Definition of the Frasnian/Famennian Stage

Transcription

Definition of the Frasnian/Famennian Stage
4了了
勿 G Ma
ppe
r
,
R Fe
i
s
t
,
R T Be
c
k
e
r a
n
d M R Ho
u
s
e
'
v
W
'
%
e
ini
f
t
i
on of
t
he
Fr
as
ni
an/
Fame
nni
a
n St
age
boundar
y
盆
Th
e bou
nd
ar
yf
or t
he Fr
a
s
n
i
an
l
Fame
nni
an St
age Gl
o
ba
l
S
t
r
at
ot
y
pe Se
c
t
i
o
n a
nd Po
i
nt
(
GSSP)
ha
s be
e
n r
at
i
fe
d勿
I
CS a
nd I
UGS a
nd i
s dr
awn i
n a s
e
c
t
i
o
n e
x
po
s
e
d ne
ar
t
he Up
pe
r Cou
mi
ac Qu
ar
r
y i
n t
he s
ou
t
he
a
s
t
er
n Mo
n-
t
a
gne Noi
r
e
,
Fr
a
nc
e
.
The po
s
i
t
i
on o
f t
he bo
u
nd
ar
y wa
s
s
e
l
e
c
t
e
d勿 t
h
e Su
b
c
ommi
s
s
i
on on De
v
o
ni
a
n St
r
at
i
gr
a
-
l
owe
r t
h
anf
or
me
r
l
y u
s
e
d bo
u
nd
ar
y l
e
v
e
l
s
f
or t
he b
as
e o
f
ph
y i
n 1
9
91 t
o c
o
i
nc
i
d
e wi
t
h t
h
e l
o
we
r bo
u
nd
ar
y o
f t
he
Hi
s
t
o
r
i
c
a
l
l
y
,
s
e
v
e
r
a
l
d
i
f
f
e
r
e
n
t
l
e
v
e
l
s
h
a
v
e
b
e
e
n
u
s
e
d
t
o
d
e
i
f
n
e
t
h
e
t
he Fame
nni
an.
I
nt
roduct
i
on
Lowe
r Pa
l
ma
t
o
l
e
pi
s
t
r
i
a
ng
ul
a
r
i
s
Zon
e i
n t
he c
o
no
do
nt
bi
o
s
t
r
at
i
gr
a
ph
y
.
The r
e
v
i
s
e
d de
ini
f
t
i
o
n o
f t
h
e l
owe
r
bou
nd
ar
y o
f t
he z
on
e
,
pr
o
po
s
e
d h
e
r
e
i
n,
e
x
c
l
u
de
s t
h
e
e
xt
r
e
me
l
y r
ar
e o
c
c
ur
r
e
nc
e
s o
f Pa
l
ma
t
ol
e
p
i
s
t
r
i
a
n
gu
l
a
r
i
s
af
e
w c
e
nt
i
me
t
r
e
s l
owe
r
,
wi
t
hi
n t
he u
pper
mo
s
t
c
on
od
ont
z
on
e
.
Th
e GS
SP al
s
o c
o
i
nc
i
de
s wi
t
h t
he bo
un
dar
y
b
et
we
e
n t
he Cr
i
c
k
i
t
es
h
ol
z
a
p
f
e
l
i
Zo
ne an
d t
he Ph
oe
n
i
x-
i
t
e
s
f
r
e
c
h
i
Zo
ne i
n t
he go
ni
a
t
i
t
e s
c
al
e
.
Th
e po
s
i
t
i
o
n o
ft
he
GS
SP i
s i
mme
di
at
e
l
y abo
v
e a ma
j
or ho
r
i
z
o
n o
f e
xt
i
nc
t
i
o
n
wi
t
h
i
n t
h
e De
v
on
i
an an
d i
s s
t
r
a
t
i
gr
a
phi
c
a
l
l
y s
ome
wh
at
10 KM
0
ba
s
e o
f t
he Fa
me
nn
i
a
n
.
I
n t
he t
y
pe a
r
e
a f
o
r t
he na
mi
ng o
f t
h
e s
t
a
g
e
s
i
n s
o
ut
he
r
n Be
l
gi
um,
p
r
e
ci
s
e d
oc
u
me
nt
a
t
i
on i
n r
ec
e
nt
y
e
a
r
s h
a
s b
e
e
n
gi
ve
n t
o a ne
w r
e
f
e
r
en
c
e s
e
c
t
i
o
n r
e
p
l
a
c
i
n
g t
he no
w-
i
nf
i
l
l
e
d c
l
a
s
s
i
c
s
e
c
t
i
o
n i
n t
h
e Se
nz
e
i
l
l
e
s r
a
i
l
wa
y c
u
t
t
i
n
g (
Bu
l
t
yn
c
k a
n
d o
t
h
e
r
s
,
1
988
)
.
La
r
g
e
l
y r
e
s
u
l
t
i
n
g f
r
o
m wo
r
k i
n t
h
e i
f
r
s
t
qu
a
r
t
e
r
o
f t
hi
s
c
e
n
t
u
r
y i
n Ge
r
-
ma
ny,
a
no
t
he
r bo
un
da
r
y wa
s us
e
d t
h
a
t
was
ba
s
e
d on t
h
e e
n
t
r
y of t
h
e
g
on
i
a
t
i
t
e Che
i
l
oc
e
r
as i
n t
he p
e
l
a
g
i
c r
e
a
l
m.
I
n t
h
e l
a
t
t
e
r
h
a
l
f o
f t
h
i
s
c
en
t
u
r
y t
he c
on
s
i
d
e
r
a
bl
e g
r
owt
h of c
on
od
o
nt
s
t
u
di
e
s ha
s
l
e
d t
o muc
h
r
e
in
f
e
me
n
t o
f t
he bi
os
t
r
a
t
i
g
r
a
ph
y.
Howe
v
e
r
,
t
h
e l
e
ve
l
t
a
ke
n a
s
t
h
e
base of t
he Famenni
an has var
i
e
d bet
ween a l
evel at t
he ba
se of t
he
c
r
e
pi
da Zon
e d
own t
o t
he ba
s
e o
f t
he Lo
we
r t
r
i
an
gu
l
ar
i
s Zo
ne
.
The
n
e
e
d f
or
a
n i
n
t
e
r
n
a
t
i
on
a
l
d
ef
i
ni
t
i
on ha
s b
e
c
ome u
r
ge
n
t
.
The Su
bc
o
m-
,Bedari
eux
A
因
区鼓习
DEVO N !A N
-
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0
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/
,气 L
e
s
..二,二,
/
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与
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U
U
A
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R
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:
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r
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n
q
e
5
/
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}
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二又弓一扮代v
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ac
O
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L一一
」
Nazai
re de
Ladarez
P R O V/刀 C E
六
Bedarieux
Aur
i
gnac
+叼ont
pel
l
i
er
子
Toul
ouse
N
,亡ausses et
V
eyran
A
+Cessenon
。
。
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L卜1
4
Magal
as
卜
、_, .、
s 尸 A I N
Fi
g
u
r
e]
Ma
p
s
s
h
o
wi
n
g t
h
e
po
s
i
t
i
o
n o
ft
e
h
Mo
n
t
a
g
u
e No
i
r
e i
n s
o
u
t
h
e
a
s
t
e
r
n Fr
a
n
c
e
a
n
d o
ft
h
e U
p
p
e
r
户u
mi
a
c_
Qu
a
r
7
y n
e
a
r C
e
s
s
e
n
o
n
,
t
h
e
s
i
t
e
o
ft
h
e
Gl
o
b
a
l
S
t
r
a
t
o
t
y
p
e S
e
c
t
i
o
n a
n
d P
o
i
n
t
(
GS
S
P
)
f
o
r
t
h
e
d
e
in
f
i
t
i
o
n o
ft
h
e b
a
s
e o
ft
h
e
Fa
me
nni
a
n St
a
ge (
i
n
s
e
t
m
a
p A)
.
Mod
i
ie
f
df
r
o
m Be
c
k
e
ra
nd o
t
he
r
s (
1
9
89
)
.
Epi
s
o
de
s
,
Vol
.
1
6
,
n
o.
4
阿 E D I TE RR A NE AN
0 一
.
100 KM
一」
-
434
mi
s
s
i
on on De
vo
ni
a
n St
r
a
t
i
g
r
a
ph
y (
SDS) ha
s gi
ve
n c
a
r
e
f
u
l
c
o
ns
i
d
e
r
-
a
t
i
o
n t
o whi
c
h l
e
v
e
l
i
s
mo
s
t a
pp
r
op
r
i
a
t
e f
o
r i
nt
e
r
na
t
i
on
a
l
c
or
r
e
l
a
t
i
on
a
d
j
a
c
e
nt
t
o t
he s
ou
t
h
e
a
s
t
e
r
n b
or
d
e
r of
t
h
e di
s
us
e
d u
pp
e
r ma
r
b
l
e q
ua
r
r
y
a
n
d i
t
d
e
c
i
d
e
d,
a
t a me
e
t
i
n
g i
n Was
h
i
n
gt
o
n i
n 1
98
9,
t
h
a
t a GSSP
1.
5 km NI
E o
f Ce
s
s
e
n
on vi
l
l
a
ge a
nd 2 1
00 m SW of Ca
u
s
s
e
s e
t
Ve
y
r
a
n.
I
t
c
a
n be r
e
a
c
h
e
d e
as
i
l
y by a pa
t
h up t
h
e hi
l
l
f
r
o
m ne
a
r t
h
e t
r
a
c
k t
o Le
s
Gr
a
n
ge
s f
a
r
mh
ou
s
e f
r
o
m t
he r
oa
d D1
3
6 be
t
we
e
n Ces
s
e
no
n a
nd St
Na
z
a
i
r
e
-
de
-
La
d
a
r
e
z
.
The g
r
ou
nd i
s o
wn
e
d by t
he c
o
mmun
e of
Ces
s
e
no
n a
n
d i
s a
l
r
e
a
d
y p
r
ot
e
c
t
e
d a
s pa
r
t
o
f a wa
t
e
r s
u
pp
l
y a
r
e
a
.
Con
-
s
e
r
va
t
i
on a
nd p
r
o
t
e
c
t
i
on of
t
he s
e
c
t
i
o
n h
a
s be
e
n a
s
s
ur
e
d by c
ommu
na
l
a
n
d d
e
pa
r
t
me
n
t
o
ic
f
i
a
l
s
.
Fr
e
e a
c
c
e
s
s f
o
r s
c
i
e
nt
i
s
t
s i
s
c
on
ir
f
me
d.
The s
e
q
ue
n
ce i
s
on
e o
f pe
l
a
g
i
c c
a
l
c
i
l
u
t
i
t
e
s
,
mos
t
l
y r
e
d t
i
nt
e
d
a
n
d wel
l
b
e
dde
d,
wi
t
h b
e
ddi
ng pr
o
ba
bl
y c
o
nt
r
o
l
l
e
d b
y Mi
l
a
n-
k
ovi
t
c
h-
Ba
nd cl
i
ma
t
i
c os
c
i
l
l
a
t
i
o
ns du
r
i
ng s
e
di
me
nt
a
t
i
on.
The
s
e
qu
e
nc
e ha
s b
ee
n de
s
c
r
i
be
d i
n pu
bl
i
s
he
d a
c
c
o
unt
s (
Ho
us
e a
nd o
t
h-
e
r
s
,
1
985;
Ma
pp
e
r
,
1
9
89;
Be
c
ke
r a
nd o
t
h
e
r
s
,
1
98
9;
Sc
hi
nd
l
e
r 1
9
90;
Be
c
k
e
r 1
99
3a
)
a
n
d i
s
de
p
i
c
t
e
d i
n igu
f
r
e 2
.
The b
ou
nda
r
y i
s dr
a
wn
be
t
we
e
n Be
ds
31
g a
n
d 3
2a a
s s
h
own i
n ig
f
ur
e
s 2 a
nd 3
.
Di
s
t
i
n
c
t
i
v
e i
s
Be
d 3 1
g whi
c
h i
s c
or
r
e
l
a
t
e
d wi
t
h t
he Up
pe
r Ke
l
l
was
s
e
r Li
me
s
t
o
ne ot
Ge
r
ma
ny a
n
d whi
c
h i
s a h
ypo
xi
c d
a
r
k gr
e
y c
a
l
c
i
l
ut
i
t
e t
o c
a
l
c
a
r
e
ni
t
e
a
b
ov
e whi
c
h i
s t
h
e mos
t
ma
r
k
e
d f
a
un
a
l
bo
un
da
r
y.
Bot
h Be
ds 31
g
a
n
d 32
a a
r
e c
ha
r
a
c
t
e
r
i
z
ed by p
e
l
a
g
i
c f
a
un
as
.
The s
e
q
ue
n
ce c
h
os
e
n s
h
ows
a c
o
mpl
e
t
e
s
u
c
c
es
s
i
o
n t
h
r
oug
h t
he
e
a
r
l
y Fr
as
n
i
a
n t
o l
a
t
e Fa
me
n
ni
a
n
.
I
t
i
s un
f
a
ul
t
e
d a
n
d ha
s n
o t
e
c
t
o
n
i
c
pr
obl
e
ms
.
Th
e be
ds
a
r
e a
pp
r
ox
i
ma
t
e
l
y ve
r
t
i
c
a
l
.
Equ
i
v
a
l
e
nt
s
e
c
t
i
ons
c
a
n be f
o
un
d e
l
s
e
whe
r
e i
n t
he a
r
e
a.
Th
e r
oc
ks
a
r
e of l
ow-
gr
a
de met
a
-
mo
r
ph
i
s
m a
nd t
he
r
ma
l
ma
t
u
r
i
t
y (
CAI 2
-
3)
a
n
d c
ompr
i
s
e a
n h
omo
ge
-
n
e
ou
s pe
l
a
gi
c c
a
l
c
i
l
ut
i
t
e s
e
q
ue
n
c
e wi
t
hou
t ma
r
l
y o
r s
ha
l
y i
nt
e
r
be
d
s
.
The
r
e i
s
a c
omp
l
e
t
e z
on
a
l s
uc
c
e
s
s
i
o
n wi
t
h a r
i
c
h f
os
s
i
l
c
o
nt
e
nt
,
e
s
pe
-
c
i
a
l
l
y o
f t
h
e bi
o
s
t
r
a
t
i
gr
a
p
hi
c
al
l
y s
i
gn
i
ic
f
a
n
t g
r
oup
s o
f c
ono
do
nt
s
,
(
UQ)
o
f Cou
r
ni
a
c
,
1
75 i
n WSW o
f t
h
e Le
s Gr
a
n
ge
s f
a
r
mho
us
e
,
a
bo
ut
s
ho
ul
d be s
ou
gh
t i
n r
el
a
t
i
on t
o t
he b
as
e of t
he Lo
we
r t
r
i
ang
ul
ar
i
s
c
o
no
do
nt
Zo
ne
.
Fi
n
a
l
b
a
l
l
ot
s a
n
d r
a
t
i
ic
f
a
t
i
on by I
CS a
nd I
UGS (
i
n
J
a
nua
r
y 1
993
)
l
ed t
o a l
e
ve
l
i
n a s
e
c
t
i
on a
t
Co
ur
n
i
a
c,
s
ou
t
h
e
r
n Fr
a
nc
e
b
e
i
n
g d
es
i
gna
t
e
d a
s t
h
e Gl
o
ba
l
St
r
a
t
o
t
y
pe Se
c
t
i
on a
nd Poi
nt
(
GSSP)
.
A br
i
ef
r
ev
i
e
w of t
he d
oc
ume
n
t
a
t
i
o
n l
e
a
di
ng t
o t
hi
s d
e
c
i
s
i
o
n i
s gi
ve
n
h
e
r
e.
The c
o
no
do
nt
a
nd g
on
i
a
t
i
t
e bi
o
s
t
r
a
t
i
gr
a
p
hi
c d
i
v
i
s
i
o
ns a
r
e
s
ho
wn i
n ig
f
ur
e
s 5 a
nd 6.
I
t
ha
s be
e
n r
e
c
og
n
i
s
e
d by t
he Su
bc
o
mmi
s
s
i
on t
ha
t
,
i
n g
en
e
r
a
l
,
Dev
o
ni
a
n s
e
c
t
i
on
s i
n pe
l
a
gi
c r
ea
l
m f
a
c
i
e
s a
r
e mo
r
e l
i
ke
l
y t
o be c
om-
pl
e
t
e t
ha
n t
ho
s
e i
n t
he n
e
r
i
t
i
c f
a
,
c
i
e
s
.
Th
e pe
l
a
gi
c f
a
c
i
e
s f
or
ms a be
t
t
e
r
ba
s
i
s f
o
r t
h
e bi
os
t
r
a
t
i
gr
a
p
hi
ca
l
p
r
e
c
i
s
i
o
n n
e
e
de
d f
or i
nt
e
r
na
t
i
o
na
l
c
or
-
r
e
l
a
t
i
on.
I
n pa
r
t
i
c
ul
a
r t
he c
o
no
do
nt
an
d g
on
i
a
t
i
t
e r
e
c
or
d
s a
r
e be
t
t
e
r i
n
t
h
os
e f
a
c
i
e
s
.
Tha
t
i
s no
t t
o i
mpl
y t
ha
t
t
he
r
e a
r
e n
ot
f
a
c
i
e
s r
i
c
h i
n ot
he
r
gr
ou
ps
,
f
o
r e
xa
mpl
e s
por
e
s a
n
d br
a
c
hi
o
po
ds
,
wh
i
c
h a
r
e v
e
r
y i
mp
or
-
t
a
nt
f
or
c
or
r
e
l
a
t
i
on
,
bu
t
i
t i
s n
or
ma
l
l
y e
a
s
i
e
r t
o c
o
r
r
e
l
a
t
e i
nt
o s
uc
h s
e
c
-
t
i
o
ns s
e
c
on
da
r
i
l
y f
r
om pr
i
ma
r
y s
e
c
t
i
o
ns i
n t
he p
e
l
a
g
i
c f
a
c
i
e
s
.
I
n t
he
C
a t e
-
e
l( a
a
‘
0了 价以
?
l
ast r
es
or
t t
he Subcommi
ss
i
on concent
r
at
ed on t
wo s
uch s
ect
i
ons,
one
at St
ei
nbr
uch Schmi
dt i
n t
he Rheni
s
h Sl
at
e M ount
ai
ns
,Ger
many
V
。理
仁
select
ed because of t
he bet
t
er docum ent
ati
on
汀
other at (
Sa
n
db
e
r
g a
n
d o
t
h
e
r
s
,
1
99
0;
Sc
h
i
n
dl
e
r
,
I
Co
ur
n
i
a
c i
n t
h
e Mo
n
t
a
g
ue No
i
r
e (
Fe
i
s
t
,
1
9
90 The l
a
t
t
e
r wa
s i
f
na
l
l
y
ssilgr
oups.
Rec
omme
nde
d s
t
r
at
ot
ype
Th
e
r
e
c
o
mme
n
d
e
d b
o
u
n
d
a
r
y
G
S
S
P b
e
t
we
e
n
t
h
e
F
r
a
s
n
i
a
n
a
n
d
Fa
me
n
n
i
a
n St
a
g
e
s (
De
v
o
ni
a
n)
i
s a
bo
ve t
he Up
pe
r Co
ur
n
i
a
c Qua
r
r
y
,
n
e
a
r Ce
s
s
e
n
on
,
Mo
n
t
a
g
ue No
i
r
e
,
Fr
a
n
c
e (
ig
f
ur
e 1
)
.
Th
e s
e
c
t
i
o
n i
s s
i
t
-
u
a
t
e
d i
n t
h
e s
o
ut
he
a
s
t
e
r
n Mon
t
a
gn
e Noi
r
e
,
D6
pa
r
t
me
n
t 1
-
1
6
r
a
u
l
t
,
Di
s
-
t
r
i
c
t
o
f Ce
s
s
e
no
n (
t
op
og
r
a
p
hi
c s
he
e
t 1:
2
5 00
0,
No.
25
4
4 E,
Mu
r
v
i
e
l
-
1
}s-
1
36ziers;
La
mb
e
r
t
'
sc
oo
r
d
i
n
a
t
e
s
:
x=1
30 37
5,
y=6
58 5
5
)
.
I
t
i
s
ammonoi
ds
, t
r
i
l
obi
t
es, t
ent
acul
i
t
es and os
t
r
acods.
Det
ai
l
e
d docu-
me
n
t
a
t
i
o
n ha
s be
e
n pr
o
vi
de
d o
f t
hi
s (
Fe
i
s
t
,
1
9
90)
.
Ge
o
c
he
mi
ca
l
wor
k a
c
r
os
s t
h
e bo
un
da
r
y a
t
Cou
r
n
i
a
c h
a
s a
l
s
o b
e
en p
ub
l
i
s
he
d
(
Goo
df
e
l
l
ow a
n
d ot
he
r
s
,
1
98
9;
Gr
a
nd
j
e
a
n a
nd o
t
h
e
r
s
,
1
9
89;
Gr
a
n
d-
j
e
a
n-
L6
c
uy
e
r a
n
d ot
he
r
s
,
1
993
;
J
oa
c
h
i
ms
k
i
a
nd Bugg
i
s
c
h,
1
9
93;
Gi
r
a
r
d a
nd o
t
he
r
s
,
1
993
) a
nd c
ur
r
e
nt
l
y ot
he
r i
n
ve
s
t
i
ga
t
i
o
ns a
r
e b
e
i
ng
NV-NN山乏哎比
ZV一
NS喊比匡
l
e
d
Fi
gur
e 2 Detai
ma
p o
f t
he之
加per
Qua
r
r
y a
t Cour
ni
ac
,
ne
ar Ce
s
s
e
no
n,
s
howi
ng t
he b
e
d
numb
e
r
i
ng an
d t
he
po
s
i
t
i
o
n o
ft
he GSSP
d
e
ini
f
n
g t
he ba
s
e o
f
I
I
I
4
-
r
!
1
-
1
}
邢1
4
0
3
.
6
,
J
7:
口 !
3
1
3
州日 .
1
3
4
1
二
t
he Famenni
an
bet
wee
n Beds 31
Pl
an
and 3
2.
Mod
i
ie
f
d
户o
m Ho
us
e an
d
o
t
he
r
s (
1
98
5)
.
12
SCA LE
O F
M E TRES
(
1
6)
F仓ur
e 3 The
F
R
A
S
N
I
A
N}
F
A
ME
N
N
I
A
N
det
ai
l
ed success
i
on
1
.
0 METRE
0.
5
o
f be
d
s a
r
ou
nd t
he
GSSP l
e
v
e
l be
t
we
e
n
be
d
s Mg a
nd 3
2a
ab
ov
et
he Uppe
r
Co
umi
ac Qua
ny
臼习
6
1
6
‘
卿 {
‘
队{
泞、护
酬川
ne
ar Ces
senon.
1
4
-
1
4
r
1
-
3
579-.
f
尹、了
1
W
7
刊?
30
15 16
3
d
,
1
e
ele
12 -9
21 -
.
91
61
51
9
d
3
护7
i
e9
,
1
8 3
32
61
+
'
6
C t d 1二
33
期 go
CM
f
BED
December 1993
435
i
s a k
e
y s
eq
ue
n
c
e f
or t
he t
hi
r
t
e
e
n
-
f
ol
d Mo
nt
a
gn
e No
i
r
e Fr
a
s
ma
n
un
de
r
t
a
k
e
n.
Ma
gn
e
t
os
t
r
a
t
i
g
r
a
phi
c wo
r
k i
n
di
c
a
t
e
s t
ha
t
t
h
e ar
ea wa
s
r
ema
g
ne
t
i
z
e
d d
ur
i
ng t
he Pe
r
mi
a
n.
z
c
o
a
t
n
e
a
d
t
i
o
a
t
n
v
(
a
M
r
i
a
o
p
u
p
s
e
s
r
e
c
1
9
t
i
8
o
9
n
)
s
,
i
s
n
u
b
N
s
o
t
a
r
n
t
h
t
i
a
A
l
m
p
a
e
r
t
i
s
c
a
o
,
f
W
w
h
e
i
s
c
t
h
e
r
h
n
a
A
v
e
u
s
b
t
e
r
a
e
l
n
i
a
r
,
e
p
a
n
l
i
d
-
E
a
t
u
U
r
o
p
p
e
a
r
n
C
R
o
u
s
r
s
n
i
a
c
(
M
e
x
a
t
p
e
p
n
e
d
r
s
a
f
n
r
o
d
m
F
o
F
s
r
t
a
e
s
r
m
1
a
9
n
9
3
Z
,
o
i
f
n
g
e
u
5
r
e
t
o
2
)
t
.
h
T
e
h
t
o
e
p
s
e
o
q
f
u
Z
e
n
o
n
c
e
1
3(
=t
op of t
he Fr
a
s
ni
a
n)
,
b
ut
on
l
y t
h
e hi
gh
e
r pa
r
t
o
f t
h
e Fr
a
s
ma
n
Corr
e
l
at
i
on of t
he pr
opo
s
e
d bo
undar
y
s
eauence i
s of conce
rn her
e.
l
evel
Bed 24a,
t
he Lower Ke
l
l
wass
er Li
mes
t
one, nas me i
owes
t
o
c
cu
r
r
e
nc
e of An
c
y
r
o
gn
at
hu
s as
y
mme
t
r
i
c
u
s,
whi
c
h i
s t
he d
e
ini
f
ng
s
pe
c
i
e
s f
or t
h
e l
owe
r bo
und
a
r
y o
f t
he Upp
e
r g
i
g
as Zo
ne i
n t
h
e z
o
na
-
t
i
on of Zi
e
gl
e
r (
1
96
2,
p.
23;
1
9
7 1
)
.
The c
o
no
don
t
f
a
una of Be
d 2
4a i
s
i
n
t
e
r
p
r
e
t
e
d a
s t
he h
i
g
he
s
t
s
a
mpl
e
d l
e
ve
l
o
f Mon
t
a
gu
e No
i
r
e Zo
ne 1
2
Th
e
b
o
u
n
d
a
r
y
l
e
v
e
l
p
r
o
p
o
s
e
d
r
e
p
r
e
s
e
n
t
s
p
e
r
h
a
p
s
t
h
e
b
e
s
t
c
o
r
r
e
l
a
t
e
d
hori
zon i
n t
he Devoni
an.
A r
evi
ew of mor
e t
han 30 i
nt
er
nat
i
onal see-
t
i
ons ha
s be
e
n p
r
es
e
nt
e
d b
y Sa
nd
be
r
g a
n
d o
t
he
r
s (
1
98
8
) i
nc
l
ud
i
n
g
l
oc
a
l
i
t
i
e
s i
n Nor
t
h Ame
r
i
ca a
n
d Eu
r
op
e.
Fu
r
t
h
e
r c
o
r
r
e
l
a
t
i
on i
s es
t
a
b-
l
i
s
he
d i
n No
r
t
h Af
r
i
ca (
Be
c
ke
r a
nd ot
he
r
s 1
988
)
,
Chi
n
a (
J
i
,
1
989
)
a
n
d Au
s
t
r
a
l
i
a (
Be
c
ke
r a
nd o
t
he
r
s
,
1
991
)
.
Th
e bou
nda
r
y c
or
r
e
s
pon
ds
t
o t
he e
xt
i
nc
t
i
o
n of a
l
l
s
p
e
c
i
e
s of t
he c
on
od
on
t
s An
c
yr
o
de
l
l
a a
nd
Oz
ar
k
o
di
n
a a
nd t
he l
o
s
s of a
l
l
but
a f
e
w s
pe
c
i
e
s
of Pa
l
mat
o
l
e
pi
s
,
Po
l
y
gna
t
hu
s
,
a
nd Anc
yr
og
nat
hu
s
,
a
cc
o
r
di
ng t
o Sa
nd
be
r
g a
n
d ot
he
r
s
(
Ma
pp
e
r
,
1
98
9,
p.
45
6,
ig
f
ur
e 4)
.
Ho
we
ve
r
,
du
e t
o t
h
e f
a
c
i
e
s
o
f t
he
Lowe
r Ke
l
l
wa
s
s
e
r a
t
Co
ur
n
i
a
c
,
wh
i
c
h a
ppa
r
en
t
l
y r
e
s
ul
t
e
d i
n t
he
e
x
t
r
e
me r
a
r
i
t
y o
f Pal
ma
t
ol
e
pi
s i
n Be
d 2
4a
,
a hi
ghe
r z
o
na
l
i
d
en
t
i
ic
f
a
-
t
i
o
n c
a
n
no
t b
e e
xc
l
ud
ed
.
No
ne
t
he
l
es
s
,
t
he l
owe
r bo
un
da
r
y of Zon
e
13 i
s n
e
c
es
s
a
r
i
l
y t
a
ke
n a
t t
he l
owes
t
oc
c
u
r
r
e
nc
e o
fP
a
I
n
i
at
ol
e
pi
s b
og-
ar
t
e
ns
i
s i
n Be
d 24
e (
ig
f
ur
e 5)
,
a
s we
l
l
a
s a
t
t
he c
oi
nc
i
de
nt
l
owe
s
t
(
1
98
8
,
p
p.
2
93
-2
94
)
.
The
r
e i
s a we
l
l
-
k
no
wn e
x
t
i
nc
t
i
on a
mo
ng g
o
m-
oc
cur
r
ence of Pa.
has
si s.
s.
The l
owest occur
r
ence of Pa.
rhenana
a
t
i
t
e
s o
f t
h
e Ge
phu
r
oc
e
r
a
t
i
da
e a
nd Be
l
o
c
e
r
a
t
i
d
a
e a
n
d t
he r
e
co
r
d f
or
bo
t
h c
o
nod
on
t
s
a
n
d go
ni
a
t
i
t
e
s a
t
Co
ur
n
i
a
c de
mo
ns
t
r
a
t
es t
h
i
s wel
l
.
Th
e l
a
s
t of t
he b
r
a
c
h
i
o
po
d At
r
y
pi
d
a
e oc
c
u
r
s
j
u
s
t
be
l
ow t
he bou
nd
a
r
y
l
e
ve
l
(
Be
ck
e
r a
n
d o
t
he
r
s
,
1
991
)
.
Amon
g t
r
i
l
obi
t
e
s t
he Da
l
ma
n
i
t
i
da
e
,
O
t
h
d
e
o
b
n
a
t
o
s
e
p
l
o
e
f
u
r
t
i
h
d
e
a
e
,
n
H
d
-
a
F
r
r
p
a
e
s
t
n
i
d
i
a
n
e
U
a
n
p
d
p
e
A
r
u
K
l
a
e
c
l
l
o
w
p
l
a
e
s
u
s
r
e
i
r
n
L
a
e
i
m
a
e
l
l
s
t
d
o
i
n
s
e
a
p
l
p
e
e
v
a
e
r
l
o
a
f
t
Be
d 31
g.
Ot
he
r
s ha
v
e doc
ume
nt
e
d t
he g
l
o
ba
l
e
xt
i
nc
t
i
o
n o
f c
or
a
l
a
卜
1
nd A at
(
Sor
a
uf a
n
d Pe
d
de
r
,
1
98
6;
Sc
r
u
t
t
o
n,
1
988
)
,
s
t
r
omat
op
o
r
o
i
d (
St
ea
r
n
,
1
98
7)
a
nd a
c
r
i
t
a
r
c
h (
Va
ng
u
es
t
a
i
ne a
n
d ot
h
e
r
s
,
1
98
3)
g
r
o
ups
t
he
r
e ha
s b
e
e
n muc
h r
e
c
e
nt
l
oc
a
l
do
c
ume
nt
a
t
i
on i
n ma
n
y a
r
e
a
s
.
he bounda
ry
c
ha
n
ge
ov
e
r of b
en
t
hon
i
c os
t
r
a
c
o
d f
a
un
a
s a
c
r
o
s
st
Cou
mi
a
c ha
s b
e
e
n pu
bl
i
s
he
d (
Let
hi
e
r
s a
nd Fe
i
s
t
,
1
991
)
.
s
e
n
s
u Ma
p
pe
r a
nd Fos
t
e
r (
1
99
3,
p.
24
,
ig
f
ur
e 2)
i
s
hi
gh wi
t
h
i
n Zon
e
1
3 i
n Be
d 3
0b,
c
o
i
nc
i
de
n
t
wi
t
h t
h
e l
owe
s
t
Pa.
bo
o
gaa
r
di
(
ig
f
ur
e 5
)
.
I
t i
s n
ot
e
wo
r
t
hy t
ha
t
s
pe
c
i
es
c
ha
r
a
c
t
e
r
i
s
t
i
c o
f t
he hi
gh
e
r pa
r
t
o
f
t
h
e Fr
a
s
ni
a
n a
t
Up
pe
r Cou
mi
a
c
,
Pal
mat
ol
e
pi
s b
og
ar
t
e
n
s
i
s
,
Pa.
wi
n
ch
e
l
l
i
,
Pa.
r
he
nan
a
,
Pa
.
bo
og
aar
di
,
Anc
yr
o
gna
t
h
u
s a
s
y
mme
t
r
i
-
cu
s
,
a
n
dA
n
c
y
r
o
de
l
l
a cu
r
v
a
t
a a
l
l
t
e
r
mi
na
t
e i
n e
i
t
h
e
r Be
ds 31
f or 31
g.
The c
o
no
do
nt
f
a
una of Be
d 31
g,
t
h
e Up
pe
r Ke
l
l
wa
s
s
e
r Li
me
s
t
o
ne
,
ha
s a
l
l
b
ut
t
wo of t
he f
or
e
go
i
n
g s
p
e
c
i
e
s (
igu
f
r
e 5
)
a
n
d i
s do
mi
n
a
t
e
d
i
n t
e
r
ms o
f i
t
s Pal
mat
ol
e
pi
s c
omp
on
e
nt
,
by Pa
.
bo
g
ar
t
e
ns
i
s a
n
d Pa.
wi
nc
he
l
l
i
.
On e
v
i
d
en
c
e of t
h
e l
o
we
s
t
(
a
nd o
n
l
y)
o
c
c
ur
r
e
n
c
e o
f An
c
y-
r
og
nat
hu
s u
bi
qu
i
t
us i
n Be
d 31
g,
t
he f
a
u
na i
s a
p
pa
r
e
nt
l
y c
o
r
r
el
a
t
i
v
e
wi
t
h t
he up
pe
r pa
r
t
o
f t
he l
i
ngu
扣r
mi
s Zo
ne (
Zi
e
gl
e
r a
n
d Sa
nd
be
r
g,
1
9
90
,
p.
21
)
,
de
s
pi
t
e t
h
e a
bs
e
nc
e of Pa.
l
i
n
gu扣r
mi
s
.
I
n t
h
e Mon-
Conodont record
Th
e
c
o
n
o
d
o
n
t
s
e
q
u
e
n
c
e
a
t
t
h
e
U
p
p
e
r
C
o
u
mi
a
c
Qu
a
r
r
y e
x
t
e
n
d
s
f
r
o
m
wi
t
h
i
n t
he mi
dd
l
e pa
r
t
of t
he Fr
a
s
n
i
a
n a
c
r
os
s
t
h
e Fr
a
s
n
i
a
n
/
Fa
me
nn
-
i
a
n bo
un
da
r
y a
n
d i
n
t
o t
h
e l
owe
r Fa
men
ni
a
n (
a
s hi
g
h a
s t
he Upp
e
r
c
r
e
pi
d
a Zo
ne i
n t
he c
ur
r
e
n
t s
a
mpl
i
ng up t
o Be
d 44)
.
Upp
e
r Co
unua
c
Fi
gur
e 4
Pho
t
o
gr
a
ph o
f t
he
t
a
g
ne Noi
r
e Fr
as
n
i
a
n z
ona
t
i
on,
Be
d 3 1
g
r
e
pr
e
s
e
n
t
s t
he h
i
g
he
s
t p
a
r
t
of Zo
ne 1
3
.
Anc
yr
og
nat
hu
s uI
nq
ui
t
u
s a
l
s
o o
c
c
ur
s i
n f
a
u
na
s o
f t
h
e
Up
pe
r Ke
l
l
wa
s
s
e
r Li
mes
t
o
ne a
t t
h
r
e
e o
t
h
e
r ne
a
r
by Mon
t
a
g
ne No
i
r
e
l
o
ca
l
i
t
i
e
s
:
Ca
us
s
e
s e
t
Ve
yr
a
n No
r
t
h a
n
d Sou
t
h
,
a
n
d Lo
we
r Cou
mi
a
c
(
Fe
i
s
t
,
1
99
0,
p
p.
1
9,
2
4,
30
)
.
Al
l t
he
s
e f
a
u
na
s a
r
e c
or
r
e
l
a
t
i
ve wi
t
h t
he
l
i
ngu
i
f
or
mi
s Zon
e
,
b
ut
l
a
c
k t
h
e n
omi
na
l
s
pe
c
i
e
s
(
Bec
ke
r a
nd o
t
h
e
r
s
,
1
98
9,
pp.
26
2,
26
5)
.
黔 --
succes
si
on above
煞
t
he Up
pe
r Cou
mi
ac
Quar
r
y.
The GSSP
71
,
翼
噢
l
i
e
s bet
ween Bed
31
g and 3
2a.
For
s
c
al
e c
o
mpa
r
e wi
t
h
t
he pr
o
il
f
e o
f t
he
be
ds gi
v
e
n i
n
igur
f
e 3.
纂墓
雌夔瓣毅井 价
鬓
粱省
岁彩孚擎
七万黔 像
一
巍砂色_
坛
32a
瓣嚎
脚
了I‘
了 I C
了 弃a
Epi
sodes,
Vol
.
I反 no.
4
盗
了峪
尸... .月,,.网 ..尸一~一 一一一~-, 一.
—
—
—
不
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一
4了6
s
pe
ci
es across t
he
Frasni
anlFamenni
n
b
ound
ar
y at Uppe
r
Co
umi
a
c Qua
r
r
y.
The GSSPfo
r t
he
b
as
e o
ft
he
LI
DW
MIDDLE COUMIAC FORMATI
ON
Fi
gur
e 5 Range
c
har
t o
f c
o
nod
ont
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2日
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271 2习
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b 1
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Fam enni
an i
s at t
he
SP indet
b
as
e o
f Be
d3
2
a.
Po
s
i
t
i
on o
f t
he
u
p
pe
rpa
r
t
o
f
如月二
eUQ己
p0 u吧b七谈
尘
Fr
as
ni
an Zone 12,
Fr
as
ni
an Zone 13
犯 比d U二d妞ern口健
口舀
I.
sy}
.
.
.
b它1巧
.
(
Ma
ppe
r
,
1
98
9)
,
t
he
Rl hQS舀t
Lo
we
r
,
Mi
ddl
e and
Up
pe
r t
r
i
ang
ul
a
r
i
s
Zo
ne
s
,
and t
he
皿」
丁
」七.d酬刀m s创曰
An.几笼白SQ
.. 口
I七. ef.尸匕.breuL
s
Lower and Mi
ddl
e
...
Pa. LVW 沦 迸 f二Pa..
曲 注匕位
c
r
ep
i
da Zone
s i
s
i
ndi
cat
ed.
Re
c
ogni
t
i
on o
f t
he
Lowe
r
,
Mi
dd
l
e an
d
功少e
r c
r
e
pi
da
R L cE P. r
h-
...
Mehl臼U
Pa rh己n口尸习1
召l
m
}
Zones i
s ci
t
ed i
n t
he
t
e
xt
,
bu
t s
pe
c
i
e
s
}
p匕 气夕
r
anges are not
shown above Bed
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r
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e
p
h
, k
s
p
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a
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t
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e
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r
red ran
ge An- RANGES OF CONODONT TAXA AT THE UPPER COUMI
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Up
p
e
r
C
o
u
t
r
u
a
c
B
e
d
3
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g
i
s
a
l
s
o n
o
t
e
wo
r
t
h
y
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i
n
h
a
y
i
n
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c
o
n
i
f
-
d
e
n
t
l
y i
d
e
n
t
i
f
i
e
d_
s
p
e
c
i
me
n o
ta
l
n
i
a
t
o
‘
e
p
‘
s
t
ra
n
g
u
t
仔
n
s冬
a
n
d o
些
q
u
e
s
t
i
o
n
a
b
l
e s
p
e
c
i
me
n o
f
t
h
e
s
a
me s
p
e
c
i
e
s
,
呼I
g
r
me
r
u
s
t
e
u i
n月臀L
,
1
9
9
0
,
p
.
3
6
)
a
mo
n
g t
h
e ma
n
y e
x
a
mpe
s
o
t
P
a
.o
g
a
r
r
e
n
s
‘
s
.
a
D
a r
a
.
wi
n
c
h
e
l
l
i
.
Al
t
h
o
u
g
h o
n
e
c
a
n
n
o
t
c
o
mp
l
e
t
e
坚e
x
c
,
u
g
e
ne
t
p
o
s
s
i
甲l
i
y。
‘
s
t
r
a
t
i
g
r
a
p
h
i
c
a
l
l
e
a
k f
r
o
m t
h
e o
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l
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n
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a
n
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t
r
e
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a
r
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c
c
u
r
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n
c
e
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o
f
P
a
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t
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e
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t
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a
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l
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r
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n t
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p
p
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r
e p
r
o
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a
b
l
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o
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u
n
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o t
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p
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r
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u
r
ma
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i
t
e
r
e
p
o
r
t
o
t
2
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n
i
f
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u
r
e
d s
p
e
c
i
me
n
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i
呼
e
n
t
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e
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s塑.
pr
a
e
t
n
a
n
g
u
t
a
ns
f
r
o
m t
h
r
s
a
mp
l
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t
h
i
n Be
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t
te
_
U
p
p
e
r
Kewa
s
s
e
r
L
i
me
s
‘
o
n
e
a Se
‘
n
-
b
r
u
c
h S
c
h
mi
d
t
(
S
a
n
d
b
e
r
g a
n
d o
te
r
s
,‘
丫
匕
匕
,乡
a
b
‘
e止!ma
y r
e
p
r
e
anot
her Fras
man occ
ur
r
ence o1尸a.
t
rangul
ars.
ini
s i
nt
er
ence i
s
ba
s
e
d on Ma
p
pe
r
'
s s
t
u
d
y i
n 1
9
90 of
t
he h
ol
ot
y
pe a
n
d t
wo p
a
r
a
t
yp
e
s of
Pa
.
pr
ae
t
r
i
a
ng
ul
ar
i
s Zi
e
g
l
e
r a
nd Sa
nd
b
e
r
g (
i
n Sa
n
d
be
r
g a
nd o
t
h
e
r
s
,
1
98
8
,
p.
3
0
4,
pl
.
1
,
ig
f
u
r
e
s 1
,
3
,
4)
,
a
l
l
f
r
om t
he hi
g
he
s
t be
d of t
he l
i
n
-
gu扣r
i
ni
s Zon
e a
t Ha
ma
r La
g
hda
d,
s
o
ut
he
r
n Mo
r
o
c
c
o.
The
s
e t
h
r
e
e
t
yp
e
s (
Pa e
l
e
me
n
t
s
) ha
v
e a
n a
r
c
h
e
d o
ut
e
r
-
p
os
t
e
no
r p
l
a
t
f
o
r
m,
whi
c
h
g
e
nt
l
y r
i
s
e
s
f
r
o
m t
he l
o
be t
oj
us
t
be
f
o
r
e t
he t
i
p,
t
he
n a
r
c
h
e
s d
own
wa
r
d.
Thi
s i
s e
x
a
c
t
l
y t
he s
a
me a
s i
n ma
n
y s
p
e
c
i
me
n
s of Pa.
t
r
i
a
ngu
l
ar
i
s i
n
t
h
e Lowe
r a
n
d Mi
d
dl
e t
r
i
an
gu
l
ar
i
s Zo
ne
s
;
ho
we
v
e
r
,
i
n t
he hol
o
t
y
pe
(
Sa
n
ne
ma
n
n,
1
955
,
pl
.
2
4,
ig
f
u
r
e 3
)
t
h
e o
ut
e
r pos
t
e
r
i
o
r pl
a
t
f
o
r
m r
i
s
e
s
mo
r
e
s
t
e
e
p
l
y f
r
o
m t
h
e l
o
b
e t
o j
u
s
t
b
e
f
o
r
e t
h
e t
i
p
,
t
n
e
n_
a
r
c
n
e
s
u
o
wn
-
wa
r
d
.
Co
n
s
e
q
u
e
n
t
l
y
,
t
h
e v
i
e
w i
s
f
a
v
o
u
r
e
d h
e
r
e
t
h
a
t
F
a
l
ma
t
o
吧S
t
r-
an
gu
l
a
r
i
s a
nd Pa.
pr
ae
t
t
i
a
ngu
l
ai
l
s a
r
e s
y
no
ny
ms
.
A s
l
i
g
ht
l
y di
f
f
e
r
e
n
t
v
i
e
w pe
r
ha
ps c
ou
l
d b
e s
upp
or
t
e
d i
n whi
c
h t
he l
a
t
t
e
r i
s t
r
e
a
t
e
d as an
i
n
t
r
a
s
pe
c
i
ic f
mo
r
ph
ot
ype
,
wh
i
c
h i
s a
ppa
r
e
n
t
l
y a c
ommo
n f
or
m o
f
Pa
.
t
r
i
an
gul
ar
i
s i
n t
he Lowe
r t
r
i
an
gul
ar
i
s Zo
ne
.
For
ms l
i
k
e t
he
hol
o
t
yp
e o
f Sa
n
ne
ma
nn s
e
em t
o be r
e
l
a
t
i
ve
l
y r
ar
e i
n t
hi
s z
on
e a
t
l
e
a
s
t
i
n t
he Mo
nt
a
gn
e No
i
r
e.
Cl
e
a
r
l
y t
he pr
ae
t
r
i
an
gul
ar
i
s mor
ph
o-
t
yp
e c
r
os
s
e
s t
h
e Fr
a
s
n
i
a
n
/
Fa
men
ni
a
n bo
unda
r
y a
t
Sc
hmi
dt
Qua
r
r
y,
a
s wa
s we
l
l
doc
u
me
nt
e
d by Sa
n
db
e
r
g a
nd ot
h
e
r
s (
1
9
88,
Ta
bl
e 1
)
.
[
The 27 s
p
e
c
i
me
ns l
i
s
t
e
d a
s Pa.
pr
ae
t
r
i
an
gul
ans i
n Be
d 1
6 a
t
Sc
hmi
dt
Qua
r
r
y oc
c
ur
wi
t
h 33
80 l
i
s
t
e
d a
s Pa.
s
ub
r
e
c
t
a (
a j
u
ni
o
r
s
yno
ny
m o
f Pa.
wi
nc
he
l
l
i
,
s
e
e Ma
pp
e
r a
n
d Fos
t
e
r
,
1
9
93,
bu
t s
p
e
ci
-
me
ns o
f Pa.
b
o
gar
t
e
ns
i
s ma
y ha
ve be
e
n i
n
cl
ud
e
d i
n t
hi
s
c
o
un
t
)
]
.
Th
us
,
e
xt
r
e
me
l
y r
a
r
e Pa.
t
r
i
angu
l
ar
i
s as t
hi
s s
p
e
c
i
e
s i
s d
e
l
i
mi
t
e
d
he
r
e ma
y oc
c
ur s
l
i
g
ht
l
y be
l
o
w t
he l
e
ve
l
of t
he l
o
we
r bo
un
da
r
y of t
h
e
Lo
we
r t
r
i
a
ngu
l
ar
i
s
Zon
e a
t bo
t
h Cou
mi
a
c a
n
d Sc
hmi
dt
q
ua
r
r
i
es (
up
t
o 1
2 a
nd 1
0 c
m be
l
ow t
h
e bo
un
da
r
y,
r
es
p
ec
t
i
v
el
y)
.
I
n bo
t
h
i
ns
t
a
n
ce
s
,
t
hi
s
i
s wi
t
h
i
n t
he co
nt
e
xt
o
f t
he domi
n
a
nt
u
ppe
r Fr
a
s
ni
a
n
f
a
u
na of t
he Upp
e
r Ke
l
l
was
s
e
r Li
mes
t
o
ne
.
Pal
mat
ol
e
pt
s t
nang
u-
l
ar
i
s do
es no
t
oc
c
u
r i
n t
h
e be
ds
e
q
ui
va
l
e
n
t t
o Be
d 3 1
g
a
t
t
h
e o
t
he
r
Mon
t
a
g
ue No
i
r
e l
oc
a
l
i
t
i
e
s c
i
t
e
d p
r
e
v
i
ou
s
l
y.
I
t f
ol
l
ows
,
f
r
o
m t
he f
or
e
goi
ng d
i
s
c
us
s
i
o
n,
t
ha
t
t
he
r
e s
ho
ul
d be
modi
ic
f
a
t
i
o
n of t
he de
f
i
ni
t
i
o
n of t
he l
owe
r bo
unda
r
y of t
h
e Lo
we
r
t
r
i
an
gul
ar
i
s Zon
e (
Zi
e
gl
e
r
,
1
962,
P.
25;
Zi
e
gl
e
r a
nd Sa
nd
be
r
g,
1
9
90,
p.
2
2) f
or
whi
c
h t
h
e s
o
l
e c
r
i
t
e
r
i
on i
s t
he ir
f
s
t
oc
c
u
r
r
e
n
c
e o
ft
h
e
no
mi
na
l
t
a
x
on.
I
ns
t
ea
d i
t
i
s pr
opo
s
e
d he
r
e t
o u
s
e f
o
r de
f
i
ni
t
i
on t
he
D ecember 1993 437
a
b
und
a
nt
o
r lo
f
od o
cc
ur
r
e
nc
e of Pal
ma
t
ol
e
pi
s t
r
i
a
ngu
l
a
r
t
s
,
t
o t
h
e
v
i
r
t
u
a
l
e
xc
l
us
i
on of ot
he
r s
pe
c
i
e
s o
f t
h
e ge
nus
,
s
t
r
a
t
i
g
r
a
phi
c
a
l
l
y
a
bov
e t
he f
au
na do
mi
n
a
t
e
d b
y t
he c
ha
r
a
c
t
e
r
i
s
t
i
c upp
e
r F
r
a
s
ni
a
n
s
pe
c
i
e
s
.
The
s
e i
n
c
l
ud
e Pal
ma
t
ol
e
pi
s wi
nc
he
l
l
i
,
Pa.
b
og
ar
t
e
ns
i
s
,
Pa.
r
h
e
nan
a,
Pa.
boo
gaar
di
,
Anc
yr
o
gn
at
hu
s as
y
mn
i
e
t
r
i
cu
s
,
Anc
y-
r
o
de
l
l
a c
ur
v
at
a (
l
a
t
e f
o
r
m)
,
a
nd i
n s
ome a
r
e
as
,
Pa.
l
i
ngu
i
f
onn
i
s a
nd
Pa
.
i
.
u
n
t
i
ane
n
s
i
s.
Thu
s
,
i
n t
h
e t
hr
e
e s
a
mp
l
i
ng i
nt
e
r
va
l
s
wi
t
hi
n Be
d
32
a a
t
Up
pe
r Co
umi
a
c (
Fe
i
s
t
,
1
99
0,
p.
36
)
a
n
d Be
d 3
2b,
Pa.
t
r
i
an-
gul
ar
i
s i
s
t
h
e on
l
y Pal
mat
ol
e
pi
s s
pe
c
i
e
s wi
t
h t
he e
xc
e
pt
i
o
n of t
r
a
n-
s
i
t
i
o
na
l
s
p
e
ci
me
ns he
r
e t
e
r
me
d Pa.
a
f.
'
de
l
i
c
at
ul
a d
e
l
i
c
al
u
l
a'
.
Re
c
o
gn
i
t
i
o
n o
f t
he Up
pe
r t
r
i
an
gul
ar
i
s Zone (
Be
ds 3
4a
,
3
5a
,
3
6)
,
t
he Lo
we
r c
r
e
pi
da Zon
e (
Be
d 37
,
on t
he u
nq
ue
s
t
i
o
n
ed oc
c
ur
-
r
e
n
c
e o
f t
he n
omi
na
l
s
pe
c
i
e
s
)
,
a
n
d Mi
d
dl
e c
r
e
pi
da Zo
n
e (
Be
ds
Non
e of t
h
e s
pe
c
i
e
s i
n Be
ds 31
a
n
d l
o
we
r a
t
Uppe
r Cou
ni
j
a
c (
ig
f
ur
e
success
i
on f
r
om t
he Mi
ddl
e Fr
as
ni
an t
o t
he M i
ddl
e Famenni
an.
41
-44)
i
s s
t
r
a
i
g
ht
f
o
r
wa
r
d
,
but
no
t
e
s
pe
c
i
a
l
l
y pe
r
t
i
n
e
nt
t
o t
h
e de
s
c
r
i
p-
t
i
on of t
he GSSP.
Goni
at
i
t
e record
Co
u
r
n
i
a
c
i
s
o
n
e
o
f
t
h
e
f
e
w k
n
o
w
n
p
l
a
c
e
s
w
i
t
h。
c
o
n
t
i
n
u
o
u
s
g
o
n
i
a
t
i
t
e
5
) r
a
n
ge
s a
bo
ve t
h
e GSSP pos
i
t
i
o
ne
d a
t
t
h
e l
owe
r b
oun
da
r
y o
f Be
d
3
2a
,
e
xc
e
pt
I
cr
i
o
du
s al
t
e
mat
us
,
whi
c
h i
s we
l
l
kno
wn t
o c
r
os
s t
he
Fr
a
s
ni
a
n
/
Fa
me
nni
a
n bo
und
a
r
y e
l
s
e
wh
e
r
e [
t
h
e de
s
i
gn
a
t
i
on Me
hl
i
n
a
s
pp.
r
e
pr
es
e
nt
s s
e
v
e
r
a
l
s
pe
c
i
e
s whos
e de
ma
r
c
a
t
i
on i
s
u
nc
l
ea
r
]
.
The
n
e
w s
p
ec
i
es
of Anc
yr
og
nat
hu
s i
n Be
d 32
b i
s pa
r
t
o
f t
he Fame
n
ni
a
n
l
i
ne
a
g
e i
n
cl
ud
i
n
g Ag.
s
i
ne
l
ami
nu
s a
nd Ag
.
c
r
y
pt
u
s a
n
d i
s
mo
r
pho
-
l
ogi
c
a
l
l
y di
s
t
i
nc
t
f
r
o
m t
he ma
i
n Fr
a
s
n
i
a
n Anc
y
r
o
gn
a
t
h
us
l
i
ne
a
g
es
t
ha
t
i
n
c
l
ud
e Ag.
as
ymme
t
r
i
cu
s a
nd Ag.
u
bi
qu
i
t
u
s
.
Cha
r
a
c
t
e
r
i
s
t
i
c
u
ppe
r Fr
a
s
ma
n s
pe
c
i
e
s of Pol
yg
nat
hu
s
,
s
uc
h a
s
Po
.
we
bbi
a
nd Po
.
de
c
or
o
s
us
,
d
o no
t r
a
ng
e a
bov
e t
he GSS
P a
n
d a
r
e r
e
pl
a
c
e
d by Po
.
br
e
vi
l
ami
nu
s a
n
d Po
.
n.
s
p.
a
bo
ve t
h
e bou
nda
r
y a
t
Co
umi
a
c
.
Th
e l
o
we
r
b
ou
nd
a
r
y o
ft
he Mi
d
dl
e t
r
i
an
gu
l
a
r
i
s Zon
e
,
de
i
f
n
e
d b
y
t
h
e l
o
we
s
t oc
c
u
r
r
e
nc
e o
f Pal
mat
ol
e
pi
s cl
a
r
k
i
an
d Pa.
de
l
i
c
a
t
u
l
a de
l
i
-
c
at
ul
a (
Zi
e
g
l
e
r
,
1
96
2,
p.
2
6;
1
9
71
,
c
ha
r
t
5)
,
i
s
a
t
t
h
e ba
s
e of Up
pe
r
Cou
mi
a
c Be
d 3
2
c.
Th
e f
o
r
me
r s
p
ec
i
e
s i
s
pe
r
f
e
c
t
l
y u
s
e
a
b
l
e f
or
de
i
f
n-
i
n
g t
h
e b
a
s
e of t
he z
on
e a
n
d t
h
a
t
pa
r
t
of t
h
e o
r
i
g
i
n
a
l
d
ef
i
ni
t
i
on i
s f
o
l
-
l
o
we
d he
r
e.
Bu
t us
e o
f t
he l
a
t
t
e
r t
ax
o
n i
s c
l
o
ude
d by a t
a
x
on
omi
c
pr
o
bl
em.
Th
a
t i
s
,
Pa.
de
l
i
c
a
t
u
l
a de
l
i
c
at
ul
a i
n t
he s
e
ns
e o
f mo
s
t
a
ut
h
o
r
s s
i
nc
e 1
96
2,
a
n
d Pa.
d
e
l
i
c
at
ul
a p
l
a
t
ys o
fZ
i
e
gl
e
r a
nd Sa
nd
be
r
g
(
1
99
0)
,
a
r
e c
h
a
r
a
c
t
e
r
i
z
e
d by p
l
a
t
f
or
m ou
t
l
i
n
e
s of t
he Pa e
l
e
me
n
t
di
s
-
t
i
n
c
t
l
y di
f
f
e
r
e
nt
f
r
om
t
ha
t
o
f t
he l
e
c
t
o
t
y
pe
│
Ammo
no
i
ds oc
c
ur
b
ot
h i
n t
he l
a
t
e
s
t
Fr
a
s
n
i
a
n (
Be
d 3 I
g
)
a
n
db
a
s
a
l
mos
t
Fa
me
n
ni
a
n (
Be
d 3
2a
)
.
P
r
es
e
r
v
a
t
i
o
n,
h
owe
v
e
r
,
i
s i
n ma
n
y c
a
s
e
s r
e
l
a
-
t
i
ve
l
y po
o
r
,
e
x
t
r
a
c
t
i
on t
e
di
o
us
,
a
n
d s
p
e
c
i
e
s
-
l
e
ve
l de
t
e
r
mi
n
a
t
i
o
n i
s
of
te
n di
ic
f
u
l
t
.
Th
e f
a
u
na
l
r
e
c
o
r
d (
ig
f
u
r
e 6)
ha
s g
r
a
du
a
l
l
y be
e
n b
ui
l
t
u
p
ov
e
r t
h
e pa
s
t
s
e
v
e
r
a
l
ye
a
r
s a
nd c
e
r
t
a
i
n
l
y c
a
n be f
u
r
t
he
r e
x
t
e
nd
e
d.
Be
d
-
by
-
be
d i
n
v
e
s
t
i
g
a
t
i
o
ns we
r
e ir
f
s
t
c
on
du
c
t
e
db
y Ho
us
e a
n
d o
t
h
e
r
s (
1
9
85
)
a
n
d u
pd
a
t
e
s
o
f f
a
u
na
l
l
i
s
t
s we
r
e s
u
p
pl
i
e
d i
n c
o
ns
e
c
u
t
i
v
e ye
a
r
s
t
o SI
DS
a
n
d l
a
t
e
r s
u
mma
r
i
s
e
d (
i
n Fe
i
s
t
,
1
99
0)
.
A d
e
t
a
i
l
e
d d
e
s
c
r
i
pt
i
o
n of t
he
Fa
me
nn
i
a
n pa
r
t
i
s pr
o
vi
de
d i
n Be
c
k
e
r (
1
9
93
a
)
.
The b
ou
nda
r
y i
t
s
e
l
f i
s c
l
e
a
r
l
y ma
r
ke
d by
t
he d
i
s
a
pp
e
a
r
a
n
ce of
t
he ge
p
hu
r
oc
e
r
a
t
i
ds a
nd be
l
o
c
e
r
a
t
i
ds wi
t
h t
he i
mme
di
a
t
e s
ubs
e
q
ue
n
t
bl
o
om of t
he t
o
r
noc
e
r
a
t
i
d Ph
oe
ni
x
i
t
e
s f
r
e
c
hi
i
n t
he ha
ema
t
i
t
e
-
e
n
r
i
c
he
d ba
s
e o
f Be
d 32
a,
bu
t
t
he s
p
e
c
i
e
s i
s k
no
wn t
o o
c
c
ur i
n l
a
t
e
Fr
a
s
n
i
a
n b
e
ds e
l
s
e
whe
r
e (
Be
c
ke
r
,
1
993
a
)
.
Thef
r
e
c
hi
(
p
a
r
t
i
a
l
r
a
ng
e
)
Zone or Ph
oe
ni
x
i
t
e
s Ge
no
z
on
e (
do H-
A)
i
s
f
ol
l
o
we
d i
n Be
d 3
5c by
t
h
e e
n
t
r
y o
f ir
f
s
t Fa
l
c
i
t
or
no
c
e
r
a
s a
nd q
ue
s
t
i
o
na
bl
e c
r
os
s
-
s
e
c
t
i
on
s of
Che
i
l
o
ce
r
as
.
Bot
h g
e
ne
r
a a
r
e de
ini
f
ng f
or
ms o
f t
he d
o 1
1
-
B.
Un
do
ub
t
e
d Che
i
l
o
c
er
a
s s
p
e
c
i
me
ns ha
ve no
t be
e
n c
o
l
l
e
c
t
e
d b
e
l
o
w
Be
d 39 a
nd ea
r
l
i
e
s
t f
a
l
c
i
t
or
no
c
e
r
a
t
i
ds (
es
pe
ci
a
l
l
y t
hos
e f
o
un
d i
n Be
d
3
6)
a
r
e s
t
i
l
l
s
ome
wh
a
t i
n
t
e
r
me
d
i
a
t
e t
o Phoe
ni
xi
t
e
s
.
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DDLE COL
I
MI
AC FORMATI
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PPERCOL
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OTTE FM .
of Pa.
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ul
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anson and M ehl
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.
s
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l
c
a
│ t
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cat
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a
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s
e
ns
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t
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)
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n Upp
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r
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um │ W alc1
tornocerasf
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│ u
m Cour
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ac Beds 32c,
33c,
and 34a i
s wr
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-
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eror
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a
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s
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r
c
h
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r
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s
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a
t
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m │ C
r
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c
k
i
t
e
s h
a
l
z
a
p
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l
f
│ Fi
gur
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c
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o Bed 39 i
n t
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he
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ar
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├──────────────────────────────────────
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二。
r
d │
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亚理 t
a
x
o
n
o
m
│
i
c
i
m
p
r
e
c
i
s
s
i
o
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二] i
n
f
we
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n
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ones
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on wi
t
h
t
he conodont
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│
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o
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t
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n,
s
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igur
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R
AYMOND】 反
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sodes.
VoL 16 no.
4
438
The Cou
r
n
i
a
c g
on
i
a
t
i
t
e s
e
qu
e
nc
e ge
n
e
r
a
l
l
y ma
t
c
h
es
f
a
un
a
l
s
uc
-
ces
si
ons des
cr
i
be
d f
r
om cl
ass
i
cal s
ect
i
ons of t
he Rheni
sh Sl
at
e
Mo
unt
a
i
ns a
nd Thur
i
ng
i
a.
Co
r
r
e
l
a
t
i
on wi
t
h t
h
e n
e
w i
nt
e
r
na
t
i
ona
l
z
on
a
t
i
on bas
e
d on t
h
e a
p
pe
a
r
an
c
e a
nd s
pr
e
a
d of ge
n
er
a (
Ho
us
e a
nd
Ki
r
c
hg
as
s
e
r
,
1
9
93;
Bec
k
e
r an
d o
t
he
r
s
,
1
9
93;
Be
c
ke
r
,
1
993
b)
i
s
s
t
r
a
i
g
ht
f
o
r
wa
r
d,
a
l
t
ho
ugh t
he
r
e a
r
e r
e
gi
ona
l
f
ea
t
ur
e
s mi
r
r
o
r
i
ng
l
o
c
a
l
f
a
c
i
e
s de
v
el
opme
n
t
s
.
For e
x
a
mpl
e
,
i
nt
e
r
na
t
i
o
na
l
ma
r
ke
r ge
n
-
er
a s
uch as Thmant
i
coceras,
Cari
noceras,
Neoni
ant
i
cocer
as and
Pl
a
y
f
or
di
t
e
s a
r
e s
t
i
l
l
l
a
c
ki
n
g i
n t
he Mon
t
a
gn
e Noi
r
e a
r
e
a.
La
t
e
Mi
ddl
e Fr
a
s
ni
a
n be
ds
at
Coumi
ac
,
be
l
ow t
he e
qui
va
l
e
n
t of t
he
Lowe
r Kel
l
wa
s
s
e
r Li
mes
t
o
ne (
Be
d 24
a
)
,
c
on
t
a
i
n Be
l
o
c
er
as
,
v
a
r
i
-
o
us ma
nt
i
c
oc
e
r
a
t
i
ds a
nd s
ome
wha
t
a
mb
i
gu
ous Co
s
t
a
ma
nt
i
c
o
ce
r
as
(
Ho
us
e an
d o
t
he
r
s
,
1
98
5;
pl
.
2,
f
i
g
ur
e 1
1
-1
2
)
.
The l
a
t
t
e
r i
s a ma
r
ke
r
f
or t
he Pl
ay
f
or
di
t
e
s a
nd Ne
omant
i
c
oc
e
r
as Ge
noz
o
ne
s (
do I
-
I a
n
d I
-
J
)
,
o
r f
or
t
h
e c
or
dat
um Zo
ne (
do 1
1
3
/
y)
of t
he c
l
as
s
i
ca
l
go
ni
a
t
i
t
e
z
ona
t
i
o
n.
I
n
de
x s
pe
ci
e
s f
or Di
vi
s
i
ons I
-
K s
o f
a
r ha
v
e o
nl
y be
e
n
f
ound above t
he Lower Kel
l
wass
er l
evel i
n Bed 25b (
Ar
chocer
as
s
p.
) a
nd Be
d 2
6b (
Ma
nt
i
c
oc
e
r
a
s ado
价n
s
e
)
.
The ir
f
s
t
e
vi
d
e
nc
e f
o
r
t
h
e l
a
t
e
s
t
Fr
as
ni
a
n Cr
i
c
ki
t
e
s Ge
n
oz
on
e (
d
o I
-
L)
co
mes
f
r
o
m Be
d
2
6c.
At
Co
un
na
c a
n
d n
ei
gh
bo
ur
i
ng s
e
c
t
i
ons
o
t t
he Mon
t
Pe
y
r
o
ux
Na
ppe a s
uc
c
es
s
i
on o
f Cr
i
c
k
i
t
e
s s
pe
c
i
e
s ca
n pos
s
i
bl
y be es
t
a
b-
l
i
s
h
e
d.
Al
l
e
a
r
l
y,
of
te
n s
ome
wha
t dou
bt
f
u
l
membe
r
s o
f t
he l
i
ne
a
g
e
ha
v
e who
r
l
f
or
ms s
i
mi
l
ar
t
o Ma
nt
i
c
oc
e
r
a
s c
or
d
at
u
m a
nd ma
y b
e
r
el
at
e
d t
o t
he Can
ad
i
a
n Cr
i
c
k.
c
or
di
for
me (
Mi
l
l
e
r
)
.
Cr
i
c
k
.
h
ol
z
a
p
f
e
l
i
,
t
he i
n
de
x s
pe
c
i
e
s of t
he c
l
as
s
i
c
al
ho
l
z
ap
f
el
t
Zo
ne
,
e
nt
e
r
s
i
n gr
e
a
t pr
o
f
us
i
on i
n t
h
e l
o
we
r pa
r
t
o
f Be
d Me a
n
d co
nt
i
nu
es
t
o
Be
d 31
g
a
nd t
o t
he e
nd of t
he Fr
a
s
ni
a
n.
Th
e a
p
pa
r
e
nt
p
at
c
hi
ne
s
s o
f
t
h
e Uppe
r Fr
a
s
n
i
a
n go
ni
a
t
i
t
e r
e
c
or
d i
s o
nl
y t
o a l
i
mi
t
e
d e
xt
e
nt
a
c
ons
e
qu
e
nc
e of s
a
mp
l
i
ng bi
a
s
.
Cl
e
a
r
l
y t
wo di
f
f
e
r
e
n
t go
ni
a
t
i
t
e
M
I
D
D
I
Z C
O
U
M
I
A
C F
O
R
M
A
T
I
O
N
a
s
s
e
mbl
a
ge
s a
l
t
e
r
na
t
e wi
t
h e
a
c
h ot
he
r
.
The f
i
r
s
t i
s c
ha
r
a
c
t
e
r
i
z
e
d by
gr
e
a
t
a
b
unda
n
c
e of Bel
oc
e
r
as t
e
nui
s
t
r
i
a
t
um (
e.
g.
i
n Beds
26
d a
n
d
31
a
)
a
nd r
a
r
i
t
y o
f ge
p
hur
o
ce
r
a
t
i
ds a
nd t
or
no
ce
r
a
t
i
ds
.
Ot
he
r be
d
s
ha
ve r
i
c
h Mant
i
c
oc
e
r
as or Cr
i
c
ki
t
e
s f
a
una
s a
c
c
ompa
ni
e
d b
y mor
e
a
bun
da
nt
t
or
no
c
e
r
a
t
i
ds
,
a
nd Buc
hi
ol
a.
Du
e t
o t
he l
a
c
k of l
i
t
h
ol
o
gi
-
ca
l
di
f
f
e
r
e
nc
e
s be
t
we
en be
d
s c
on
t
a
i
n
i
ng t
h
e t
wo f
a
un
al
t
y
pe
s i
t
i
s
i
nf
e
r
r
e
d t
ha
t c
ha
ngi
ng a
s
s
e
mbl
a
g
es
mi
r
r
or luc
f
t
u
a
t
i
o
ns i
n t
he
t
r
op
hi
c s
t
r
u
c
t
u
r
e
.
Es
pe
c
i
a
l
l
y t
he Upp
e
r Ke
l
l
wa
s
s
e
r Li
me
s
t
o
ne
eq
ui
va
l
e
nt
s wi
t
h t
he
i
r
r
i
c
h Cr
i
c
ki
t
e
s a
nd pe
l
ec
y
pod f
a
una
s a
s
s
oc
i
-
a
t
e
d wi
t
h unus
ua
l
l
y l
a
r
g
e os
t
r
a
c
ods but
wi
t
ho
ut
Bel
oc
e
r
as ma
y
ha
ve be
e
n de
pos
i
t
ed unde
r e
ut
r
o
phi
c e
c
ol
ogi
ca
l
c
o
ndi
t
i
o
ns
.
Thi
s
s
ug
ge
s
t
s t
ha
t
b
e
l
oc
e
r
a
t
i
ds t
h
r
i
ve
d d
ur
i
n
g ol
i
go
t
r
op
hi
c pe
r
i
o
ds
.
The c
ho
s
e
n s
t
r
a
t
ot
ype h
as
on
e o
f t
he b
es
t
kn
own go
ni
a
t
i
t
e
r
ec
o
r
ds a
r
ou
nd t
h
e Fr
a
s
ma
n
/
Fa
me
nma
n bo
un
da
r
y a
nd i
s t
h
e be
s
t
kno
wn a
ny
wh
e
r
e f
or
i
nd
e
x s
pe
c
i
e
s oc
c
u
r
r
i
ng i
n b
e
ds e
xa
c
t
l
y a
t
t
he
bo
und
a
r
y.
Howe
ve
r
,
i
t
i
s f
a
r f
r
o
m b
e
i
n
g a
n i
d
ea
l
a
mmono
i
d l
oc
a
l
i
t
y
d
ue t
o a
s
p
e
c
t
s o
fp
r
e
s
e
r
v
a
t
i
on a
n
d d
i
f
f
i
c
u
l
t
r
ec
o
ve
r
y o
f l
a
r
ge f
a
u
na
s
.
Thi
s mi
s
f
o
r
t
un
e c
a
n pa
r
t
l
y b
e ba
l
an
c
e
d by t
he wi
de r
a
nge o
f ot
he
r
g
on
i
a
t
i
t
e-
be
a
r
i
ng a
nd e
a
s
i
l
y c
o
fe
l
a
t
a
bl
e l
o
c
a
l
i
t
i
es
n
e
a
r
by i
n t
he
Mo
nt
a
gn
e Noi
r
e,
whi
c
h s
upp
l
y a
d
di
t
i
o
na
l
i
nf
o
r
mat
i
on on s
t
r
a
t
i
-
g
r
a
ph
i
c
a
l
r
an
g
es a
nd oc
c
u
r
r
en
c
e
s i
n s
l
i
g
ht
l
y di
f
f
e
r
e
nt
f
a
c
i
e
s s
e
t
t
i
ng
s
The
s
e wi
l
l
be d
oc
ume
n
t
e
d el
s
e
whe
r
e (
Be
c
ke
r a
nd Ho
us
e
,
1
9
94 i
n
pr
ess
).
Tr
i
l
obi
t
es
Amo
n
g
b
o
t
t
o
m-
l
i
v
i
n
g b
i
o
t
a
s
,
t
r
i
l
o
b
i
t
e
s
a
r
e
mo
s
t
f
r
e
q
u
e
n
t
a
n
d
d
i
v
e
r
s
i
f
i
e
d i
n t
he La
t
e Fr
a
s
ni
a
n s
t
r
a
t
a a
t Cour
ni
a
c.
Of e
i
gh
t f
a
mi
l
i
es
}
U
P
P
E
R C
O
U
M
I
A
C F
O
R
M
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T
I
O
N
│
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AN FAMENNI
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l
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I
c
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e
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a
l
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c
I
d}
’
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a
l
b
I
c
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a
l
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I
c
I
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l
b
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c
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l
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c
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d
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l
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d
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l
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l
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l
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nf
er
r
e
d ││
ra
nge │ Fi
gu
r
e7 Ra
nge c
har
t s
ho
wi
n
g t
he t
r
i
l
obi
t
e r
e
c
o
r
df
r
om Be
d2
3 t
o Be
d 39 ar
o
und t
he GSSP i
n t
he s
e
c
t
i
o
n a
t
t
he Uppe
r Quany at
Co
un
z
i
ac
.
Modi
ie
f
df
r
o
m dat
a o
f R Fe
i
s
t (
i
n Fe
i
s
t
,
19
90
)
.
[)Prpmhpr 1993
439
k
nown gi
oba
l
l
y t
o he pr
e
s
e
n
t at
t
he Up
pe
r Ke
l
l
wa
s
s
er
e
xt
i
n
ct
i
on
c\cnt (
Fei
s
t
.]
990:
1
991)
.Si
x Occur i
n t
he st
r
a
t
ot
ype s
ect
i
on
(
Pi
oc
t
i
da
e
,
AUl
a
c
opl
e
ur
i
d
a
e.
Odon
t
o
pl
C
U
r
i
da
e
.
Ha
r
p
et
i
da
c
,
Da
l
-
ma
ni
t
i
d
a
e a
n
d Pha
c
op
i
da
e)
a
nd o
f
a t
ot
a
l
of 1
3 s
p
ec
i
es
kn
own
,
doba
l
i
y,
ni
ne ar
e r
ep
r
es
e
nt
e
d a
t
COL
I
l
f
ua
c (
s
e
e f
i
-u
r
e 7)
.
Al
l
b
ut
t
h
r
e
e f
a
mi
l
i
e
s a
n
d川 s
pe
c
i
es
di
s
a
ppe
a
r w i
t
hi
n Zon
e 1
3
.
o
r a
t
t
h
e
ba
s
e of
t
h
e Upp
er
Kel
l
wa
s
s
e
r Li
mes
t
on
e e
qUI
Va
l
en
t
(
Be
d 3}
9)
.
Th
e
l
a
t
t
e
r d
oes
n
ot
yi
e
l
d a
ny t
r
i
l
o
bi
t
es
,
n
o
r d
o t
h
e S
u
c
c
e
e
di
n
g be
ds oft
h
e
Lo
we
r t
r
i
a
n
gt
dar
i
.
s
Zon
e (
Be
d 32
a
-
b)
,
a f
a
c
t
wh
i
c
h h
as
b
e
e
n o
bs
e
r
v
e
d
i
n a
l
l
kn
o
wn F
r
a
s
ma
n
/
Fa
r
n
e
r
i
ma
n bo
un
da
r
y s
e
c
t
i
on
s wo
r
l
d wi
d
e.
Re
c
ov
e
r
y do
e
s n
o
t
t
a
k
e pl
a
c
e e
a
r
l
i
e
r t
h
a
n f
r
om t
h
e Mi
ddl
e t
r
i
a
n
gul
ar
i
,
}
Zo
ne o
nwa
r
ds
wh
e
n s
ol
e
l
y ph
a
c
op
i
ds oft
h
e g
e
n
u
s A劝Ar
a
t
i
o
p
s o
c
c
u
r
(
Be
ds 32c
-e)
.
Thus t
he del
’
i
ned basal
Famenman at t
he bas
e of t
he
Lower
t
r
i
a
m}
i
i
l
ar
i
s Zone
.
ca
nn
ot
be pr
e
ci
s
e
l
y r
ec
og
r
u
s
e
d L
I
S
1
1
1
"
t
r
i
l
o
b
i
t
e
s
.
B
y c
o
n
t
r
a
s
t
.
t
h
e b
a
s
e
o
f
t
h
e t
o
p
mo
s
t
F
r
a
s
n
i
a
n Ke
l
l
wa
s
s
e
i
Li
i
rc
,
,
t
one l
e
ve
l
c
an b
e l
o
ca
t
e
d wi
t
h con
s
i
de
r
a
bl
e pr
e
c
i
s
i
on by t
h
e
ma
j
o
r ex
t
i
n
ct
i
on a
f
f
ec
t
i
n.
, t
r
i
l
obi
t
es
a
t
t
ha
t
l
ev
e
l
(
Fe
i
s
t
a
nd
Schi
ndl
er,
1994)
.
Fi
n
e
-
s
c
a
l
e i
n
t
r
a
z
on
a
l
s
ub
di
v
i
s
i
on a
n
d bi
os
t
r
a
t
i
g
r
a
phi
c
al
c
o
r
i
-
c
-
l
a
t
i
on oft
h
e l
a
t
e
s
t
Fr
a
s
ni
a
n a
t
CoL
u
ni
a
c i
s pr
o
vi
de
d by ev
ol
ut
i
o
na
r
y
s
p
e
ci
es
of
、
t
he p
r
oe
t
i
d Pal
pe
b
,
一
(
di
a wh
i
ch a
r
e c
ha
r
a
c
t
e
r
i
z
e
d b
y a t
r
en
d
i
n r
e
du
c
t
i
on o
f t
h
e p
al
p
eb
r
al
l
o
be a
n
d i
n t
h
e s
t
r
a
i
g
ht
e
ni
n
g of t
h
e
f
a
c
i
a
l
s
u
i
u
r
es
.
Th
e a
n
c
e
s
t
r
al
f
o
r
m,
Pal
.
l
at
e
pa
l
pe
br
al
i
s b
ei
n
I
g a
l
r
e
ad
y
pr
e
s
en
t
b
el
ow t
h
e Lowe
r Ke
l
bv
a
s
s
c
r[f
i
nes
t
on
e c
qL
]
i
\
a
l
e
n
t
(
Be
d
24
a)
,
ha
s be
e
n r
e
c
ov
er
ed a
s hi
,
,
h a
s i
n t
h
e mi
d
dl
e of Zon
e]
3 (
Be
d
29
d)
.
I
t
g
I
a
ve r
i
s
e t
o Pa
l
.
pal
pe
br
al
i
s whi
c
h i
s a
bu
n
da
n
t川Be
ds
3 1
。
t
o 3 1
e and i
s al
s
o known f
r
oi
n l
at
es
t Fr
as
i
nan s
ect
i
ons i
n t
he Rhen-
i
s
h Sc
hi
cf
c
r
1
(
,
c
bf
i
-
g
c a
n
d Ha
r
z (
Fe
i
s
t
a
nd S
ch
i
n
dl
er
,
1
99
4)
a
n
d ma
y
Oc
c
ur
i
n t
he Ca
n
ni
m,
Bas
i
n of
We
s
t
e
r
n Au
s
t
r
al
i
a (
K Mc
Na
ma
r
a
.
o
r
al
co
mmun
i
c
a
t
i
on)
Th
e l
as
t
r
e
pr
e
s
cr
i
t
a
t
i
v
e of
t
h
e l
i
n
e
a
g
e i
s
Pal
.
b
r
e
(
-
(i
oe v
\
h
i
c
h ci
ne
r
s t
h
e s
t
r
a
t
ol
yp
e s
e
c
t
i
o
n i
n Be
d 3 1
c an
d f
,
t
h
us
mar
k
-
i
ng p
r
e
ci
s
el
y t
h
e l
as
t
ox
yg
e
na
t
e
d l
c
v
el
-
bo
t
t
oi
n en
vi
r
on
me
nt
pr
i
o
r t
o
t
he h
yp
oxi
c ov
er
t
u
r
n o
f t
h
e Up
pe
r Ke
l
l
wa
s
s
e
r
Li
me
s
t
on
e
.
Th
i
s
s
pe
c
i
e
s i
s a
t
l
e
a
s
t
of mod
e
r
a
t
e v
a
l
u
e f
o
r l
ong d
i
s
t
a
n
c
e c
o
r
r
el
a
t
i
on a
s
i
t
ha
s be
e
n r
e
c
ov
e
r
e
d i
n t
he Rh
e
ni
s
h Sc
h
i
e
f
e
r
g
eb
i
r
g
e (
a
t
St
embr
u
ch
Sc
hmi
dt
)
a
i
记i
n t
h
e Ha
r
z (
Ac
kc Va
l
l
e
y)
i
n a
n e
qu
i
v
a
l
en
t
p
os
i
t
i
on
a
n
d a
s
s
oc
i
a
t
e
d t
h
e
r
e wi
t
h Pal
i
)
i
at
ol
e
pi
s l
i
n
guU
i
)
r
i
n
i
s (
Sch
i
ndl
e
r
.
1
99
0)
.
Cons
e
qu
e
nt
l
y Pal
.
br
e
c
ci
ae a
p
pe
a
r
s
t
o be l
i
mi
t
e
d t
o t
h
e l
owe
r
p
ar
t
of
t
h
e l
i
n
gui
Mi
-
mi
s Zon
e
.
Th
i
s s
up
por
t
s t
h
e r
e
s
u
l
t
s obt
a
i
ne
d by
gr
a
p
hi
c c
or
r
e
l
a
t
i
o
n,
wh
i
c
h i
n
di
ca
t
e
d t
he p
r
e
s
e
n
c
e of l
i
n
gi
t
咖r
MI
A
Zo
ne e
qui
va
l
e
nt
s a
t
Cour
ni
a
c i
n Be
ds 3 I
f a
n
d g (
Ma
p
p
er
i
n Be
c
k
e
r
a
nd o
t
h
e
r
s
,
1
98
9,
p.
26
2)
.
Conc
o
mi
t
an
t
l
y wi
t
h t
he Pal
pe
b
r
al
i
a l
i
n
e
,
a
n
ot
h
e
r e
v
ol
ut
i
on
a
r
y
t
r
e
n
d i
s
s
e
en i
n S
u
c
c
e
e
di
ng po
pul
a
t
i
on
s o
f
Cr
Y
pho
ps ac
u
t
i
ce
ps a
n
d
t
hi
s t
r
e
n
d i
s of bi
os
t
r
a
t
i
g
r
a
ph
i
c
a
l
s
i
gni
f
i
c
a
n
c
e.
Thi
s gr
ou
p,
whi
c
h i
s
r
a
t
he
r
c
ommon i
n Be
d
s 3 1
a t
o 3 1
f
,
e
xh
i
b
i
t
s a s
p
e
c
t
a
c
ul
a
r
r
e
duc
t
i
on
i
n t
h
e i
ne
a
n nu
mb
e
r of i
t
s
e
y
e l
e
n
s
e
s wh
i
ch d
r
op f
r
om t
e
n l
en
s
e
s
i
n
Be
d 3 1
a t
o on
l
y t
h
r
e
e l
e
ns
e
s i
n Be
d 3 1
f be
f
o
r
e i
t
s
ina
f
l
e
x
t
i
nc
t
i
on
(
Fe
i
s
t
,
1
991
)
.Thi
s
CV
OI
L
I
t
i
o
n ma
y r
e
pr
e
s
e
nt:
、f
ur
t
he
r
po
t
e
n
t
i
al
f
oi
t
i
n
e
-
s
c
a
l
e h
i
os
t
r
a
t
i
g
r
a
p
hi
c
al
s
u
b
di
v
i
s
i
o
ns a
nd c
or
r
e
l
a
t
i
on dur
i
ng a
p
e
r
i
o
d of
'
wo
r
l
d
-
wi
de bi
ot
i
c c
r
i
s
i
s
.
Ot
her f
os
s
i
l
gr
oups
Pa
l
y
n
o
mo
r
p
h
s
h
a
v
e
b
e
e
n
o
b
t
a
i
n
e
d
f
r
o
m t
h
e
U
p
p
e
r
K
e
l
l
wa
s
s
e
r
l
e
v
e
l
(
Be
d 3 1
-)
bu
t
a
r
e t
o
o ba
d
l
y p
r
e
s
e
r
v
e
d f
or
pr
ec
i
s
e i
de
nt
i
ic
f
a
t
i
on
:
f
ur
-
t
h
e
r wor
k i
s r
e
qu
i
r
e
d h
e
r
e
.
The s
t
r
a
t
i
g
r
a
phi
c
a
l
l
y i
mp
or
t
an
t
e
nt
or
no
〕
-
zoaccans ar
e a ra
t
her mi
nor f
aunal
el
ement
}
de
t
ai
l
ed wor
k i
s cur
-
r
e
nt
l
y b
e
i
n
g c
a
r
r
i
e
d ou
t
b
y F Le
t
hi
e
r
s
.
S
ol
i
t
ar
y r
ug
os
e c
or
a
l
s
,
ga
s
-
t
r
opods,
or
t
hoconi
c and br
evi
coni
c naut
i
l
oi
ds
,
cr
i
noi
ds and r
ar
e is
f
h
s
c
a
l
es
r
ep
r
e
s
e
n
t
f
ur
t
he
r a
cc
e
s
s
o
r
y f
os
s
i
l
g
r
ou
ps S
t
i
l
l
t
o be i
nv
e
s
t
i
-
"
a
t
e
d i
n de
t
a
i
l
.
Br
a
c
hi
op
od
a a
n
d bi
v
al
v
e r
e
c
o
r
ds of C Bab
i
n an
d P R
Ra
ch
e
b
oeu
t
h
a
v
e be
e
n pr
e
s
e
nt
e
d (
i
n F
ei
s
t
.
1
99
0)
.
Ri
c
h h
or
n
o
ct
en
i
d
f
au
na
s
ha
v
e be
e
n de
t
e
r
mi
n
ed b
y M Tr
u
vol
s
-
Ma
s
s
or
n (
i
n Fe
i
s
t
.
1
9
90)
}
t
h
e Hoi
n
oc
i
e
nu
s u
l
t
i
mu
s Zon
e i
s i
f
r
s
t
r
e
c
og
ni
z
ed i
n Be
d 2
6c
and t
her
ef
or
e s
eems t
o cor
r
el
at
e wi
t
h t
he Cri
cki
t
es Genozone.
0s
t
r
a-
cod dat
a of
'
F Let
hi
er
s have been l
i
s
t
e
d (
i
n Fei
st
, 1
990)
,
and mor
e
t
ha
n 30 di
f
f
e
r
en
t
s
p
e
ci
e
s ar
e r
ec
o
gni
s
e
d i
n t
he t
opmo
s
t
b
e
ds of t
h
e
Epi
sM es
,、初/
.I6,,
,
(
,
.4
F
r
as
ma
n.
a
r
n
om,
t
he b
e
nt
ho
ni
c f
o
r
ms
.
t
he Fr
a
s
ni
a
n
[
Fa
n
i
e
nn
i
a
n
bou
n
da
r
y i
s
c
ha
r
a
c
t
er
i
z
e
d b}
a ma
j
or ex
t
i
n
ct
i
o
n s
i
n
ce 0
5 pe
r
c
e
n
t
of
al
l
r
e
c
o
r
de
d t
a
xa山s
a
p
p
ea
r
t
h
e
r
e (
Le
t
hi
e
r
s
a
n
d F
e
i
s
t
.
1
9
91
)
.Thu
s
.
t
h
e Co
ur
ni
a
c s
t
r
a
t
ot
yp
e be
a
r
s
e
vi
de
n
c
e(
)
【a
n ex
t
r
a
or
di
n
a
r
y b
r
e
a
dt
h
off
a
un
al
r
e
p
r
e
s
e
n
t
a
t
i
on e
n
abl
i
ng c
or
r
e
l
a
t
i
on i
n
t
o r
e
-i
n
-
l
e
s wi
t
h be
t
t
er
s
po
r
e an
d a
cr
i
t
a
r
c
h r
e
c
or
ds
.
Th
e SDS vi
e
ws
t
h
i
s
d
oc
ume
nt
a
t
i
on a
s
t
he b
es
t
ofa
ny oft
he l
ev
el
s
i
t
ha
s
r
e
c
omme
nd
e
d l
o
t
-
b
0L
mda
r
y s
t
r
a
t
o-
t
ypes i
n t
he Devoi
nan.
Che
mos
t
rat
i
g
raphy
As
y
e
t
g
e
o
c
h
e
r
n
i
c
a
l
m
e
t
h
o
d
,
d
o
n
o
t
p
r
o
v
i
d
e
a
n
u
n
a
i
r
i
b
i
2
t
-
I
O
U
S
g
u
i
d
e
t
o c
o
r
r
el
a
t
i
on i
nt
e
r
n
a
t
i
on
a
l
l
y.
a
l
t
h
ou
gh t
he
r
e i
s gr
e
at
p
o
t
e
nt
i
a
l
e
s
pe
-
ci
a
l
l
y wi
t
h s
t
r
o
nt
i
um i
s
o
t
o
pe c
h
er
n
os
t
r
a
t
i
gr
a
p
hy a
ppl
i
e
d wi
t
hi
n t
he
f
r
a
r
n
ewor
k o
f c
o
no
do
nt
bi
o
s
t
r
a
t
i
gr
a
ph
y (
e
.
g.
Ru
pp
e
l
a
nd ot
h
e
r
s
.
1
9
93)
.
Ge
oc
l
i
c
i
ni
c
al
me
t
h
o
ds d
o.
ho
we
v
e
r
,
pr
ov
i
de a
n i
mpo
r
t
a
n
t
cl
u
e t
o e
nv
i
r
onme
n
t
a
l
i
n
t
er
pr
e
t
a
t
i
o
n.
The s
t
r
a
t
o
t
y
pe s
e
c
t
i
o
n a
t
C0
1
1
1
1
1
j
a
c h
as
b
e
en i
nv
e
s
t
i
ga
t
e
d f
or
6 "CI
"C v
al
u
es
b
y J
o
a
c
hi
ms
k
i
a
n
d Bug
gi
s
ch(1
9
93)
whe
r
e:
、
po
s
i
t
i
v
e S
h
i
f
t
of (
3
-
1
T wa
s
n
ot
e
d a
t
t
h
e
b
oun
d
a
r
v.
Rar
c-
c
a
r
t
h e
l
emen
t
s
i
RLI
-
-
)
h
av
e b
ee
n i
n
ve
s
t
i
ga
t
e
d b
y
Gr
an
dj
e
a
n-
L&L
l
y
e
r a
n
d o
t
h
er
s(1
9
93)
u
s
i
n
g i
n
di
vi
du
a
l
De
von
i
a
n
c
on
o
don
t
s
,
bu
t
n
o a
n
or
n
al
y a
t
t
h
e b
oun
d
ar
y wa
s no
t
ed a
l
t
h
ou
gh t
he
REF p
a
t
t
er
ns
di
d n
ot
c
onf
or
m wi
t
h t
h
os
e of
mod
e
r
n s
e
a wa
t
e
r
.
Th
e
pu
r
s
u
i
t
O
f i
t
-
i
di
l
l
m an
oma
l
i
e
s
ha
s
n
ot
be
en S
uc
c
e
s
s
f
ul
a
t
Co
un
na
c
(
Gi
r
ar
d and ot
her
s
, 1
993)
, nor at St
ei
nbr
uch Schmi
dt (
McGhee and
o
t
h
er
s
,
1
9
84)
.
Th
os
e cl
a
i
me
d a
t
t
he b
ou
nd
j
r
y i
n t
h
e Ca
n
ni
ng Ba
s
i
n
ha
v
e p
r
ov
e
d t
o he f
r
om a muc
h I
nL
he
r
s
t
r
a
t
i
g
r
a
phi
c l
e
v
el
(
Be
c
k
er
a
nd ot
h
er
s
,
1
9
91
:
Ni
c
ol
l
a
nd Pl
a
yf
or
d.
1
9
93)
a
l
t
h
ou
gh t
h
e
r
e i
s
a
we
a
k i
r
i
d
i
u
m a
n
on
i
a
l
v a
pp
a
r
e
nt
l
y a
t
t
h
e c
or
r
e
c
t
l
e
ve
l
a
t
Xi
a
ng
t
i
a
n
(
Wa
l
l
g a
nd ot
he
r
s
,
1
99 1
)a
nd ot
h
e
r
el
e
me
n
t
a
n
oma
l
i
e
s
,
bu
t
t
h
e c
a
u
s
e
i
s uncer
t
ai
n.
Ther
e has been no l
i
nk of r
ui
cr
ot
ekt
i
t
es wi
t
h t
he bound-
a
r
y s
t
r
a
t
o
t
yp
e a
n
d r
e
c
or
d
s el
s
e
wh
e
r
e a
r
e hi
gh
er
i
n t
h
e Fa
me
n
ni
a
n
(
hi
gh wi
t
hi
n t
h
e Lower
t
r
i
an
gi
dar
i
s Zo
ne a
t
S
en
z
c
i
l
l
c
s
i
n Bel
g
i
u
m:
Cl
a
e
ys
a
nd ot
he
r
s
,
1
99
2.
f
i
gu
r
e I
;Lo
we
r
cr
e
j
)
i
d
o Zon
e i
n S
o
ut
h
Ch
i
n
a:
Wa
ng
,
1
9
92)
.
Re
l
at
i
ons
hi
p t
o t
he Uppe
r Ke
l
l
was
s
e
r Eve
nt
The mai
n s
e
di
me
n
t
a
r
y ma
r
ke
r
oft
h
e GSSP i
s t
he t
o
p of
t
h
e di
s
t
i
n
c
-
t
i
v
c l
ev
e
l
k
no
wn i
n Ge
r
ma
n
y a
s t
he Up
pe
r
Kel
l
wa
s
s
e
r Li
me
s
t
on
e
(
Wa
l
l
i
s
e
r
a
n
d ot
he
r
s(1
98
9)
.
Thi
s da
r
k h
yp
oxi
c l
i
me
s
t
on
e a
pp
e
a
r
s
t
o
r
eDr
cs
ent ai
l acme i
n t
he sm-
cad of a di
st
i
nct
i
ve 九ci
es whi
ch i
n
many sect
i
ons ,
I
ot
)
al
l
y i
s pr
eci
sel
y const
r
ai
neo ny conoc
i
oni
mos
-
t
r
a
t
i
g
r
a
phy.
I
n p
a
r
al
l
el
wi
t
h t
he wor
k of t
he SDS i
n r
e
c
en
t
ye
a
r
s h
a
s
b
ee
n t
h
e r
e
c
o
gni
t
i
on of
a
n i
mp
or
t
a
n
t
ex
t
i
nc
t
i
on ev
e
n
t n
e
a
r t
he
Fr
as
ni
a
n
/
F
ar
n
en
ni
an b
oun
da
r
y.
Some of
一
t
h
es
e ex
t
i
nc
t
i
on
s we
r
e
l
i
s
t
e
d i
n t
he p
r
e
vi
ous
s
e
c
t
i
ons
.
Th
e r
e
c
omme
n
da
t
i
on o
f t
h
e Su
b-
c
ommi
s
s
i
on f
o
r a GSSP f
a
l
l
s
a
t
a l
e
ve
l
i
mme
di
a
t
e
l
y a
bo
ve t
h
e a
c
me
of ext
i
nct
i
ons
.
t
hat i
s at t
he bas
e of t
he Lower t
r
i
ongul
ori
s Zone.
Th
e mos
t
pr
e
c
i
s
e d
oc
ume
n
t
a
t
i
on f
o
r t
hi
s
(
Be
ck
e
r a
n
d ot
h
e
r
s
,
1
98
9)
h
a
s b
e
en f
ol
l
owe
d by da
t
a a
s
s
e
mbl
e
d f
or
t
he Sub
c
ommi
s
s
i
on a
n
d
i
l
l
us
t
r
a
t
e
d h
e
r
e (
f
i
gu
r
e
s 5
-7)
.
Fol
l
owi
nI
g e
s
pe
c
i
al
l
y t
h
e wo
r
k of
Sa
n
db
e
r
g a
n
d ot
he
r
s(
1
9
88)
t
hi
s
l
ev
e
l
h
a
s be
e
n wi
d
el
y t
r
a
c
e
d i
n
t
er
-
n
a
t
i
ona
l
l
y.
The
r
e
ha
s
be
e
n mu
c
h d
e
ba
t
e on t
he C
a
us
e of
t
he s
e
di
men
t
a
r
y
p
e
r
t
u
r
ba
t
i
on r
e
pr
e
s
en
t
ed by t
h
e Up
pe
r
Kel
l
wa
s
s
e
r Li
mes
t
on
e.
The
mat
t
er cannot be s
ai
d t
o be r
esol
ved.
I
ndeed,
s
ome member
s of t
he
Su
bc
ommi
s
s
i
o
n e
a
r
l
i
e
r
t
ook t
h
e vi
e
w t
ha
t
a mor
e a
p
pr
op
r
i
a
t
e bo
un
d-
a
r
y mi
g
ht
b
e c
ho
s
e
n a
wa
y f
r
om t
h
e s
u
d
de
n f
a
un
a
l
a
n
d s
e
di
men
t
a
r
y
c
h
an
g
cj
us
t
b
el
ow t
h
e ba
s
e oft
h
e Lo
we
r
t
r
i
an
gul
ar
i
s Zon
e b
ut
i
n t
he
e
n
d,
t
h
e e
a
s
e ofi
n
t
e
r
na
t
i
o
na
l
c
on e
l
a
t
i
on ba
s
e
d on t
he f
a
un
a
l
c
ha
n
ge
s
l
e
d t
o t
hi
s bo
un
da
r
y b
e
i
n
g r
e
c
oi
nm
e
t
i
de
d.
Fou
r ma
i
n g
r
ou
ps of hy
po
t
h
es
e
s ha
v
e
b
e
en i
nv
ok
e
d t
o e
xp
l
ai
n
t
h
e f
a
un
a
l
an
d s
e
di
me
nt
a
r
y c
h
an
g
es
a
r
o
un
d t
he b
a
s
e of t
he Lowe
r t
r
i
-
an
gu
l
ar
i
s Zon
e
.
Fi
r
s
t
l
y,
c
a
u
s
e
s r
e
l
a
t
e
d t
o a bol
i
d
e i
mp
a
c
t
o
r i
n
a
pa
ct
s
(
San
d
be
r
- a
n
d o
t
h
er
s
1
989
,
Mc
La
r
e
n a
n
d Goo
df
e
l
l
ow 1
9
90
)
.
S
ec
-
on
dl
y,
a s
p
r
e
a
d of
'
a
n
oxi
c c
on
di
t
i
on
s on c
on
t
i
ne
nt
a
l
s
h
el
v
es
a
}
s
o
c
i
-
a
t
e
d wi
t
h pos
s
i
bl
e t
e
c
t
on
i
c e
v
e
nt
s
an
d o
c
e
an ov
e
r
t
u
r
n (
Wi
l
d
e a
n
d
440
Be
r
r
y
,
1
98
4}
Bu
1
g
gi
s
c
h
,
1
9
91
)wi
t
h a t
r
an
s
gr
e
s
s
i
o
n f
ol
l
owe
d by
q
ui
c
k r
e
g
Z
,
r
e
s
s
i
o
n a
t
t
h
e L
i
pp
e
r bou
n
da
r
y (
Sa
n
db
e
r
g a
n
d o
t
he
r
s
,
1
98
8)
.
Thi
r
dl
y.
t
h
es
e s
e
c
on
d ev
e
nt
s bu
t
a
s
s
oc
i
a
t
e
d wi
t
h a p
e
a
k of c
o
l
d c
l
i
-
i
na
t
i
c c
on
di
t
i
on
s
,
r
es
ul
t
i
n
g i
n a r
i
s
e of
'
t
h
e p
yc
n
oc
hn
e (
Cop
p
er
,
1
98
6)
.
Fou
r
t
hl
y.
t
h
e s
a
n
i
e
,
bu
t
wi
t
h t
he Upp
e
r Ke
l
l
wa
s
s
e
r
Li
me
s
t
o
ne
r
ep
r
e
s
en
t
i
n
g a p
ea
k of h
ot
cl
i
ma
t
i
c c
on
di
t
i
o
ns
,
r
es
ul
t
i
ng al
s
o i
n a r
i
s
e
of t
h
e py
cn
o
cl
i
n
c a
nd p
r
o
ba
bl
e di
s
r
u
pt
i
on oft
h
e t
r
o
ph
i
c t
i
e
r
i
n
g,
pa
r
-
t
i
cu
l
a
r
l
y t
ha
t
of
t
he pl
a
n
kt
on (
Be
ck
e
r
a
n
d Ho
us
e
.
1
99
4)
;
t
h
i
s l
a
t
t
e
r
t
h
e
or
y wa
s f
i
r
s
t
i
nv
ok
e
d f
o
r De
vo
ni
a
n a
n
oxi
c ev
e
n
t
s b
y Be
ck
e
t
(
1
9
92)
.
Re
l
a
t
e
d h
yp
ot
he
s
e
s i
nv
ol
v
i
n
g c
l
i
ma
t
i
c wa
r
mi
ng h
a
ve b
e
en
s
ug
g
es
t
ed b
y Th
omps
o
n a
n
d Ne
wt
o
n(1
98
9)
a
nd Or
mi
s
t
on a
n
d
Kl
a
p
p
er(
1
9
92)
.
I
t
i
s -
e
ne
r
a
l
l
y r
e
c
og
ni
s
e
d t
h
a
t
t
h
e c
ol
l
a
ps
e of
s
t
r
o-
ma
t
op
or
o
i
d r
e
e
l
'
s
y
s
t
er
ns
r
i
c
a
r t
he en
d of
t
h
e Fr
as
ni
a
n l
e
d t
o e
x
t
i
n
c
-
t
i
ons(
)
{
’ass
oci
at
ed f
aunas
.
Hous
e(1
985) has dr
awn a
t
t
ent
i
on t
o t
he
ma
n
y s
i
mi
l
a
r e
v
e
nt
s a
t
o
t
h
e
r l
e
ve
l
s i
n t
h
e Dev
on
i
a
n s
uI
,
,
gc
s
t
i
i
l
g t
h
a
t
a
n i
n
t
e
r
p
r
e
t
a
t
i
o
n e
n
ab
l
i
n
I
c
,
a c
ommon h
yp
ot
he
s
i
s i
s t
o b
e pr
e
f
e
r
r
e
d.
Wh
a
t
e
v
er
t
he c
a
u
s
e
(
s
)
oft
h
i
s
s
e
di
me
n
t
a
r
y p
e
r
t
u
r
ba
t
i
on ma
y b
e,
t
h
e SDS t
ook t
h
e、i
ew t
h
a
t
t
h
e hy
p
oxi
c p
er
t
ur
b
a
t
i
ons
bc
l
o\
N
t
h
e b
a
s
e
of
t
h
e I
-
o\
x
,
c
r
t
r
i
mi
gul
ar
i
s Zon
e r
e
s
ul
t
e
d i
n a c
ons
i
d
c
r
j
bi
e f
a
un
al
c
ha
n
t
,
eov
e
r
an
d a
n h
or
i
z
on wh
i
c
h ma
y b
e c
or
r
e
l
a
t
e
d i
nt
e
r
na
t
i
on
a
l
l
y
wi
t
h pe
r
ha
p
s mor
e pr
e
ci
s
i
on t
ha
n a
n
y ot
he
r
i
n t
he De
v
oni
a
n.
I
t
i
s
i
n
t
h
e l
i
,
,
h
t
oft
h
i
s
v
i
ew t
ha
t
Cour
ni
a
c wa
s r
e
c
ommen
de
d f
or t
h
e GSSP
t
o del
i
ne t
he bas
e of t
he Fai
nenni
an St
a-c.
Fe
i
s
t
,
R,
ed,1
990,
The Fr
as
ni
an
/
Famenni
an boundar
y and a
dj
a
ccnt
s
t
r
at
a ol
t
he eas
t
e
r
n Mon
t
a
gue Noi
r
e,
Fr
a
nce:
Rj
GS Subcommi
ss
i
on on
Devoni
an St
r
at
i
gr
aph
y,
Gui
de Book,
69即 .
1
-
c
i
s
t
,
R,
1
99 1
,
Th
e l
at
e Dcvoni
a
n t
r
i
l
obi
t
e C
r
i
si
s
:
Hi
s
t
or
i
cal
Bi
ol
ogy,
v.
5,
pp.1
97
-
-
21
4
Fe
i
s
t
,
R,
and Schi
ndl
e
r
,
E
-
,
1
994,
Tr
i
l
obi
t
e
s dt
-
1
1
-
i
li
f
!
,
t
he Fr
a
sni
an KcI
l
was
s
er
cr
i
s
i
s i
n Eur
op
ean La
t
e De
von
i
a
n cepha
l
opo
d l
i
ni
cs
i
on
es:
Cour
i
er
For
s
chungsi
ns
t
i
t
ut
Senck
enhcr
g,
v.
1
69,
1
95-223.
Gi
r
a
r
d.
C,
Rocc
hi
a,
R,
Fei
s
t
,
R,
Fr
oge
t
,[
一
,a
nd Robi
n,
F
.
.
1
993,
No cvi
denc
e
of i
mp
act
a
t t
he Fr
a
sni
an
/
Famenni
an bOUnda
r
v i
n t
he s
i
r
at
ot
vp
c ar
ea. S
ou
t
he
r
n Fr
anc
e-
I
nt
er
di
s
ci
pl
i
nar
y Conf
e
r
en
ce on Gl
obal
BO
、
L
1
n
dai
\
1 Fvci
ns
,
Ki
el
ce
,
27-29 Scpt
c[
i
i
bc
r 1
993.
Abs
t
r
ac
t
.
Goodf
el
l
ow,
W 「
)
,Gel
ds
et
z
ei
,
I
F I
F J,
Mc
Lar
en,
D J
,
Or
c
har
d,
M J
,
and
Kk甲per
,
G,
1
989,
'
]
'
he Fr
as
ni
an
/
Famenni
an ext
i
nc
t
i
on-
cur
r
ent
1
-
C.
S
UI
t
S
and pos
s
i
bl
e caus
es.
i
n McMi
l
l
an,
N J
,
Embr
y,
A F and Gl
as
s
.
1
) J
,
c(
I
s
.
,
Devoni
a
n‘
)
}t
h
e Wor
l
d:
Canadi
an Soci
e
t
}(
)
!Pe
t
r
ol
eum (
wol
ogi
s
t
s, Mcni
oi
r 1
4,
}
.
3,
pp.
9-
-
2 1
.
Gr
andi
c
an,
1
'
.
Al
ba
r
e
de,
F.
an
d Fei
s
t
,
R,
1
989,
REE N
ar
i
a
l
i
ons aci
os
s t
he
1
了
r
是
1
、
1
1
1
之
1
1
1
/
1
了
之
汇
l
l
l
cl
l
l
l
l
之
川 b
ol
-
I
n
c
l
ar
v.
F
er
r
a八hs
t
r
a
ct
、
一\.
I
}
pp.
1
8
4
Gr
andj
ean-
1
.
6
cu
yei
,
P,
Fe
i
s
t
,
R,
and Al
hai
&l
c,
F,
1
993,
Ra
t
e c
at
(
h cl
e
ment
s
i
n ol
d bi
ogeni
c apal
i
t
cs
:
Geoc
hemi
ca c
t
(’
o
,
mocl
i
ci
ni
ca八ct
}
l
.
v.
}7
.
1
)
p
. 2507-
-
2514.
House, M R} 1985, Cor
r
el
ati
on ofmi
d-Pal
acozoi
c ar
nmonoi
d c\ ol
ui
i
onarv
eve
nt
s wi
t
h gl
obal
s
edi
ment
ar
I
v
p
er
t
ur
ba
t
i
ons:
Nat
ur
e,
Lon
dor,、3 1
'
,
pp.
1
7 -
22.
HOL
I
S
C
,
M R,
an
d Ki
r
chgas
s
er
,
W 下,1
993,
Devoni
;
川9‘
川l
a
t
i
t
e
bi
OS
I
I
-
al
i
gr
aph} a
nd t
i
mi
ng〔
)
{
、t
a
ci
cs mov
ement
s i
n t
he Fr
a
s
ni
a
n of
R eferences
e
as
t
er
n No
r
t
h Amer
i
ca,
i
n Ha
i
l
wood,
E
"
.
A.
and Ki
dd,
R 1
3
,
eds,H i
0
i
RC}
01
1
.
1
1
[
1
01
1 St
r
at
i
gr
aphy,
Geol
ogi
c
al
Soci
et
} Spe
ci
al
Publ
i
cat
i
on,
no.
7
0,
f
3
cck
er
,
R T,
1
992,
Zur Ker
ni
t
ni
s、
〔
川f
i
e川hCr
V-
St
l
I
l
C L
i
nd Annul
a
t
a-
I
.
ondon,pp.267-292
Schi
ef
et
i
m Nor
ds
au
er
l
an
d t
Ohci
-
(
I
c\on,
Rhei
ni
s
ches Schi
ct
cr
geb
i
r
gc,
Hous
e,
M R,
Kf
i
r
chgas
s
er
,
Pr
i
c
e,
J D,
al
l
(
]
Wade,
G,]
985.
Goni
a
t
i
t
es hom
GK 4611
Hohcnl
i
n
nhm,
,
)
:
Be
r
l
i
n
er Gcowi
s
s
ens
chal
-
li
f
ch
e Ahl
-
r
an
dl
Unge
r
Fr
a
sni
al
l
(
Uppe,
一Devoni
an)
a
r
i
d ad
j
a
cent
s
t
r
at
a of
t
he Mon
t
a
gn
e Noi
r
e:
F
}
,
\.
30 一pp.
3-
41
Fl
er
c
yni
c
a.、.]
、pp I
一]
9.
Bec
ker
,
R T}
1
993a.
St
r
a
l
i
gr
a
phi
s
che Gl
i
cd
er
r
j
I
1
2 und Ammonoi
de
en-
E Q}
1
989,
Oi
l t
i
r
e Fr
aqi
j
an
/
1
-
ani
cr
i
ni
an t
r
i
as
s c
xt
i
nct
i
on e\cr
i
t
i
n Sout
h
FAI
L
I
CI
I
i
n
n Neh(
l
ent
i
-
1
1
1
1 (
Ol
vi
dewn 1
1
)
wn FI
V
I
OI
NI L
i
n
d Nor
d-
Al
-
r
i
k
a:
( I
nt
r
a:(01
1
1
I
CI
-
FOt
S
Cl
l
Ul
)
,S
l
T
Ni
t
L
u Scn
cl
k
cnhc
q-
}
. ]
7,
pp.
-75-1
2
01
COL
I
l
i
c
r For
s
chUngS
i
n
St
i
t
i
r Scr
i
ck
cr
i
ber
g,
v
.
1
55,
405 pp,
,
l
oa
chi
mski
,
M Nt and Bu
ggi
s
c
h,
W,
1
993, Anoxi
c event
s i
l
l
t
h
e l
at
e
Bec
ker
,
R T,
1
993b,八c
oxi
a,
CI
L
I
S
UI
t
I
C cha
ngcs
,
and Upp
er Dc\
oni
a
n【
(
)
Fr
a
sn}
a
r-
-
-
Caus
es of t
he FI
飞
1
,
1
1
]
an/
1
了
是
1
l
l
l
cl
l
l
l
l
之
t
l
l t
a
unal
cr
i
s
i
s
:
GC01
0
2).、
l
ower
mos
t Car
boni
t
el
-
OL
I
S9l
obal
ammor
r
oi
d di
v
er
s
i
t
y,
m Hous
e
,
M R,
21
,pp.675-678
c
d.
,
The ammor
r
oi
dea:
cr
i
vi
i
-
or
unci
n,
cc
ol
o
gy,
an
d evol
u
t
i
ona
r
y cha
ng
e
:
Kl
a
pper
,
(;
,1
98
9,
The Mont
a
gne Noi
r
e Fr
as
ni
an (
Uppe
r Dc}
oni
aj
l
f
Svs
t
el
l
i
at
i
c、八s
s
oci
a
t
i
on s
pe
ci
al
v.
47,
Cl
ar
endon Pr
es
s
,
Oxl
md,
pp. conodont
}
L
I
CCCS
S
I
OI
I
,
i
l
l
Mc
Mi
I
l
an,
N J,
Fmhr
y.
A F and Gl
as
s
,
D J
}
】15-161.
cds
.
,
Devoni
an ol
'
t
hc Wor
l
d:
Ca
nadi
an Soci
et
y of
Pel
r
ol
cur
n Ge
ol
og
i
s
t
s
,
Be
cker
,
R T,
Fei
s
t
,
R,
Fl
ai
s
.
G,
HOI
.
I
s
C}
M R,
and Kl
app
cr
,
G,
1
989,
Menr
oi
r
1
4,
}.
3,
pp.
449-468.
Fr
a
sn
i
a
n/
Fame
r
i
ni
an ext
i
nct
i
(
川 C}ent
s i
n t
h
e DeN
or
l
i
a
l
l
al
( (
川川i
ac,
Kl
apper
,
G.
and Fo}
t
e
r J
r
,
C T,
1
993,
Shap
e anal
ys
i
s of Fr
as
ni
an s
peci
cs of
s
out
hc
r
n Fr
anc
e:
Cor
npl
e,尽endL
I
S C
I
C I
'
Aca
&r
ni
e des Sci
enc
es
,
Pa
r
i
s,
t
h
e Lat
e DcNoni
an c
onodont
vcnus Pal
ma
t
ol
e
pi
s
:
Pal
eont
ol
ogi
ca
l
Scr
.
1
1
,
\下)
( pp.
259-
-
266
Soci
e
t
y.
Memoi
r 32(
J
our
nal
of
Pal
eont
ol
og
y,
v.
67,
no.
4.
}L
l
pp.
)
一
3-
} pp.
Be
cker
,
R T.
and HOL
I
S
'
e,
M R,I
L
)
94,
Kel
l
wa
ss
er eve
nt
s a
nd g
oni
a
t
i
t
c
Lct
hi
ci
s
,「
;
,and Fei
s
t
,
R,
1
991
,La c
r
i
s
e des os
t
r
ac
odc} hent
hi
C
I
L
I
C,
}川
}
t
i
c
cc
ss
i
ons i
n t
he Mor
amune Noi
r
e wi
t
h cor
r
un
ews on p
os
s
i
hl
c pas
s
age Fj
a}
ni
cn-Far
r
i
cnui
an C
I
C COL
I
A
l
l
i
a
c (
NI
ont
aLl
i
c Noi
r
e,
Fi
anc
e
C Al
S
i
t
t
i
ons
:
Cour
i
er For
s
c
hun
gs
i
nS
t
i
(
L
i
t
Scr
i
ckc
r
i
h
er
g,
v.
1
69,
pp.
45-77
met
i
di
on
al
e)
:
COl
np
t
C
S RCF
I
C
I
L
I
S (
I
C I
’
A
ca
demi
c des Sci
enc
es
,
Pa
l
i
s
,
S
c
i
.
Becke‘
一
,R T.
HOL
I
S
C,
M R,
and As
hoUt
i
,/
\ R,
1
988,
Pot
en
t
i
al
s
(
r
a
t
ot
ype
1
1
,
v.
3 1
2,
pp.
1
057-1
063-
J
r
,
G R,
Gi
l
mor
e,
J S}
Or
t
h,
〔一J
,
and Ol
s
en,
F
'
,
1
984,
No
s
ec
t
i
on l
or t
h
e Fr
as
ni
an/
1
-
mi
r
enni
an hounda",
at
El
At
r
ous,
Tal
i
t
a
l
l
. McGhee M orocco: DOCL
I
F
Y
I
CI
TI SUbm i
ned t
o t
he Int
er
nat
i
onal SUbcom mi
ssi
on on
gcoc
her
ni
cal
evi
den
ce f
or al
l
as
t
e
r
oi
dal
i
mpa
ct
at
l
at
e Fl
e%oni
a
n r
na}
s
Dev
oni
an St
r
a
l
i
gr
a
phy,
Rennes
,
1
988-
6 pp.
ext
i
nct
i
on hor
i
zon:
Nal
ui
c,
Lon
don.、308,
pp.
629-63!
.
Beck
er
,
R T.
HOL
I
S
C,
M R,
Ki
i
chga
s
}
cr
,
W 1
'
,
and Pl
a
yf
or
d,
P F
- 1
991,
Mcl
-
a
r
cn,
D J,
a
nd Goodl
'
el
l
ow,
W D.
1
990,
Geol
ogi
c
al
ar
i
d hi
ol
ogi
c
al
Sedi
ment
ar
y a
nd I
'
aUn
al
Chang
es a(
r
os
s t
he Fr
a
s
ni
an/
Famenni
a
n c
ons
e
qu
ence
s 01
gi
a
nt
i
mpa
ct
s:
Annua
l
Revi
e
w Far
t
h ai
r
(
]
Pl
ar
l
el
m-
y
hOUndar
y i
n f
i
l
e Can
ni
ng RaJn ol
'
Wcs
t
cr
i
i
Au
st
r
a
l
i
a:
1
1 i
s
t
oi
i
c
al Sci
enc
es
.
v.]8}
pp.1
23
-1
7]
.
Bi
ol
ogy,
\.
5一pp.
1
83- 1
90
Ni
c
ol
l
,
R S,
and Pl
aNt
or
d,
1
)
F,
1
993,
Upper Devoni
an i
n山Hi
l
l al
l
(
)
I
l
l
al
i
c
s}
conodont zonaii
on and the Frasnian/Famenni
an 1
1
01
-
I
ndarv i
n ti
r
e
Be
cker
,
R 1,
Hous
e,
M R一and Ki
r
chLas
s
以w一
丁,I
99飞,De
voni
a
丁
I9ol
l
i
a
nt
e
bi
os
t
r
a
t
i
D-
aphy a
nd t
i
mi
ng ol
'
I
'
a
ci
cs I
l
l
menl
ent
s i
n t
he Fr
js
ni
an of
t
he Cal
mi
ng Ha
si
n-
Wes
t
er
n Aus
t
i
a
l
i
a:
Pal
a
eogeogr
apl
i
v
Canni
n
g Ba
mn,
Wes
t
er
n Aus
ual
i
a,
i
n f
l
ai
l
wood,
F A,
a
nd Ki
dd,
R B, Pal
ac
ocl
i
n
l
a
I
01
02y,
Pa
l
aC
OCCO]
02},、.1
04,
pp.
1
05-1
1
3
eds
.
,
Hi
gh r
CS
Ol
U
t
i
Ol
l S
t
i
a
l
i
2r
a
phy:
GeOI
O}i
Cdl
SOCI
Ct
}
l
Spe
ci
al
Or
nl
i
s
t
on,
A R,
an
d Kl
a
ppcr
,
G.
1
99-
2,
Pa
l
eo
cl
i
ma
r
c
.
c
ont
r
ok oi
l
Uppe
r
Publ
i
c
at
i
on,
no,
7
0,
pp,
293
-;
-l
?
Devoni
an I
OUr
c
e r
oc
k s
e
qu
enc
es and s
l
a
ck
ed ext
i
nc
t
i
ons:
51
h Nor
t
h Br
ans
on,
E B.
an
d Mehl
,
M Gi
. 1
93
4,
Conodoni
s f
r
om t
h
e Gi
a
s
s
y Cr
eek
Amer
i
can Pa
l
e
ont
ol
ogi
ca
l
Conve
nt
i
on,
Abs
t
r
a
ct
s a
nd Pr
og
r
a
m, Shal
e 01
'M I
SSOUri Uni
ver
si
t
v of Mi
}soui
.
i
St
UdI
CS, V.
8, 1
7 1
-259
Pal
eont
ol
ogi
c
al
So
ci
et
y Sp
eci
a
l
Publ
i
ca
t
i
on no.
6,
p.
2
2T
j
i
mpr
i
nt
1
9331
.
Ruppel
,
S C,
J
ames
,
F W一and Bar
r
i
ck,
J F,
1
993,
F
l
i
gh r
c}
ol
li
f
O[
l
s
t
r
ont
i
L
l
i
n
BL
I
,Li
s
ch,
W,
1
991
,
The gl
ob
al
Fr
a}
ni
an/
Far
r
i
c
nni
an '
Kcl
l
wa
}
}
er Eve
nt
'
i
s
oi
opc c
he
mos
t
r
a
t
i
gr
aphy of
'
t
he Pal
eoz
oi
c Usi
ng con
odont
s:
Geol
ogi
cal
Gcol
ogi
s
che RL
i
nds
chaU,
V.
80
/1
,pp.
49-72.
Soci
et
y(
)
}
、
Ame
r
i
ca,
Abst
r
act
s wi
山 Pi
ogr
ams,
v.
25(
6)
.
pp.
A473-474 BUI
t
VI
l
Ck,
1
'
.
Dr
e
es
cr
i
,
R,
Gr
oc
}
s
ens
,
F,
St
r
uve
,
W,
We
ddi
ge
,
K,
Wer
ner
,
R,
San
dbe
r
g,
C A,
Sc
hi
n
dl
er
.
E .Wal
l
i
s
er
,
0 H, a
nd Zi
egl
er
,
W,
1
990,
a
nd Zi
ei
ler
f
,
W,
N88八r
d
ennes (
Bel
gi
um)
and Fi
l
cl
Hi
l
l
} (
Feder
al Pr
opos
al
f
or t
h
e Fr
as
ni
an/
Pamenni
an }
I
t
St
Ci
I
I
I
I
I
-
L
i
ch SCI
I
I
I
I
i
dt
(
Er
r
s
.
c ar
c
a,
Repu
bl
i
c of
Ger
ma
ny)
:
Con
ner Foi
-
S
CI
l
L
U
I
LS
i
l
l
S
t
i
t
l
i
t
Sc
r
i
ckci
i
bc
q
u,
v.
1
02
-
Kel
l
er
wa
l
d,
Rhei
n.
Schi
ct
er
g
chi
r
ge,
Ger
many)
mi
d ma
t
ci
i
al
f
or t
he pp.
9- 1
55
cor
r
es
pondi
ng f
i
el
d t
r
i
p on Sep
t
e
mbe
r 1
8,
1
990.
Document
Cl
a
c}s,
P,
Ca.
}
i
cl
一‘
J
-
G,
and Ma
i
gol
i
s,
S,
V,
1
992,
Mi
kr
ot
ect
i
t
cs and mas
s
(
UnpUb
l
i
s
he山submi
t
t
ed t
o [
t
i
c Subcommi
ss
i
on on Dcv
oma
n
c\
I
i
n
ct
i
ol
l
s
:
C\i
d
ci
l
cc f
ol;
‘Lat
e Dev
ol
l
i
al
l
as
t
cl
oi
d i
mpac
t
:
Sci
enc
e,
V.
St
r
a
t
i
gr
l
a
phy OCS,
I
UGS)
,
Fr
ankf
ur
t
/
M,
18 pl
)
.
2}
7,
pp.
1
1
02一!川4
Sa
ndbe
r
g,
C A,
Zi
cl
,l
e
r
,
W,
Di
cc
}
cn,
R,
and But
l
er
,
J 1
- 1
988,
La
t
e Fr
a
muan
Copp
er
,
P,
1
98
6 Fr
a
s
ni
a
n/
Fame
nni
a
n ma
s
s ext
i
n
ct
i
on a
l
l
(
]
col
d-Ni
ncr
mas
s ex
t
i
n
ct
i
on:
Conodont
e
ven
t s
t
r
ai
i
}
l
r
a
pf
r
\,
g
dol
i
a
l
c
hang
es an
d pos
}
i
bl
e
oce
ar
i
s:
Geol
m-
,
y,
v.
1
4,
pp.
835- 83
9.
ca
L
I
S
CS
:
COUr
i
er Fot
,
.
,
chL
i
nL
S
i
r
l
s
t
i
t
i
l
t
Sen
cken
her
g,
v.
1
02,
p
p.
263-3
07
Oecem ber /993
441
Sa
nnemann,
D,
1
955,
Bei
t
r
ag z
ur Unt
er
gl
i
ede
r
ung des Obe
r
devons nac
h
Wal
l
i
s
e
r
,
0 H,
Gr
oos
-
Uf
f
enor
de,
H,
Sc
hi
ndl
er
, E,
and Zi
e
gl
e
r
, W,
On t
he
Conodont
e
n:
Ne
ues J
ahr
buc
h f
t
i
r Ge
ol
ogi
e und Pal
bont
ol
ogi
e
,
Uppe
r Kel
l
wa
ss
e
r Hor
i
z
on (
Bounda
r
y Fr
as
ni
an/ Famenni
an)
:
Cour
i
e
r Ab
h
a
n
d
l
u
n
g
e
n
,
v
.
1
0
0
(
3
)
,
p
p.
3
2
4
-3
31
.
For
s
chungs
i
ns
t
i
t
ut Senc
kenbe
r
g,
v. I 1
0,
pp.
247
-255.
Schi
ndl
e
r
,
E,
1
990, Di
e Ke
l
l
wa
ss
er
-
Kr
i
s
e (
hohe Fr
a
s
ne-
St
uf
e,
Obe
r
devon)
:
Wang, K,
1
992, Gl
as
s
y mi
c
r
os
pher
u
l
e
s (
mi
c
r
ot
ekt
i
t
es
) f
r
om an Uppe
r
Gbt
t
i
Dge
r Ar
be
i
t
en z
ur Ge
ol
o
gi
c und Pal
don
t
ol
ogi
e,
v.
46, 11
5 pp. De
voni
a
n l
i
mes
t
one:
Sci
enc
e, v.
256, pp.
1
547-155
0.
Scr
ut
t
on,
C T,
1
988,
Pa
t
t
e
r
ns of e
xt
i
nct
i
on and s
ur
vi
val i
n Pa
l
a
eozo
i
c
Wa
n
g
,
K,
Or
t
h
,
C J
,
At
t
r
e
p J
r
,
M,
Ch
a
t
t
e
r
t
o
n
,
B D E,
Ho
n
g
f
e
i
,
H,
a
n
d
c
o
r
a
l
s
:
S
y
s
t
e
ma
t
i
c
s As
s
o
c
i
a
t
i
o
n S
p
e
c
i
a
l
Vo
l
u
me n
o.
3
4,
p
p
.
6
5
-
8
8
.
Ge
l
ds
e
t
z
er
,
H H J
, 1
991,
Geoch
emi
c
al
evi
den
ce f
or a cat
as
t
r
ophi
c bi
ot
i
c
So
r
auf
,
J E,
a
nd Pedde
r
,
A E H,
1
986,
La
t
e Devoni
an r
u
gos
e cor
a
l
s a
nd t
he
e
ve
nt at t
he Fr
as
ni
an/
Fa
menni
a
n boundar
y i
n s
out
h Chi
na:
Geol
ogy,
v Fr
as
ni
an/
Fa
me
nni
a
n cr
i
s
i
s:
Cana
di
a
n J
our
nal
of Ea
r
t
h Sc
i
ence.
v.
23,
p.
1
9,pp.776-779.
1265-1287.
Wi
l
de,
P,
and Ber
r
y,
W B N,
1
984, Des
t
a
bi
l
i
za
t
i
on of t
he o
cea
ni
c dens
i
t
y
St
e
am,
C W,
1
987, Ef
f
ect
of t
he Fr
a
s
ni
a
n/
Famenni
a
n ext
i
nc
t
i
on ev
ent on
s
t
r
uc
t
ur
e and i
t
s s
i
gni
ic
f
anc
e t
o ma
r
i
ne '
ext
i
nc
t
i
on'
eve
nt
s
:
s
t
r
omat
opor
oi
ds
:
Ge
ol
ogy,
v.
1
5, pp. 677-679.
Pal
ae
ogeogr
a
phy,
Pal
a
eocl
i
mat
ol
ogy,
Pal
a
eoe
col
o
gy,
v.
48
,
pp.
143-162.
Thomps
on,
J B, and Ne
wt
on,
C R,
1
98
9,
La
t
e Dev
oni
a
n mas
s e
xt
i
nc
t
i
on:
epi
s
odi
c cl
i
ma
t
i
c cool
i
ng or war
mi
ng? i
n McMi
l
l
a
n, N J
, Embr
y,
A F Zi
e
gl
e
r
, W,
1
962,
Taxi
onomi
e und Phyl
ogeme Ober
devoni
s
cher
and Gl
as
s
,
D J
,
e
ds
.
,
Devoni
a
n of t
he Wor
l
d:
Cana
di
a
n Soc
i
e
t
y of Conodon
t
e
n und i
br
e s
t
r
at
i
gr
a
phi
s
c
he Be
deut
ung:
Abl
i
andl
un
gen d
es
Pe
t
r
ol
eum Geol
ogi
s
t
s
,
Memoi
r 1
4, v.
3,
pp.
29
-34.
He
ss
i
s
c
he
n Landes
a
mt
e
s f
6r Bodenf
or
s
chung,
v.
38, 1
66 pp.
Va
ngue
s
t
ai
ne, M, DeCl
ai
r
f
a
yt
,
T,
Rouha
r
t
, A,
and Smee
s
t
er
s
,
A,
1
983,
Zi
e
gl
e
r
,
W, 1
971
, Cono
don
t St
r
at
i
gr
a
phy oft
he Eur
ope
an De
voni
an:
Zonat
i
on par a
cr
i
t
ar
chs du Fr
a
s
ni
en Su
p6
r
i
eur
- Fanmenni
an I
nf
6
r
i
e
u
r
Ge
ol
ogi
c
al
Soci
et
y ofA
mer
i
c
a Me
moi
r 12
7,
pp.
227-284.
dans l
es bas
S
i
Ds de Di
nant
, Namur
,
Her
ve et Ca
mpi
ne (
1
36v
oni
e
n
Zi
e
gl
e
r
,
W, and Sandbe
r
g,
C A,
1
990,
The l
a
t
e Devoni
an conodont zo
nat
i
on:
Supd
er
i
eu
r de Bel
gi
qu
e)
:
Annal
es Soci
6
t
6 g6ol
ogi
que de Bel
qi
qu
e
,
v.
Cour
i
e
r For
s
chungs
i
ns
t
i
t
ut Se
nck
enber
g,
v. 1
2 1
,
1
06,pp.1
21
-1
71.
11
5 pp.
口
Dr Gi
l
be
r
t Ma
ppe
r (
t
o
p r
i
gh
t
)
i
s Pr
o
f
e
s
s
o
r o
f Ge
ol
og
y at
t
he Un
i
-
v
e
r
s
i
l
y o
f I
owa,
I
o
wa Ci
t
y
,
I
owa 5
22
42,
US
A.
Hi
s r
e
s
e
ar
c
h c
o
nc
e
n-
t
r
at
e
s o
n Si
l
u
r
i
an and De
v
on
i
an c
o
nod
on
t t
ax
o
no
my an
d bi
os
t
r
at
i
g-
r
a
phy
,
wi
t
h s
pe
ci
al
e
mph
as
i
s on t
h
e Fr
as
ni
an
.
He i
s c
ur
r
e
nt
l
y
i
nv
ol
v
e
d wi
t
h me
t
ho
ds o
fm
or
ph
ome
t
r
i
c a
nal
y
s
i
s o
f co
no
do
nt
s a
nd
t
he
i
r us
e i
n gr
a
phi
c c
or
r
e
l
at
i
on
.
He i
s a Ti
t
ul
ar Me
mb
er o
ft
he S
ub-
c
o
mmi
s
s
i
o
n o
n De
v
oni
an St
r
a
t
i
gr
a
p勺 a
nd was i
t
s Vi
ce
-
Chai
r
man
fr
o
m 1
97
6 t
o 1
9
84
.
Dr Rai
mund Fe
i
s
t (
bo
t
t
om r
i
gh
t
)
i
s Di
r
e
c
t
o
r o
f Re
s
e
ar
c
h wi
t
h t
he
Fr
e
n
c
h Nat
i
on
al
Sc
i
e
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r
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