO
Co.:
Russian
Gulch State
Park,
Arup
L89243
(LD).
MONTEREY
CO.:
Carmel
Valley,
Tavares et
al.
920
(FH).
SAN
FRANCISCO
Co.:
San
Francisco,
Sutro
Heights,
Herre 268
(us).
SAN
MATEO
CO.:
Pilarcitos
Creek
Canyon,
Thompson
4743
(FH).
SANTA
BARBARA
CO.:
Figueroa
Mountains,
Arup
L89231
(LD);
Santa
Cruz
Island,
Weber
&
Bratt 657
(Bratt),
Arup
L89241
(LD);
Santa
Cruz Moun-
tains,
Pescadero
Point,
Herre
718
(FH, US).
OREGON.
COOS
CO.:
Sunset
Bay
State
Park, Arup
L89245
(LD).
CURRY
CO.:
Humbug
Mtn. State
Park,
Arup
L89244
(LD).
LINCOLN
CO.:
3 mi.
Nof
Newport,
Wetmore 24593
(MIN).
WASHINGTON.
JEFFERSON
CO.:
Ruby
Beach,
Arup
L89249
(LD).
WHATCOM
CO.:
Larrabee
State
Park,
Ryan
3397
(wwB).
CALOPLACA
STANTONII
W.
A. Weber
ex
Arup sp.
nov.
(FIG.
3-4,
12-14)
TYPE:
U.S.A.
CALIFORNIA.
SANTA
BARBARA
CO.:
Santa
Cruz
Island,
Fraser
Point,
on
poorly
consolidated,
crum-
bling
rock
near
the
ocean,
1986,
Weber,
Bratt
&
Hogan,
Lich.
Ex.
Colo
No. 672
(holotype
LD).
Thallus
crustaceous
vel
subsquamulatus,
verruculosus
vel
exasperatus,;
continuus
vel ex areolis
dispersis
com-
positus,
fulvus,
plerumque
opacus,
interdum
structuris
isi-
diiformibus
simplicibus
vel
sparse
ramosis
instructus;
me-
dulla
copiose
crystallifera;
prothallus
deest.
Apothecia
pauca
vel
numerosa,
plerumque
sessilia,
0.5-
1.5
mm
diam.,
atroferruginea
vel
brunneola,
margo
pro-
1992]
ARUP: CALOPLACA
455
rOr
CAI
Adpg
C1
~s;?2
FIGURES
12-13.
Caloplaca
stantonii.
-
12. A
well-developed
form from the
type locality;
note the
areoles
lifting
from
substrate,
revealing
the white medulla
(arrow);
California,
Arup
L89224
(LD).
-
13.
A
form
showing
characteristic,
almost
globose,
isidia-like
outgrowths (arrow)
from the
edge
of the
areoles;
Oregon, Arup
L89223
(LD).
Bars
=
1
mm.
prius
primitus
crassus
demum
tenuescens,
margo
thallinus
tenuissimus, suppressus,
sporae
polaribiloculares,
14.7-
19.6
x
5.9-7.7
jum,
septum
1.5-2.5
jum.
Etymology.
-The name of
the new
species
was
proposed
by
Mrs.
C. Bratt
and
Dr. W. A.
Weber
and
refers
to Dr.
Carey
Stanton. Dr. Stanton
owned
most of
Santa Cruz
Island,
where the
type
specimen
was
collected,
and allowed the establishment of
a
research station
on
the
island.
For a
long
period
of
time he
encouraged-and
supported
financially-
biological
as
well as historical
research
on
the Chan-
nel Islands.
Thallus
(Fig. 12-13)
ca. 1-5 cm or sometimes
more,
often
coalescing
to
cover
larger
areas,
crustose
to
subsquamulose,
verruculose to
rough,
composed
of scattered
to
slightly
discrete
areoles,
or
more
or
less continuous
with a
few
irregular
cracks;
areoles
0.3-1
mm
broad,
0.3-1.2
mm
thick,
most often
thick
due
to
well-developed
medulla,
sometimes
lifting
from substrate at
one
end,
curving
upwards,
exposing
white medulla
(Fig. 12),
never
becoming
sorediate
but
occasionally
with
simple
to
sparsely
branched,
somewhat
globose,
isidia-like
structures
on
edges
of
areoles
(Fig.
13);
in
cross section show-
ing
an
up
to 100
gm
thick
amorphous, epinecral
layer;
cortex of
anticlinal thick-walled
hyphae
with
oblong
cells
(Fig. 3),
continuous
algal layer,
and
thick medulla with a
large
amount
of
crystals
(to
some
degree
soluble
in
HC1,
but not
in
HNO3
or
KOH); margin
of scattered areoles
or more or less
continuous and
definite;
thallus color
varying only
slightly,
from
some shade of
yellow
to brownish-
yellow,
mostly
dull;
prothallus
absent.
Apothecia
(Fig.
12-13)
few to
abundant,
some-
times
lacking,
rarely
crowded,
lecanorine,
mostly
sessile but
occasionally
adnate or
raised,
0.5-1.5
mm,
more or
less round to
slightly irregular,
disk
more or
less
plane
to
convex,
sometimes
flexuose,
?
.
.7
".
"-
' "
,,
.3
Il
?
r?.
o•
.
.
,
.
-
,•
•
*? .J4
*#
y~'\
i.4,;-
..
..
•-'q
.
.,
.
•
\
',
,
.
•,••.z
•
•..
,-.
?r
:.'
".
ll
•,•~
~ ~~
?
,
-,•",2"
"."
k
?
"
•
-•"•
,
",?''•
-
t
.'
."i
'
.
'
.I•
..
" I
FIGURE
14.
Caloplaca
stantonii.
Cross section
showing
the
unusually
thick
cortex of
the thalline
margin; Oregon,
Arup
L89222
(LD).
Bar
=
50
,m.
456
THE
BRYOLOGIST
[VOL.
95
V
I
/
D
\0
'
\
I
*4
FIGURE
15.
The known distribution
of
Caloplaca
stantonii.
dark-ferruginous,
brownish-red
to
brownish; proper
margin
in
young
apothecia
often 125-200
Am
but
with time
becoming
thinner
(Fig.
12),
varying
from
25-75
Am
in
mature
apothecia,
smooth
but often
flexuose,
slightly
raised above
disk
or at same
level,
concolorous
with or
somewhat
paler
than
disk,
con-
sisting
of
thick-walled,
tightly
packed radiating
hy-
phae
with
oblong
cells
(Fig.
4);
thalline
margin
hard-
ly
visible
or
very
thin,
to 50
jtm
thick,
restricted to
base
of
apothecia,
cortex
unusually
thick
(Fig.
14),
(30-)45-75(-90)
Am;
hymenium
(75-)85-100(-110)
,gm
thick,
hyaline,
epihymenium
sordid brownish-
orange, granular
inspersed;
hypothecium
(115-)140-
190(-215)
gm
thick,
hypothecium
and lower
part
of
hymenium
heavily
inspersed
with oil
droplets,
up
to
14
gm
broad; paraphyses
simple
to
sparsely
branched
above,
1.5-2.5
gm,
the
tips
only slightly
thicker,
3.0-4.5
Am;
asci
cylindrical,
51-72
x
14-
19
,m,
with 8
spores; spores
polaribilocular,
ellip-
soid,
(14.0-)14.7-19.6(-20.6)
x
(5.0-)5.9-7.7(-8.8)
,gm, septum
distinct
or
not,
(0.8-)1.5-2.5(-3.0)
,m
(extreme
values
within
parentheses,
=
ca. 6%
of the
variation); pycnidia
mostly
present,
sparse
to abun-
dant,
somewhat
immersed or even
with
thallus,
fer-
ruginous
to
dark
red;
conidia
bacilliform,
3.5-4.0
x
1.0-1.5
Am.
Chemistry.
-
Apothecia
with
parietin,
chloro-
emodin,
fallacinal,
and smaller
amounts
of
emodin,
xanthorin,
and one unknown
anthraquinone;
thal-
lus with
small
amounts of unknown
anthraquinone.
Thallus and
apothecia
K
+
purple,
C- or C+
slowly
rose to
red,
Pd-, I-;
medulla
K-,
C-,
Pd- and
I-.
Ecology.
--Caloplaca
stantonii
is
saxicolous but
seems
to
prefer poorly
consolidated
and
crumbling
rocks. It is
so far known
only
from acidic
rocks and
grows
on both vertical
and horizontal
surfaces.
The
species,
like
C.
bolacina,
may
be
found both on
littoral rocks
and
in
the
coastal mountains
but seems
to
have a
much narrower vertical
distribution.
So
far no
locality
exceeds 70
m
above sea level.
Cal-
oplaca
stantonii
seems to
avoid
very
exposed
sites
1992]
ARUP:
CALOPLACA
457
and
is
best
developed
a little bit
away
from
the
shore,
but
does not
occur
in
really
sheltered
habitats.
Distribution.
-
Caloplaca
stantonii
has so
far been
found
only
in
California and
Oregon,
in
rather few
localities
(Fig.
15).
It
is
probably
rather uncommon
but
will
surely
be found
in
several
new
places,
e.g.,
Mexico,
in
the
years
to
come.
I
doubt, however,
that
it will
be
found
in
inland localities
since
it
seems
to be
restricted to
the
Coast
Range.
Remarks. -The
oldest
collections
of C.
stantonii
are
not from
Santa
Cruz
Island,
the
type locality,
but
in
FH
there are
some rather old
specimens
from
Santa Cruz
Mountains,
near San
Francisco,
and from
Clifton,
south of Los
Angeles.
The
oldest
one was
collected
by
A.
C.
Herre
in
1903. These
specimens
were all
determined as
Caloplaca
bolacina.
Also
some
specimens
in
ASU
from
the 1970's were filed
under
C.
bolacina. The two
species
bear
resem-
blance
to one
another,
but
C.
stantonii
may
be rec-
ognized fairly
easily by
the
ferruginous
apothecia
contrasting against
the
sordid-yellow
to
tawny-yel-
low thallus.
Furthermore,
the
verruculose to
sub-
squamulose
structure of the thallus is not
similar to
the
inflated
areoles of C. bolacina.
The
species
resembling C.
stantonii
most
in
morphological
respects
is
perhaps Caloplaca
fla-
vovirescens
(Wulf.)
DT. &
Sarnth.
They
share the
sordid-yellow
thallus
and
the
ferruginous
apothecia.
Caloplacaflavovirescens
differs,
however,
in
several
important
features
in
the
apothecia
(e.g.,
hypothe-
cium,
spores,
cortex width
in
thalline
margin)
as
well
as
chemistry
and
presence
of
crystals
in
the
medulla.
Selected
specimens.
-U.S.A.
CALIFORNIA.
LOS
ANGELES
Co.:
Clifton
by
the
Sea,
Hasse 1402
(Fn).
MENDOCINO
CO.:
Russian
Gulch State
Park,
Arup
L89221, L89225,
L89226
(LD).
ORANGE
CO.:
South
Laguna,
Nash &
Rundel 10305
(ASU).
SAN
FRANCISCO
CO.:
San
Francisco,
Lands
End
in
Lincoln
Park,
Nash
11337
(ASU);
Santa
Cruz
Mountains,
Sutro
Heights,
Herre 261
(FH).
SANTA BARBARA
CO.:
Santa
Cruz
Island,
Fraser
Point,
Weber & Bratt
L-80185
(LD);
Trailer
Barranca,
Arup
L89224,
L89227
(LD).
OREGON.
coos
co.:
Sunset
Bay
State
Park,
Arup
L89220, L89222,
L89223,
L89228
(LD).
ACKNOWLEDGMENTS
I
thank
my
supervisor,
Ingvar
Kiirnefelt,
for valuable
comments on
how
to
improve
this
article,
Dr. William
A.
Weber for
his
cooperation,
Mrs. Charis C. Bratt who
made
staying
on Santa Cruz Island
possible,
Dr.
Ove
Almborn
who
helped
me with the Latin
diagnosis,
and
finally
all
the curators who
kindly
loaned me
herbarium material.
LITERATURE
CITED
ARUP,
U. 1992.
Caloplaca
marina
and
C.
rosei,
two
difficult
species
in
North America. THE
BRYOLOGIST
95: 148-160.
,
S.
EKMAN,
L.
LINDBLOM
&
J.-E.
MATTSSON.
1992.
High
Performance
Thin
Layer
Chromatography,
HPTLC,
an advanced method for
screening
lichen sub-
stances.
Lichenologist (in press).
BRODO,
I.
M.,
W.
J.
NOBLE,
T.
AHTI
& S.
CLAYDEN. 1987.
Lichens
new to
North America from
the
flora of
British
Columbia,
Canada.
Mycotaxon
28:
99-110.
CULBERSON,
C. F. 1972.
Improved
conditions
and
new
data for
the identification
of lichen
products
by
a
stan-
dardized
thin-layer chromatographic
method.
Journal
of
Chromatography
72: 113-125.
.
1974. Conditions for the use
of
Merck silica
gel
60
F254
plates
in
the
standardized
thin-layer
chro-
matographic
technique
for lichen
products.
Journal
of
Chromatography
97:
107-108.
FINK,
B.
1935.
The
Lichen
Flora of
the United States.
Ann
Arbor.
HASSE,
H.
E.
1913. The Lichen Flora
of
Southern
Cal-
ifornia. Contributions from The
United
States
Na-
tional
Herbarium,
Vol.
17,
Part
1.
LAUNDON,
J.
R.
1992. New British
species
of
Caloplaca.
Lichenologist
24: 1-5.
NOBLE,
W. 1982. The
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