1399
A
REVISION
OF T H E GENERA
GRANT
D.
OF T H E
HYPODERMATACEAE
1
DARKER
Plant Research Institute, Canada Department of Agriculture, Ottawa, Canada
Received December 2, 1966
A reclassification of the genera and species of the H y p o d e r m a t a c e a e w i t h special
emphasis on the conifer-inhabiting species is presented. T h e 22 genera fall i n t o 7
groups as follows. A. T h e Lirula complex on Abies a n d Picea i n c l u d i n g 3 genera:
Virgella, gen. nov., w i t h rod-shaped spores segregated from Hypoderma, I s t h m i ella, gen. nov., w i t h bifusiform spores from Bifusella, a n d L i r u l a , gen. nov., w i t h
clavate spores from Hypodermella. B. T h e Hypodermella complex comprising
Hypodermella v . T u b . , subcuticular o n Larix, l i m i t e d to the t y p e species; a group
of 3 subepidermal genera o n Pinus i n c l u d i n g Davisomycella, gen. nov., w i t h
clavate spores, Ploioderma, gen. nov., w i t h rod-like spores, a n d Elytroderma
D a r k e r w i t h 2-celled spores; a n d a s u b h y p o d e r m a l genus, Lophodermella v . H o h n .
on Pinus. C. T h e Bifusella complex w i t h Bifusella v . H o h n . o n Pinus, Duplicaria
F c k l . on Empetrum, a n d a new subepidermal genus, Soleella, o n Pinus. D. T h e
Hypoderma complex comprising Hypoderma D C . ex S a i n t A m a n s emend. D e N o t . ,
M e l o d e r m a , gen. nov., a segregate from Hypoderma h a v i n g Lophodermium
affinities, Lophodermium C h e v . , and Lophomerum Ouellette a n d M a g a s i , a recent
phragmosporous segregate from Lophodermium. E. T h e genus Coccomyces D e N o t .
F. T h e Colpoma complex w i t h Colpoma W a l l r . , Xyloschizon S y d . , a n d Bifusepta
D a r k e r . G. T h e Rhytisma complex w i t h Rhytisma F r . , Placuntium v . H o h n . , a n d
Nymanomyces P . H e n n .
Introduction
T h e P h a c i d i a l e s i n m o d e r n classifications c o m p r i s e a r a t h e r heterogeneous
order of A s c o m y c e t e s w i t h s t r o m a t i c a s c o l o c u l a r ascocarps e m b e d d e d i n a n d
connate w i t h t h e host tissues a b o v e a n d b e l o w . T h e i r a s c i are u n i t u n i c a t e .
T h r e e families, the C r y p t o m y c e t a c e a e , P h a c i d i a c e a e , a n d H y p o d e r m a t a c e a e
are recognized b y v o n A r x a n d M u l l e r (1954).
T h e H y p o d e r m a t a c e a e differ f r o m t h e o t h e r t w o f a m i l i e s i n s e v e r a l m a r k e d
c h a r a c t e r i s t i c s . I n general, h o w e v e r , t h e y are d i s t i n g u i s h e d b y t h e i r ascocarps
w h i c h possess a s h i e l d - l i k e c o v e r i n g l a y e r w h i c h opens b y a n e l o n g a t e d s l i t ,
except i n the genus Coccomyces, w h e r e the o p e n i n g s are o r t e n d to be r a d i a l .
A m o r e o r less definite o p e n i n g m e c h a n i s m is present i n s o m e species, w h i l e i n
others the o p e n i n g is m e r e l y f o r t u i t o u s , b y means of a m e c h a n i c a l t e a r i n g of the
fungus a n d host tissues. T h e h y m e n i a l l a y e r is u s u a l l y m o r e o r less c u p - l i k e a n d
flat or t e n d s to b e c o m e so a t m a t u r i t y . T h e s u b h y m e n i u m is u s u a l l y p l e c t e n c h y m a t o u s w i t h cells e l o n g a t e d , m o r e o r less p a r a l l e l w i t h t h e base of t h e a p o t h e c i u m a n d o c c u p i e s the b u l k of t h e space b e t w e e n t h e h y m e n i u m a n d t h e b a s a l
s t r o m a t i c tissue. T h e b a s a l s t r o m a is p s e u d o p a r e n c h y m a t o u s a n d c o m p o s e d of
m o r e or less i s o d i a m e t r i c cells w h i c h f o r m a tissue i n c o n t a c t w i t h a n d p e n e t r a t i n g the i n n e r host cells. I n m o s t species of H y p o d e r m a t a c e a e o n conifer needles
the b a s a l s t r o m a is r e l a t i v e l y r e d u c e d i n t h i c k n e s s , whereas i n species s u c h as
Colpoma quercinum, i t is f a i r l y s h a r p l y d e l i m i t e d a n d of c o n s i d e r a b l e t h i c k n e s s .
T h e asci of the H y p o d e r m a t a c e a e range f r o m b r o a d l y s a c c a t e t o c l a v a t e t o
a l m o s t c y l i n d r i c a l , a n d m a y e v e n be r a t h e r l o n g - s t a l k e d , as i n t h e genus
Hypoderma.
T h e p o r e of the ascus n e v e r t u r n s b l u e w i t h i o d i n e . T h e ascospores
v a r y f r o m b a c i l l a r t o fusoid o r b i f u s i f o r m , t o c l a v a t e o r c y l i n d r i c a l - f i l i f o r m , a n d
f r o m one-celled t o p h r a g m o s p o r o u s , a n d are s u r r o u n d e d b y a g e l a t i n o u s s h e a t h .
A l l are h y a l i n e w i t h the e x c e p t i o n of Nymanomyces,
w h i c h has b r o w n spores.
' C o n t r i b u t i o n N o . 573 from the P l a n t Research Institute.
Canadian Journal of Botany. Volume 45 (1967)
1400
CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
T h e p y c n i d i a of t h e H y p o d e r m a t a c e a e , w h e r e k n o w n , f a l l i n t o t h e f a m i l y
Leptostromataceae.
I n t h e o l d e r classifications of t h e H y p o d e r m a t a c e a e few c h a r a c t e r s h a v e been
a c c e p t e d o n a> p r i m a r y generic basis except t h e gross a p p e a r a n c e of t h e ascocarp.
T h i s has s e r v e d m e r e l y t o segregate t h e Coccomyces, Colpoma, a n d
Rhytisma
complexes f r o m t h e l a r g e b u l k of less c o n s p i c u o u s H y p o d e r m a t a c e a e w h i c h has
been f u r t h e r s u b d i v i d e d o n the basis of spore shape, size, a n d s e p t a t i o n . I n the
o l d e r schemes of c l a s s i f i c a t i o n t h i s v i r t u a l l y a m o u n t e d t o a p r i m a r y s e p a r a t i o n
o n spore s h a p e a l o n e . C o n s e q u e n t l y these species h a v e been p l a c e d a l m o s t
w i t h o u t e x c e p t i o n i n t h e f o u r genera Lophodermium,
Hypoderma,
Hypodermella,
a n d Bifusella. M a n y of o u r present difficulties m a y be a s c r i b e d t o t h e c o n t i n u e d
a t t e m p t s of w o r k e r s t o assign n e w species to these o l d g e n e r a w h i l e d i s r e g a r d i n g
t h e m o r p h o l o g i c a l n a t u r e of s u c h s t r u c t u r e s as ascocarps, a s c i , a n d p y c n i d i a .
O u r present d i l e m m a stems f r o m t h e fact t h a t , as t h e genera h a v e become
larger, o v e r l a p p i n g i n spore c h a r a c t e r s has o c c u r r e d , a n d r e l a t i o n s h i p s b e t w e e n
g r o u p s of species h a v e b e c o m e l o s t i n the m u l t i p l i c i t y of species w h i l e c e r t a i n
o b v i o u s s i m i l a r i t i e s i n o t h e r s t r u c t u r e s h a v e been neglected o r m i n i m i z e d .
N e v e r t h e l e s s , t h e g r a d u a l a c c r e t i o n of k n o w l e d g e has b r o u g h t t o l i g h t a n u m b e r
of r e l a t i o n s h i p s w h i c h i t is n o l o n g e r p o s s i b l e t o i g n o r e , a n d t h e c u r r e n t s t u d y
has g r o w n o u t of a desire t o a t t e m p t t h e c r e a t i o n of m o r e w o r k a b l e species
g r o u p i n g s of the H y p o d e r m a t a c e a e , w i t h s p e c i a l e m p h a s i s o n the c o n i f e r i n h a b i t i n g species. A s a n e x a m p l e , t h e a c c e p t a n c e o f v o n H o u n d ' s genus
Lophodermella,
w i t h s l i g h t e m e n d a t i o n , a p p e a r s l o n g o v e r d u e . F u r t h e r m o r e , i t is
p r o p o s e d t o segregate c e r t a i n o t h e r species o r g r o u p s of species i n t o n e w g e n e r a
w h o s e c i r c u m s c r i p t i o n s a r e b a s e d p r i m a r i l y o n c o m b i n a t i o n s of c h a r a c t e r s o t h e r
t h a n spore shape.
Historical Review
T h e h i s t o r y o f t h e c l a s s i f i c a t i o n of t h e H y p o d e r m a t a c e a e has been o u t l i n e d
s e v e r a l t i m e s i n recent y e a r s . F o r t h e m o r e d e t a i l e d a c c o u n t s t h e w o r k s of
B i s b y (1923), D a r k e r (1932), T e h o n (1935), a n d T e r r i e r (1942) s h o u l d be
c o n s u l t e d . I n r e v i e w i t is sufficient t o r e c a l l t h a t t h e species of H y p o d e r m a t a c e a e
k n o w n p r i o r t o t h e F r i e s i a n s t a r t i n g d a t e , 1 J a n , 1821, for t h e F u n g i C a e t e r i i n
t h e I n t e r n a t i o n a l C o d e of B o t a n i c a l N o m e n c l a t u r e , were p l a c e d m a i n l y i n t h e
genus Hysterium
F r i e s (1823). O n l y t h r e e genera of t h e H y p o d e r m a t a c e a e i n
use t o d a y were r e c o g n i z e d d u r i n g t h e p e r i o d o f F r i e s ' S y s t e m a M y c o l o g i c u m
a n d E l e n c h u s F u n g o r u m , 1 8 2 1 - 1 8 3 2 : Hypoderma
D C . , 1805, w h i c h w a s
a c c e p t e d b y G r a y , M 6 r a t , a n d S a i n t - A m a n s i n 1821, Rhytisma
F r . , 1823, a n d
Lophodermium
C h e v . , 1826.
A s o r i g i n a l l y d e n n e d b y D e C a n d o l l e i n L a m a r c k a n d D e C a n d o l l e i n 1805,
a n d f u r t h e r e x p a n d e d i n 1815, t h e genus Hypoderma D C . w a s c r e a t e d for those
species o f Hysterium
a n d related forms g r o w i n g under a n d b r e a k i n g through
t h e e p i d e r m i s (used i n a b r o a d sense) a t m a t u r i t y . I t is o f interest t o n o t e t h a t
u n d e r Hypoderma a n d Rhytisma, w h i c h he t r e a t e d u n d e r t h e n a m e Xyloma, D e
C a n d o l l e h a d i s o l a t e d f a i r l y c o m p l e t e l y t h e t h e n k n o w n species w h i c h w e r e l a t e r
to b e c o m e t h e H y p o d e r m a t a c e a e . U n d e r Hypoderma
D C . w e r e segregated
species, m o s t o f w h i c h w e r e l a t e r t r a n s f e r r e d t o Lophodermium,
Colpoma,
Hypodermella,
a n d p o s s i b l y Duplicaria,
t o g e t h e r w i t h o n e species of Leptostroma, a n d one species of Hysterographium,
H. fraxini, w h i c h t e n d s t o r e m a i n
DARKER: REVISION OF T H E HYPODERMATACEAE
1401
red d u r i n g e a r l y d e v e l o p m e n t a n d t h u s n o d o u b t f i t t e d i n t o t h e c o n c e p t of
C a n d o l l e ' s genus. C h e v a l l i e r (1826) e s t a b l i s h e d Lophodermium
for essenIlv the s a m e g r o u p of o r g a n i s m s t r e a t e d b y D e C a n d o l l e u n d e r
Hypoderma
d i n c l u d e d s i x s t i l l c u r r e n t l y a c c e p t a b l e Lophodermium
species, one Hypoderella* ^
HyP°d
species, b u t a c c o r d i n g t o F r i e s , E l e n c h u s 2 : 1 4 5 , 1 8 2 8 ,
C h e v a l l i e r assigned t o Lophodermium
herbarum C h e v . s p e c i m e n s referable to
jjypoderma commune F r . a n d n o t t o Hysterium
herbarum F r . A l t h o u g h b o t h
Hypoderma a n d Lophodermium
were v a l i d l y p u b l i s h e d d u r i n g t h e F r i e s i a n
oeriod, 1821-1832, F r i e s h i m s e l f c o n t i n u e d t o t r e a t t h e i r species m a i n l y u n d e r
Hysterium. U n f o r t u n a t e l y , Lophodermium
is i l l e g i t i m a t e (see d i s c u s s i o n b e l o w ) .
T w o o t h e r D e C a n d o l l e species, Hypoderma
quercinum a n d H. crispum,
were
transferred b y W a l l r o t h (1833) t o his n e w genus Colpoma.
r?
a
anc
two
erma
T h e genera Hypoderma
a n d Lophodermium
were n o t c l e a r l y c i r c u m s c r i b e d
u n t i l D e N o t a r i s (1847) f i n a l l y segregated t h e i r species. I n t h e s a m e p u b l i c a t i o n
he established t h e n e w genus Coccomyces based o n Hysterium
tumidum F r . , b u t
b y a c u r i o u s c h o i c e he a t t a c h e d t h r e e r a d i a t e l y d e h i s c i n g species n o w g e n e r a l l y
placed i n Coccomyces t o t h e first s e c t i o n of the genus Lophodermium.
T h e first o f
these, r e n a m e d Lophodermium
phacidium
D e N o t . , is a n o b l i g a t e s y n o n y m o f
phacidium
coronatum F r . Coccomyces tumidum a n d P . coronatum are s y n o n y mous a n d t h e t y p e of Coccomyces is c o n s i d e r e d t o be C. coronatus ( S c h u m . ex
F r . ) D e N o t . , t h u s c i t e d l a t e r i n D e N o t . , E r b a r . c r i t t . i t a l . 1: 2 3 6 , 1 8 5 9 .
D u r i n g t h e l o n g i n t e r v a l b e t w e e n t h e w o r k of D e N o t a r i s a n d t h e s t u d i e s o f
von H o h n e l i n 1917 o n l y three m o r e genera of H y p o d e r m a t a c e a e w e r e d e s c r i b e d .
These c o m p r i s e d : Duplicaria
F c k l . , 1870, Hypodermella
v . T u b . , 1895, a n d
Nymanomyces
P . H e n n . , 1899.
V o n H o h n e l (1917a) erected a n e w order, t h e P h a c i d i a l e s v . H o h n . , c o m p o s e d
of t h e o l d f a m i l i e s P h a c i d i a c e a e a n d H y p o d e r m a t a c e a e w i t h w h i c h he u n i t e d a
n u m b e r of genera f r o m o t h e r families. H e t h e n a r b i t r a r i l y s u b d i v i d e d t h e o r d e r
i n t o six f a m i l i e s , p r i m a r i l y o n t h e basis of t h e p o s i t i o n of t h e f r u i t i n g b o d y i n t h e
s u b s t r a t u m . T h i s r e s u l t e d i n t h e s p l i t t i n g of m a n y of t h e o l d genera i n t o t w o o r
even t h r e e genera a n d r e q u i r e d t h e c r e a t i o n of a n u m b e r of n e w n a m e s . N o
a t t e m p t w a s m a d e t o d i s t r i b u t e a l l of t h e k n o w n species i n t o t h e genera of t h e
newly created skeletal framework although types were established for the new
genera. I n v a r i o u s papers d u r i n g 1917 v o n H o h n e l r e e s t a b l i s h e d o r c r e a t e d t h e
f o l l o w i n g genera: Lophodermina
a n d Lophodermellina
from
Lophodermium;
Hypodermellina
f r o m Hypoderma;
Bifusella
a n d Placuntium
from
Rhytisma;
a n d Lophodermella
f r o m Hypodermella.
Bifusella
v. Hohn. and
Placuntium
E h r e n b g . ex v . H o h n . h a v e g a i n e d general a c c e p t a n c e a m o n g t h e H y p o d e r m a taceae b y m o s t m y c o l o g i s t s , a n d i t is the belief of t h e w r i t e r t h a t t h e a c c e p t a n c e
of Lophodermella v . H o h n . is l o n g o v e r d u e . O f t h e r e m a i n d e r , N a n n f e l d t (1932)
c o n s i d e r e d t h e genus Hypodermellina
v . H o h n . t o be a m e m b e r of t h e H e m i s phaeriales w h i l e v o n A r x a n d M u l l e r (1954) t r e a t e d i t u n d e r t h e P h a c i d i a c e a e .
O n the o t h e r h a n d t h e genera Lophodermina
v . H o h n . a n d Lophodermellina
v.
H o h n . h a v e g e n e r a l l y been rejected a l t h o u g h t h e y a r e b o t h v a l i d l y p u b l i s h e d
t a x a a n d as s u c h are a v a i l a b l e i n the e v e n t t h a t s o m e o n e w i s h e s t o use t h e m
w i t h a n e m e n d e d c i r c u m s c r i p t i o n . A l t h o u g h v o n H o h n e l ' s i n s e r t i o n of the
H y p o d e r m a t a c e a e i n t h e P h a c i d i a l e s w a s a d o p t e d i n a m u c h r e s t r i c t e d sense b y
N a n n f e l d t (1932) u n d e r the n a m e P h a c i d i a c e a e , v o n H o h n e l ' s use of t h e p o s i t i o n
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CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
of t h e f r u i t - b o d y i n the h o s t tissue as a p r i m a r y basis for the s e p a r a t i o n of the
P h a c i d i a l e s v . H o h n . i n t o families has n o t g a i n e d r e c o g n i t i o n b y m o s t m y c o l o gists.
V o n H o h n e l d i d succeed, h o w e v e r , i n f o c u s i n g a t t e n t i o n o n t h e need for s t u d y
of t h i s large a n d heterogeneous g r o u p . H e , h i m s e l f (19176, c ) , b r o u g h t further
o r d e r t o t h e P h a c i d i a l e s w h e n he r e m o v e d his first f a m i l y , the S c h i z o t h y r i a c e a e ,
t o the T h r a u s m a t o p e l t i n e a e T h e i s s e n of t h e H e m i s p h a e r i a c e a e T h e i s s e n .
D u r i n g the f o l l o w i n g y e a r , v o n H o h n e l (1918) p o i n t e d o u t the s h a r p d e m a r c a t i o n e x i s t i n g b e t w e e n the H y p o d e r m a t a c e a e a n d the H y s t e r i a c e a e w h i c h u p t o
t h a t t i m e h a d been t r e a t e d t o g e t h e r u n d e r t h e H y s t e r i a l e s . I n t h i s p a p e r he
r e m o v e d t h e H y s t e r i a c e a e t o the S p h a e r i a l e s a n d m o r e r e c e n t l y M u l l e r a n d v o n
A r x (1962) h a v e t r a n s f e r r e d t h e m to the D o t h i o r a l e s . P e t r a k (1947) r e s t r i c t e d
v o n H o h n e l ' s s e c o n d f a m i l y t o Leptopeltella v . H o h n . a n d Leptopeltis v . H o h n .
a n d a d d e d t h r e e n e w genera. T h i s f a m i l y has been a c c e p t e d a n d r e v i s e d b y
E . M u l l e r a n d v o n A r x (1962) a n d b y v o n A r x (1964) u n d e r t h e s l i g h t l y
m o d i f i e d n a m e , L e p t o p e l t a c e a e , a f a m i l y w h i c h t h e y a t t a c h to the D o t h i o r a l e s
also.
T e h o n (1935) m a d e t h e o n l y serious a t t e m p t t o a p p l y the p r i n c i p l e s o u t l i n e d
b y v o n H o h n e l . A l t h o u g h i n his studies he professed to f o l l o w v o n H o h n e l ' s
s c h e m e of c l a s s i f i c a t i o n , one is left w i t h the i m p r e s s i o n t h a t he c o u l d n o t q u i t e
b r i n g h i m s e l f to a c c e p t i t i n i t s e n t i r e t y . O t h e r w i s e , his k e y to genera w o u l d
h a v e reflected t h e p r i m a r y s p l i t t i n g of t h e genus a n d the s e p a r a t i o n of i t s n e w
c o m p o n e n t s a m o n g three of v o n H o h n e l ' s s i x families. I n s t e a d , he first d i v i d e d
t h e species of Lophodermium
i n t o t w o groups o n the basis of the presence o r
absence of a l i f o r m tissue a n d o n l y s e c o n d a r i l y o n the p o s i t i o n of the h y s t e r o t h e c i u m i n the host tissue a n d m a d e n o m e n t i o n of the d i s t r i b u t i o n of his genera
i n v o n H o h n e l ' s families. A n a n a l y s i s of his k e y shows t h a t a l i f o r m tissue is
present i n a l l s u b c u t i c u l a r species, i n some i n t r a e p i d e r m a l species, b u t i n n o
s u b e p i d e r m a l species. T h e r e is w i t h i n t h e H y p o d e r m a t a c e a e a r a t h e r definite
c o r r e l a t i o n b e t w e e n the d e p t h of i n s e r t i o n of the h y s t e r o t h e c i u m a n d its c o l o r .
I n general, s u b c u t i c u l a r species are b l a c k , w h i l e c e r t a i n deep-seated species are
a l m o s t colorless, a n d a p p a r e n t l y the a b i l i t y to detect the a l i f o r m tissue is
s o m e w h a t d e p e n d e n t o n c o l o r . O n e s u b e p i d e r m a l species o n Pinus
strobus
w h i c h has a c o m p a r a t i v e l y t h i n e p i d e r m i s shows a l i f o r m tissue q u i t e w e l l
d e v e l o p e d . T h e q u e s t i o n arises as t o w h e t h e r T e h o n w o u l d h a v e c r e a t e d a n e w
genus for t h i s h a d he o b s e r v e d i t . I n t h e 1935 p u b l i c a t i o n T e h o n segregated
three n e w genera, Dermascia
f r o m Lophodermium,
a n d Epidermella
and
Locelliderma
f r o m Hypoderma. T h e s e t h r e e genera a n d a l l n e w species d e s c r i b e d
b y h i m i n t h i s p a p e r are i n v a l i d since t h e y d o n o t c o m p l y w i t h a r t i c l e 43 of t h e
I n t e r n a t i o n a l C o d e of B o t a n i c a l N o m e n c l a t u r e w h i c h requires a L a t i n d e s c r i p t i o n for v a l i d p u b l i c a t i o n . M o r e o v e r , his n e w c o m b i n a t i o n s u n d e r Dermascia are
l i k e w i s e i n v a l i d since t h e y were m a d e i n a n i n v a l i d genus. A l l of his n e w
c o m b i n a t i o n s u n d e r Lophodermina,
Lophodermellina,
a n d Dermascia h a v e been
t r e a t e d u n d e r t h e i r o r i g i n a l names i n Lophodermium
b y most workers. H i s new
species i n these t h r e e genera, h o w e v e r , h a v e been transferred b y N a n n f e l d t
(1936) a n d T e r r i e r (1942) to the genus Lophodermium
where, w i t h the exception
of Lophodermina
septata T e h o n , r e c e n t l y v a l i d a t e d as Lophomerum
septatum
O u e l l e t t e , t h e y s t i l l r e m a i n i n v a l i d a w a i t i n g L a t i n d e s c r i p t i o n s . T h e basis for
I
DARKER: REVISION OF T H E HYPODERMATACEAE
1403
the s e p a r a t i o n of Epidermella
T e h o n f r o m Hypoderma
was never clearly stated
b y its a u t h o r a n d as t h e genus is i n v a l i d l y p u b l i s h e d i t s species s h o u l d be
treated u n d e r t h e i r e s t a b l i s h e d names i n Hypoderma.
O n the other hand, the
d e s c r i p t i o n of his monospecific genus Locelliderma,
based o n
Hypoderma
mpslodesmi
C e s . o n t h e grass, Ampelodesmos
tenax, suggests t h a t t h e fungus
p r o b a b l y belongs o u t s i d e t h e H y p o d e r m a t a c e a e , p o s s i b l y i n o r n e a r t h e genus
Scirrhia. I t s h o u l d be r e e x a m i n e d . M e a n w h i l e , i t s o n l y v a l i d n a m e is Hypoderma
ampelodesmi C e s .
I n a l a t e r paper, T e h o n (1939) d e s c r i b e d s e v e r a l n e w species, a l l a c c o m p a n i e d
b y L a t i n d e s c r i p t i o n s . O n e of these, h o w e v e r , Epidermella
hansbroughii T e h o n ,
is i n v a l i d because i t was p l a c e d i n a n i n v a l i d genus. I t m a y be c a l l e d Hypoderma
hansbroughii ( T e h o n ) ex D a r k e r s p . n o v . [ = Epidermella
hansbroughii
T e h o n , M y c o l o g i a 3 1 : 688. 1939]. T e h o n ' s L a t i n d e s c r i p t i o n w i l l serve t o
validate it i n its new position.
N a n n f e l d t (1932), i n h i s e x t e n s i v e s t u d y of t h e i n o p e r c u l a t e D i s c o m y c e t e s ,
treated a p p r o x i m a t e l y . . 5 8 genera of v o n H o h n e l ' s P h a c i d i a l e s a n d , i n a d d i t i o n ,
a b o u t 30 o t h e r related genera. A f t e r r e m o v i n g t h e d i s c o r d a n t elements elsewhere
a n d r e l e g a t i n g a n u m b e r of others t o s y n o n y m y , t h e f a m i l y P h a c i d i a c e a e sensu
N a n n f e l d t , w a s r e d u c e d t o 14 genera, m o s t l y h y p o d e r m a t a c e o u s i n c h a r a c t e r .
I n a r e e x a m i n a t i o n of t h i s f a m i l y , T e r r i e r (1942) d i v i d e d i t i n t o t h r e e f a m i l i e s :
the r e v i v e d f a m i l y H y p o d e r m a t a c e a e c o m p o s e d of 10 genera a n d t h e R h y t i s maceae w i t h 3 genera together m a k i n g u p t h e o r d e r H y p o d e r m a t a l e s , a n d t h e
P h a c i d i a c e a e sensu stricto w i t h 5 genera. F i n a l l y v o n A r x a n d M u l l e r (1954) a n d
M u l l e r a n d v o n A r x (1962) r e a l i g n e d u n d e r t h e P h a c i d i a l e s 3 f a m i l i e s , t h e
P h a c i d i a c e a e w i t h 8 genera, t h e C r y p t o m y c e t a c e a e w i t h 2 genera, a n d t h e
H y p o d e r m a t a c e a e i n c l u d i n g T e r r i e r ' s R h y t i s m a c e a e w i t h 11 genera. I n t h e
H y p o d e r m a t a c e a e t h e y i n c l u d e d : Bifusella v . H o h n . , 1 9 1 7 ; Clithris ( F r . ) , 1 8 2 2 ;
Coccomyces D e N o t . , 1847; Duplicaria
F c k l . , 1 8 7 0 ; Elytroderma
D a r k e r , 1932;
Hypoderma D C . ex S a i n t A m a n s e m e n d . D e N o t . , 1847; Hypodermella
v. Tub.,
1895; Lophodermium
C h e v . , 1 8 2 6 ; Nymanomyces
P . H e n n . , 1899;
Placuntium
E h r e n b g . ex v . H o h n . , 1917; a n d Rhytisma F r . , 1823. T o these s h o u l d be a d d e d
Lophodermella v . H o h n . , 1917, Xyloschizon S y d . , 1922, k n o w n o n l y t o t h e w r i t e r
t h r o u g h t h e o r i g i n a l d e s c r i p t i o n , a n d t w o r e c e n t l y d e s c r i b e d genera,
Bifusepta
D a r k e r , 1963, a n d Lophomerum
O u e l l e t t e a n d M a g a s i , 1966.
a
T h e h y p o d e r m a t a c e o u s f u n g i h a v e been assigned o v e r t h e years t o v a r i o u s
groups of A s c o m y c e t e s . I f w e r e t u r n t o D e N o t a r i s (1847) w e find t h a t he p l a c e d
t h e m i n t h e P y r e n o m y c e t e s A s c i g e r i , a n d i n c l u d e d t h e m as a n i n t e g r a l p a r t of
the H y s t e r i a c e i , w h i c h he f u r t h e r s u b d i v i d e d o n t h e basis of c o l o r i n t o t h e
P h a e o s p o r i i a n d t h e H y a l o s p o r i i . R e h m (1887) e l a b o r a t e d t h e c l a s s i f i c a t i o n of
the H y s t e r i a c e a e C d a . , i n c l u d i n g t h e H y p o d e r m i e a e , a n d a g a i n (1912) u n d e r t h e
H y s t e r i i n e a e S c h r o t . w h i c h h e t r e a t e d as s y n o n y m o u s w i t h t h e H y s t e r i a c e a e
C d a . a n d intermediate between the Discomycetes a n d the Pyrenomycetes. V o n
H o h n e l (1917) u n i t e d t h e H y p o d e r m a t a c e a e w i t h t h e P h a c i d i a c e a e i n t h e
P h a c i d i a l e s v . H o h n . w h i c h h e r e g a r d e d as D i s c o m y c e t e s a p p r o a c h i n g t h e
Dothideales o n the one h a n d a n d the Pezizales o n the other. I n the following
y e a r , 1918, he m a d e t h e first c l e a r - c u t d i s t i n c t i o n b e t w e e n t h e H y p o d e r m a taceae a n d t h e H y s t e r i a c e a e a n d u n i t e d t h e l a t t e r w i t h t h e L o p h i o s t o m a c e a e i n
a n e w f a m i l y , t h e H y s t e r o s t o m a c e a e of t h e P y r e n o m y c e t e s . N a n n f e l d t (1932)
1404
CANADIAN JOURNAL OF BOTANY. VOL. 45. 1967
assigned t h e P h a c i d i a c e a e sensu N a n n f e l d t t o t h e H e l o t i a l e s . T e r r i e r (1942)
p l a c e d his o r d e r H y p o d e r m a t a l e s c o m p o s e d of t h e H y p o d e r m a t a c e a e a n d the
R h y t i s m a c e a e u n d e r t h e A s c o h y m e n i a l e s of N a n n f e l d t a n d r e g a r d e d t h i s o r d e r
as b e i n g r e l a t e d t o N a n n f e l d t ' s D r e p a n o p e z i z o i d e a e of t h e D e r m a t e a c e a e i n t h e
H e l o t i a l e s . T h e P h a c i d i a c e a e , o n t h e o t h e r h a n d , w e r e t r e a t e d as A s c o l o c u l a r e s
w i t h affinities t o t h e D o t h i o r a c e a e . G a u m a n n (1949) suggested t h a t the
H y p o d e r m a t a c e a e h a d a r i s e n p h y l o g e n e t i c a l l y from t h e P l e c t a s c a l e s b y w a y of
t h e D e r m a t e a c e a e . I n a d i s t i n c t l y different c l a s s i f i c a t i o n , L u t t r e l l (1951, 1955)
c o n s i d e r e d the P h a c i d i a c e a e t o be H e l o t i a l e s b e l o n g i n g i n the series U n i t u n i catae, subseries D i s c o m y c e t e s , g r o u p I n o p e r c u l a t i . V o n A r x a n d M u l l e r (1954),
w h i l e q u e s t i o n i n g t h e t r u e d i s c o m y c e t o u s n a t u r e of the P h a c i d i a l e s , p r o v i sionally placed the order between the Dothiorales a n d the Helotiales, a n d
c o n s i d e r e d t h e m t o b e u n i t u n i c a t e as p o i n t e d o u t b y L u t t r e l l b u t w i t h a l l
f a m i l i e s b e l o n g i n g to the A s c o h y m e n i a l e s i n t h e sense of N a n n f e l d t . E . M u l l e r
a n d v o n A r x (1962) t r e a t e d t h e P h a c i d i a l e s as a d i s t i n c t o r d e r of t h e D i s c o m y cetes w h i c h l i k e the H e l o t i a l e s h a d a r i s e n f r o m a pezizaceous ancestor. G a u m a n n
l a t e r (1964) suggested t h a t t h e ancestors o f t h e P h a c i d i a l e s are t o be s o u g h t
a m o n g the s i m p l e P y r o n e m a c e a e o r A s c o c o r t i c i a c e a e , t h a t t h e y h a v e d e v e l o p e d
p a r a l l e l w i t h the P e z i z a l e s a n d H e l o t i a l e s b u t i n d e p e n d e n t l y , a n d t h a t t h e y d o
n o t b e l o n g i n t h e D i s c o m y c e t e s . F u r t h e r m o r e , he n o t e d t h a t the ascocarps of
t h e P h a c i d i a l e s d e v e l o p i n a m a n n e r s i m i l a r t o those of t h e D o t h i o r a l e s ; y e t , he
b e l i e v e d t h a t t h e t w o orders were n o t c l o s e l y r e l a t e d because of t h e u n i t u n i c a t e
a s c i of the f o r m e r a n d t h e b i t u n i c a t e a s c i of the l a t t e r . H e b e l i e v e d t h e t e r m s
a s c o h y m e n i a l a n d a s c o l o c u l a r t o h a v e o n l y d e s c r i p t i v e v a l u e a n d to be u n i m p o r t a n t i n c l a s s i f i c a t i o n . I n the H y p o d e r m a t a c e a e , w h i c h he c o n s i d e r e d t o be
a s c o l o c u l a r , he n o t e d t h a t c e r t a i n species a t m a t u r i t y a p p e a r to be a s c o h y m e n i a l . I n a s i m i l a r m a n n e r he c o n s i d e r e d p a r a p h y s o i d s a n d p a r a p h y s e s t o
i n t e r g r a d e . I n a recent p u b l i c a t i o n w i t h t a x o n o m i c i m p l i c a t i o n s G o r d o n (1966)
r e p o r t e d the o c c u r r e n c e of 3 b a s i c a n d 2 i n t e r m e d i a t e t y p e s of c e n t r u m o n t o g e n y
a m o n g the H y p o d e r m a t a c e a e a n d i n t h i s p a p e r discussed 2 b a s i c a n d 1 i n t e r mediate type i n detail.
Morphological Characters
T h e s t r o m a t a m a y range f r o m s i m p l e m o n o a s c o c a r p i c t y p e s a f r a c t i o n of a
m i l l i m e t e r i n d i a m e t e r t o large p o l y a s c o c a r p i c s t r u c t u r e s u p t o 4 c m o r m o r e i n
d i a m e t e r . T h e a s c o c a r p itself, s t r i c t l y s p e a k i n g , occupies a l o c u l e w i t h i n a
s t r o m a a n d a p p e a r s as a s m a l l c u p u l e w i t h i t s associated s u r r o u n d i n g tissues
f i r m l y a t t a c h e d t o t h e s t r o m a . F o r purposes o f d i s c u s s i o n reference to the
a s c o c a r p w i l l be a s s u m e d to i n c l u d e t h e c l o s e l y a t t a c h e d s t r o m a t i c tissues as
w e l l . I n t h e d e s c r i p t i v e p o r t i o n of t h i s w o r k the t e r m h y s t e r o t h e c i u m has been
used i n reference t o t h e s i m p l e r ascocarps w h i c h s p l i t o p e n b y a l o n g i t u d i n a l
s l i t . A l t h o u g h t h e t e r m m o r e p r o p e r l y refers t o the fruit b o d y of a m e m b e r of t h e
H y s t e r i a c e a e i t s use has been c o n t i n u e d here as a m a t t e r of c o n v e n i e n c e s i n c e
t h e r e a p p e a r s to be n o o t h e r s u i t a b l e d e s c r i p t i v e t e r m a v a i l a b l e .
T h e ascocarps t h e m s e l v e s v a r y i n s h a p e s u p e r f i c i a l l y f r o m s m a l l , a l m o s t
c i r c u l a r forms l i k e Lophodermium
sphaerioides to e x t r e m e l y e l o n g a t e d , m o r e o r
less s t r a i g h t f o r m s l i k e L. ponderosae a n d the n e r v i s e q u i o u s f r u i t bodies o n
Abies a n d Picea, o r t o e l o n g a t e d , m o r e o r less c u r v e d o r b r a n c h e d forms l i k e
those of c e r t a i n Colpoma species, o r t o c o m p a r a t i v e l y large r o u n d o r p o l y g o n a l
DARKER: REVISION OF T H E HYPODERMATACEAE
1405
ocarps l i k e those of Coccomyces. A l l o f these are u s u a l l y c o n s i d e r e d m o n o scocarpous, a n d a p p a r e n t l y consist of u n i l o c u l a r s t r o m a t a . W h e t h e r these d o
a c t u a l l y represent i n d i v i d u a l ascocarps is o p e n t o q u e s t i o n a n d requires m o r e
d e t a i l e d s t u d y , b u t t h e y are enclosed i n a c o m m o n s t r o m a t i c s t r u c t u r e a n d , to
the u n a i d e d eye, s h o w w h a t a p p e a r s to be a s i n g l e h y m e n i a l surface. I n t h e socalled p o l y a s c o c a r p i c f o r m s w h i c h i n c l u d e species of Placuntium,
Nymanoyces, a n d Rhytisma, w i t h t h e e x c e p t i o n of R. punctatum,
t h e l a r g e , m o r e o r less
c o m m o n s t r o m a opens i n t o n u m e r o u s h y m e n i a l surfaces s e p a r a t e d b y s t e r i l e
tissue. I n Placuntium
t h e s t r o m a m o s t f r e q u e n t l y r u p t u r e s t o expose a large
elongated o r c i r c u l a r h y m e n i u m w i t h sterile i s l a n d s of s t r o m a t i c tissue s u p porting portions of the d a r k cover scattered through it.
I n cross-sectional v i e w t h e ascocarps v a r y f r o m species l i k e
Lophodermium
juniperinum
w h i c h b u l g e o u t w a r d s f r o m t h e h o s t tissue t o species l i k e Hypodermella sulcigena w h i c h are d e e p l y i m m e r s e d a n d e x t e r n a l l y s h o w l i t t l e e v i dence of t h e i r presence.
L e n g t h , b r e a d t h , a n d w i d t h of t h e ascocarps are c h a r a c t e r i s t i c i n the
m o r p h o l o g y of i n d i v i d u a l species.
C o l o r of t h e ascocarps is c o n s t a n t u n d e r fixed c o n d i t i o n s a n d v a r i e s f r o m
b l a c k i n t h e m o r e e x t e r n a l l y s i t u a t e d a s c o c a r p s , s u c h as those o f
Lophodermium
nitens a n d L. juniperinum,
t h r o u g h v a r i o u s shades of b r o w n i n c e r t a i n nerv i s e q u i o u s species o n Abies to a l m o s t colorless f o r m s i n t h e deep-seated ascocarps of Hypodermella
concolor a n d r e l a t e d species. C o l o r is u n d o u b t e d l y d u e to
c h e m i c a l reactions w h i c h d a r k e n the fungus tissues o r t h e associated host
tissues, a n d is d e f i n i t e l y c o n c e r n e d w i t h the p r e v e n t i o n of w a t e r loss. T h e
e x t e r n a l p a r t s of the a s c o c a r p are more affected. W h e n m o i s t u r e s u p p l i e s are
a d e q u a t e the need for t h i s w a t e r b a r r i e r d i s a p p e a r s . A s m a l l c o l l e c t i o n of
Lophodermium
pinastri
o n Pinus
resinosa s h o w s t h i s v e r y s t r i k i n g l y . O n
needles c a u g h t i n a s m a l l r o c k depression filled w i t h w a t e r , t h e u p p e r c l y p e a l
area r e m a i n e d p r a c t i c a l l y colorless a n d a p p e a r e d p a r e n c h y m a t i c , r a t h e r t h a n
p s e u d o p a r e n c h y m a t i c a n d c a r b o n o u s as i n ascocarps w h i c h h a d d e v e l o p e d
n o r m a l l y o n the forest floor.
m
T h e p o s i t i o n of t h e a s c o c a r p , t h a t i s , t h e d e p t h of i n s e r t i o n i n t h e h o s t
tissues, a p p e a r s t o be a c o n s t a n t feature for each species. V o n H o h n e l e v e n
m a d e i t t h e basis for d i v i d i n g his P h a c i d i a l e s i n t o f a m i l i e s a n d T e h o n used i t as
a basis for generic separations i n his t r e a t m e n t of t h e genus Lophodermium.
I t is
t r u e t h a t i n s e v e r a l instances groups of species s h o w t h e s a m e host p o s i t i o n a n d
i n t h e present p a p e r a few s u c h g r o u p s h a v e b e e n i s o l a t e d as genera b u t i n n o
instance was this considered the p r i m a r y factor i n the separation. A n excellent
e x a m p l e of the c o n s t a n c y of the p o s i t i o n of t h e a s c o c a r p i n t h e host tissues is t o
be f o u n d i n Hypoderma desmazierii w h i c h m a i n t a i n s t h e s a m e h o s t p o s i t i o n a n d
other characters w h e n transferred b y i n o c u l a t i o n from its n o r m a l N o r t h
A m e r i c a n host, Pinus strobus, t o t h e h a r d pines, Pinus resinosa a n d P. banksiana.
T h e c l y p e a l l a y e r f o r m s t h e u p p e r c o v e r i n g of t h e a s c o c a r p a n d o n t h e o u t e r
s i d e is u n i t e d w i t h t h e host tissue. I t is v a r i a b l e i n c o l o r , as m e n t i o n e d a b o v e ,
a p p a r e n t l y d e p e n d i n g s o m e w h a t o n t h e d e p t h of i n s e r t i o n i n t h e h o s t tissue.
A t m a t u r i t y t h e c l y p e u s s p l i t s l e n g t h w i s e t o expose t h e h y m e n i u m . T h i s m a y be
v e r y b r o a d l y exposed w i t h the c l y p e u s f o l d i n g b a c k , as i n t h e n e r v i s e q u i o u s
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CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
species o n Abies a n d Picea, o r i t m a y be v e r y n a r r o w l y exposed b y a n e l o n g a t e d
s l i t o r pore, as i n m a n y Hypoderma a n d Lophodermium
species. I n s o m e species,
for e x a m p l e Hypoderma
desmazierii,
a n d i n c e r t a i n s t r a i n s of
Lophodermium
pinastri, the s t r o m a t i c c l y p e u s , s o m e t i m e s c o m b i n e d m a r g i n a l l y w i t h the b a s a l
tissues, m a y e x t e n d w e l l b e y o n d t h e area of t h e i n n e r a s c o c a r p h y m e n i u m f o r m i n g a s o r t of shelf of d a r k s t e r i l e tissue. E v e n i n species s u c h as
Hypodermella
ampla, i n w h i c h t h e presence of a b a s a l l a y e r is m o r e difficult to d e m o n s t r a t e ,
there is a w e l l m a r k e d s u b e p i d e r m a l tissue e x t e n d i n g b e y o n d t h e h y m e n i a l area.
W i t h i n the c l y p e u s s p e c i a l i z e d tissues m a y d e v e l o p w h i c h a c t as a n o p e n i n g
m e c h a n i s m . I n t h e i r h i g h e s t d e v e l o p m e n t these tissues are c o m p o s e d of a l o n g
b a n d of p a r e n c h y m a t o u s tissue e x t e n d i n g t h e l e n g t h of the a s c o c a r p . T h e
a c t u a l o p e n i n g proceeds f r o m t h e o u t s i d e as a n o t c h e d fissure a n d as i t d e v e l o p s
the p a r e n c h y m a cells elongate i n t o the fissure as filaments, often septate, a n d
e x t e n d i n g a t r i g h t angles t o t h e l i n e of dehiscence. T e r m i n a l l y these filaments,
the " p e r i p h y s e s " of T e h o n , a r e m o r e o r less s w o l l e n a n d gelatinous a n d f o r m t h e
edges of the t w o l i p s of the o p e n a s c o m a . T h e g e l a t i n o u s , p a r e n c h y m a t i c s l i t
b a n d aids n o t o n l y i n t h e dehiscence of the a s c o c a r p b u t also serves t o seal off
t h e h y m e n i u m u n d e r a d v e r s e c o n d i t i o n s . T h e r e are a m o n g species of the
H y p o d e r m a t a c e a e v a r i o u s m o d i f i c a t i o n s i n t h e s t r u c t u r e of t h e o p e n i n g m e c h a n i s m . I n s o m e species t h e c o v e r is m u c h t h i c k e n e d i n the l i p r e g i o n a n d as t h e
n o t c h e d fissure d e v e l o p s f r o m t h e o u t s i d e t o w a r d s t h e i n n e r c a v i t y t h e d a r k
e x c i p l e flattens o u t . T h i s flattening m a y be f u r t h e r a c c e n t u a t e d b y areas of m o r e
o r less p a r e n c h y m a t o u s tissue o n t h e i n n e r face o f t h e c o v e r o n each side of t h e
t h i c k e n e d l i p s . A s these areas s w e l l , pressure is exerted a n d the c o v e r i n g tissues
are s t r e t c h e d further. S u c h e x t r e m e d e v e l o p m e n t s m a y be o b s e r v e d i n a n u m b e r
of Lophodermium
species w h e r e i t a i d s i n k e e p i n g the h y m e n i u m c o v e r e d a l m o s t
u n t i l m a t u r i t y . O n t h e o t h e r h a n d , t h e o p e n i n g m e c h a n i s m m a y be n o t h i n g
m o r e t h a n a s m a l l core of p a r e n c h y m a t o u s tissue l o c a t e d s o m e w h e r e a l o n g t h e
m i d d l e of the c o v e r i n g l a y e r . T h r o u g h t h i s p a r e n c h y m a the r u p t u r e e a s i l y t a k e s
p l a c e as i n Lophodermium
berberidis i l l u s t r a t e d b y N a n n f e l d t (1932) or i n
Hypodermella
ampla. D e v e l o p m e n t a l s t u d i e s m a y e v e n t u a l l y s h o w s o m e f o r m of
p r e f o r m e d s t r u c t u r e i n t h e m a j o r i t y of t h e H y p o d e r m a t a c e a e a l t h o u g h a t
present a n u m b e r o f species are c o n s i d e r e d t o l a c k a n y o p e n i n g m e c h a n i s m
w h i c h foreshadows t h e s l i t a n d i n s u c h instances t h e o p e n i n g t a k e s place m o r e
o r less f o r t u i t o u s l y a l o n g the l i n e of greatest stress. I n a few species, t h e r e is a
d e f i n i t e t h i n n i n g of t h e e x c i p l e a l o n g the t o p of t h e a s c o m a , f o r m i n g a s o r t o f
f a u l t - l i n e a l o n g w h i c h the o p e n i n g o c c u r s .
A m o n g the c h a r a c t e r s w h i c h a fungus s h o u l d possess for i n c l u s i o n i n t h e
H y p o d e r m a t a c e a e T e h o n s t i p u l a t e d t h e presence of a n o s t i o l a r l i n i n g of
p e r i p h y s e s . U n f o r t u n a t e l y , m a n y of T e h o n ' s o w n species fail t o m e e t t h i s test.
F u r t h e r m o r e , t h e use of the t e r m p e r i p h y s e s i n T e h o n ' s m e a n i n g is u n f o r t u n a t e
s i n c e t h e y h a v e n o t h i n g to d o w i t h t h e t r u e p e r i p h y s e s o c c u r r i n g i n the P y r e n o m y c e t e s . I n t h e l a t t e r the p e r i p h y s e s d e v e l o p w i t h i n a p e r i t h e c i u m as o u t g r o w t h s from t h e p e r i t h e c i a l w a l l a n d f o r m a l i n i n g of t h e u p p e r w a l l s a n d often
e x t e n d i n t o t h e necks of b e a k e d p e r i t h e c i a . I n t h e H y p o d e r m a t a c e a e the p e r i p h y s i s - l i k e s t r u c t u r e s are o n l y associated w i t h t h e o p e n i n g m e c h a n i s m .
T h e b a s a l l a y e r f o r m e d after t h e s p l i t t i n g o f t h e s t r o m a d u r i n g d e v e l o p m e n t
m a y c o n s i s t of n o t h i n g m o r e t h a n a t h i n tissue a d j a c e n t t o t h e i n n e r host
DARKER: REVISION OF T H E HYPODERMATACEAE
1407
issues a n d c o n t i n u o u s w i t h the fungus cells w h i c h h a v e p e n e t r a t e d a n d m o r e o r
less b r o k e n d o w n t h e host cells as, for e x a m p l e , i n Lophodermium.
I n a few
era
s Colpoma, i t m a y be h i g h l y o r g a n i z e d . S o m e t i m e s t h e r e is a
d a r k e n i n g of t h e cells of t h e b a s a l l a y e r . C l o s e e x a m i n a t i o n , h o w e v e r , m a y
reveal t h a t t h i s d a r k e n e d a r e a is p a r t i a l l y d u e t o t h e presence of d i s c o l o r e d host
cells. T h e t h i c k n e s s of the b a s a l l a y e r c a n be m o s t e a s i l y d e t e r m i n e d a t the
angles of t h e ascocarps a t the m a r g i n s a t t h e p o i n t of u n i o n w i t h the c l y p e u s .
T h e angles t h e m s e l v e s are o f t e n rilled w i t h loose p a r e n c h y m a t i c tissue, o r
c o l l e n c h y m a , or w i t h p r i s m a t i c tissue e x t e n d i n g f r o m t h e c l y p e u s t o t h e b a s a l
tissue.
T h e a s c o c a r p p r o p e r lies b e t w e e n t h e c l y p e a l a n d b a s a l s t r o m a t i c tissues. A t
the angles w h e r e these t w o tissues meet, t h e s u b h y m e n i a l tissues m a y be o b served b e n d i n g u p a l o n g the sides of the c l y p e u s . T e r r i e r (1942) c a l l e d s p e c i a l
a t t e n t i o n t o t h e c u p u l e n e s t l i n g i n the s t r o m a of Colpoma quercinum a n d s u g gested t h a t i t offers the best e v i d e n c e for the s t r o m a t i c n a t u r e of t h e f r u i t i n g
bodies of the H y p o d e r m a t a c e a e . W h i l e n o t so o b v i o u s , t h e s a m e s t r u c t u r e is
easily o b s e r v e d i n m a n y of the species of the f a m i l y .
A s c o g e n o u s tissue d e v e l o p s i n a flat or c u r v e d z o n e , u s u a l l y near the l o w e r side
of the a s c o m a l tissues b e t w e e n the c l y p e u s a n d the b a s a l l a y e r , a n d f r o m t h i s
zone arise t h e a s c i a n d p a r a p h y s e s to f o r m the h y m e n i u m . B e l o w the h y m e n i u m
is the p l e c t e n c h y m a t o u s s u b h y m e n i a l l a y e r c o m p o s e d of a c o m p a c t f i l a m e n t o u s
tissue w h o s e m a r g i n s t e n d t o c u r v e u p w a r d s t o f o r m t h e w a l l of the a s c o m a . O n
its o u t e r face i t merges s o m e w h a t i n t o t h e c l y p e u s w h i l e i n w a r d l y i t merges
i m p e r c e p t i b l y i n t o the h y m e n i u m .
T h e a s c i are s i m p l e u n i t u n i c a t e s t r u c t u r e s a n d m a y be b r o a d a n d s o m e w h a t
saccate o r c l a v a t e o r r a t h e r d e f i n i t e l y c y l i n d r i c a l , a n d w i t h o u t b l u e i n g of the
ascus t i p b y i o d i n e . I n s o m e genera t h e y b e c o m e d i s t i n c t l y s t a l k e d w i t h the
ascospores g a t h e r e d i n the u p p e r c l a v a t e e n d . I n size t h e asci of the H y p o d e r m a taceae range r o u g h l y f r o m a b o u t 50 t o 250 p, i n l e n g t h a n d a b o u t 5 t o 45 p i n
w i d t h . T h e spores are e i g h t i n n u m b e r , o r i f four, w i t h f o u r a b o r t e d spores
u s u a l l y r a t h e r c l e a r l y e v i d e n t . T h e scolecosporous f o r m s are u s u a l l y fascicled
b u t t h e r e is a t e n d e n c y for " s i s t e r " spores t o be s o m e w h a t p a i r e d . T h i s p a i r i n g
m a y be o b s e r v e d w h e r e t h e p a i r s t e n d t o s l i p d o w n i n t o the ascus g i v i n g the
ascus t i p a s o m e w h a t a s y m m e t r i c a l a p p e a r a n c e . A m o n g the s h o r t e r s p o r e d
species t h e r e is o f t e n a b i s e r i a t e a r r a n g e m e n t of t h e spores w h i c h are a g a i n
f r e q u e n t l y i n p a i r s a l t h o u g h b a s a l l y the a r r a n g e m e n t m a y be m o r e o r less
u n i s e r i a t e . I n Hypoderma
hedgcockii the four spores w h i c h d o n o t a b o r t are
a r r a n g e d u n i s e r i a t e l y w h i l e the four a b o r t e d spores are s c a t t e r e d a b o u t i r r e g u larly.
s
u
c
n
a
A s c o s p o r e s m a y be s h o r t , r o d - l i k e , f u s i f o r m , r a t h e r b r o a d l y e l l i p t i c a l ,
c l a v a t e , b i - t o t r i f u s i f o r m , o r l o n g needle-shaped, a n d are s u r r o u n d e d b y a
g e l a t i n o u s s h e a t h . I n e a r l y d e v e l o p m e n t the spores of the l o n g - s p o r e d species
m a y be c o i l e d i n the ascus b u t these u s u a l l y s t r a i g h t e n o u t as t h e ascus m a t u r e s
a n d elongates further. T h e spores are u s u a l l y h y a l i n e , a l t h o u g h i n
Placuntium
a n d c e r t a i n Hypoderma
species t h e y m a y a p p e a r s l i g h t l y y e l l o w i s h , a n d i n
Nymanomyces
t h e y are d i s t i n c t l y b r o w n . W i t h i n t h e f a m i l y , ascospores v a r y i n
size f r o m a b o u t 20 p. i n l e n g t h t o 160 p, o r m o r e a n d f r o m a b o u t 1 p t o 18 p. i n
w i d t h . M o s t g e n e r a h a v e s i m p l e amerospores b u t i n Hypoderma t h e t e n d e n c y is
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CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
t o b e c o m e t w o - c e l l e d , w h i l e t w o - c e l l e d spores are t h e r u l e i n
Elytroderma.
S e v e r a l genera w i t h s i m p l e , c o n t i n u o u s ascospores are best d e s c r i b e d as
scolecosporous. O t h e r s , n o t a b l y Lophomerum,
Bifusepta,
and
Xyloschizon,
possess p h r a g m o s p o r o u s scolescospores. S p o r e g e r m i n a t i o n i n the a m e r o s p o r o u s
a n d scolecosporous t y p e s is u s u a l l y preceded b y s e p t a t i o n n e a r the center of t h e
spore. G e r m i n a t i o n t h e n proceeds f r o m a n y p a r t o f t h e spore as i n t h e d i d y m o s p o r o u s a n d p h r a g m o s p o r o u s t y p e s . I n d i s t i l l e d w a t e r t h e spores send o u t
s i m p l e g e r m tubes w h i c h m a y f o r m s w e l l i n g s i n t o w h i c h the p r o t o p l a s m g a t h e r s .
T h e s e s w e l l i n g s m a y be c u t off b y a cell w a l l f r o m the g e r m t u b e a n d f r o m t h e
s w o l l e n area a n o t h e r g e r m t u b e m a y be s e n t o u t . T h i s m a y be r e p e a t e d t w o o r
t h r e e t i m e s as t h o u g h the spore were s e a r c h i n g for a s t o m a t a l e n t r a n c e . I n
Bifusella faullii a n d Hypodermella
mirabilis the d i s t i n c t i v e p a l m a t e s t r u c t u r e s
f o r m e d a t the ends of t h e g e r m t u b e s a r e p r o b a b l y a n o t h e r m e t h o d for f i n d i n g
e n t r a n c e i n t o the host.
T h e p a r a p h y s e s are u s u a l l y s i m p l e , o c c a s i o n a l l y once b r a n c h e d n e a r t h e t i p ,
s e p t a t e , f i l a m e n t o u s , s t r a i g h t o r s o m e t i m e s m o r e o r less c o i l e d a t t h e t i p o r
sometimes thickened, a n d usually surrounded b y a t h i n gelatinous sheath
w h i c h m a y i n s o m e species b e c o m e r a t h e r p r o m i n e n t . O c c a s i o n a l l y , e s p e c i a l l y
n e a r the base, anastomoses b y s i m p l e b r i d g i n g m a y t a k e place b e t w e e n a d j a c e n t
p a r a p h y s e s . T h e p a r a p h y s e s often a p p e a r t o exceed t h e asci i n l e n g t h , a l t h o u g h
at m a t u r i t y the asci usually project up through the paraphyses following a more
o r less r a p i d e l o n g a t i o n .
T h e p y c n i d i a o f the H y p o d e r m a t a c e a e are m e m b e r s of the L e p t o s t r o m a taceae a n d h a v e been assigned t o s u c h genera as Leptostroma,
Leptothyrium,
Leptothyrella,
Hypodermina,
Labrella,
Melasmia,
Colpomella, a n d
Crandattia,
a l t h o u g h i t is d o u b t f u l t h a t a n y t r u e Leptothyrium
o r Leptothyrella species w i l l
be f o u n d to b e l o n g here. T h e p y c n i d i a c o n s i s t of s i m p l e c u s h i o n s o f p a r e n c h y m a t i c tissue f r o m w h i c h arise a p a l i s a d e - l i k e tissue of s p e r m a t i o p h o r e s b e a r i n g
s i m p l e b a c i l l a r s p e r m a t i a . T h e o n l y species a t present k n o w n to possess a
definite c o v e r i n g l a y e r o v e r t h e p y c n i d i a are Leptostroma berberidis N a n n f . , t h e
species of Bifusella as redefined b e l o w , a n d Placuntium
andromedae ( P e r s . ex
F r . ) v . H o h n . T h e p y c n i d i a m a y be s m a l l , s i m p l e , m o r e o r less r o u n d e d , d a r k
s t r u c t u r e s l i k e those of Leptostroma pinastri D e s m . , o r e l o n g a t e d , o f t e n m o r e o r
less c o n t i n u o u s , s o m e w h a t c o n c o l o r o u s s t r u c t u r e s w h i c h s o m e t i m e s t e n d t o
d a r k e n after s p e r m a t i a l d i s c h a r g e l i k e t h e p y c n i d i a o f t h e " n e r v i s e q u i o u s "
species of H y p o d e r m a t a c e a e o n Abies, o r t h e y m a y be q u i t e n u m e r o u s a n d t e n d
t o coalesce b e c o m i n g m o r e o r less l a b y r i n t h i n e o r effused as i n t h e i m p e r f e c t
s t a t e of Bifusella
linearis
(Pk.) v . H o h n . , or small, simple, elliptical and
c o n c o l o r o u s a n d n e v e r d a r k e n i n g as i n t h e p y c n i d i a of Hypodermella
ampla
( D a v i s ) D e a r n . , o r l a r g e , a p p r o a c h i n g t h e a s c o c a r p i n size, a n d s h i n i n g b l a c k as
i n t h e p y c n i d i a of Hypoderma pini ( D e a r n . ) D a r k e r . T h e s p e r m a t i a l states o f
Coccomyces, Rhytisma,
a n d Colpoma are g e n e r a l l y r e p o r t e d as f a l l i n g i n t o t h e
f o r m genera Leptothyrium,
Melasmia,
a n d Colpomella r e s p e c t i v e l y . F o r t h e
m o s t p a r t , t h e p y c n i d i a h a v e t h e s a m e p o s i t i o n i n t h e host tissues as h a v e t h e i r
perfect states. I t s h o u l d be n o t e d , h o w e v e r , t h a t i n Lophodermium
pinastri, w i t h
ascocarps s u b c u t i c u l a r i n t h e c e n t e r a n d s u b e p i d e r m a l a t t h e m a r g i n s , the
p y c n i d i a are s u b e p i d e r m a l , whereas a m o n g t h e " n e r v i s e q u i o u s " species o n
Abies, i n c e r t a i n species w i t h ascocarps i n t r a e p i d e r m a l i n the c e n t e r a n d
s u b e p i d e r m a l a t the m a r g i n s , t h e p y c n i d i a are i n t r a e p i d e r m a l .
DARKER: REVISION OF T H E HYPODERMATACEAE
1409
T h e spermatiophores are borne a t the tips of somewhat swollen, often pearshaped cells a t t h e surface o f t h e b a s a l p a r e n c h y m a t o u s c u s h i o n . T h e s m a l l
n e r m a t i a are c u t off a c r o g e n o u s l y f r o m t h e t i p s o f t h e s p e r m a t i o p h o r e s , p r o b a b l y i n succession j u d g i n g f r o m t h e v a s t n u m b e r s p r o d u c e d , a l t h o u g h t h i s p o i n t
has n e v e r been d e f i n i t e l y e s t a b l i s h e d . D i s p e r s a l is u s u a l l y b y m e a n s o f a l e n g t h wise s l i t o r b y a r u p t u r e o f t h e m a r g i n o f t h e p y c n i d i u m . I n c e r t a i n o f t h e
" n e r v i s e q u i o u s " species o n Abies t h e s p e r m a t i a m a y emerge t h r o u g h t h e s t o m a t a l openings a n d effuse o v e r t h e surface o f t h e needle. V e r y o c c a s i o n a l l y i n
Bifusella faullii,
c i r r h i of spermatia m a y be observed s t a n d i n g above the
s t o m a t a after m a i n t e n a n c e i n m o i s t c h a m b e r s i n t h e l a b o r a t o r y . I n s i z e t h e
s p e r m a t i a v a r y f r o m a b o u t 0.5-1.2 n i n w i d t h i n different species a n d i n l e n g t h
from a b o u t 3 . 0 - 3 . 5 n i n Bifusella
faullii
t o 1 2 - 2 0 n i n Bifusella
linearis.
T h e p y c n i d i a m a t u r e 1 y e a r after i n f e c t i o n i n t h e " n e r v i s e q u i o u s " species o n
Abies, whereas t h e ascocarps r e q u i r e 2 y e a r s t o r e a c h m a t u r i t y . I n t h e m a j o r i t y
of H y p o d e r m a t a c e a e , h o w e v e r , t h e p y c n i d i a a p p e a r 1 o r 2 y e a r s after i n f e c t i o n
a n d a r e c l o s e l y f o l l o w e d b y t h e ascocarps d u r i n g t h e s a m e season.
I n a d d i t i o n t o m i c r o s c o p i c s t u d i e s , field o b s e r v a t i o n s o n gross m o r p h o l o g y o f
the fungus, age o f diseased needles, a n d s o o n , a r e a l l a h e l p f u l p a r t i n t h e
c o m p l e t e c i r c u m s c r i p t i o n of a n i n d i v i d u a l species. N o d o u b t a l l of t h e H y p o d e r m a t a c e a e a r e m o r e o r less p a r a s i t i c , a l t h o u g h t h e degree o f p a r a s i t i s m is
e x t r e m e l y difficult t o assess i n v o l v i n g as i t does n o t o n l y t h e r e l a t i v e suscept i b i l i t y o f t h e h o s t w h i c h m a y v a r y f r o m p l a n t t o p l a n t a n d t h e degree o f
v i r u l e n c e of t h e fungus itself b u t also t h e effects d u e t o v a r i a t i o n of t h e e n v i r o n m e n t a l factors a s w e l l . I t is n o t s u r p r i s i n g , therefore, t h a t i t is r a t h e r d i f f i c u l t t o
r e p r o d u c e t h e diseases caused b y these f u n g i i n a r t i f i c i a l i n f e c t i o n e x p e r i m e n t s .
W e i r (1916) w a s successful i n b r i n g i n g a b o u t i n f e c t i o n w i t h h i s Hypoderma
deformans. D a r k e r (1932) r e p o r t e d s a t i s f a c t o r y i n f e c t i o n e x p e r i m e n t s w i t h
Hypodermella
laricis v . T u b . a n d H. concolor ( D e a r n . ) D a r k e r , b u t w i t h m o s t
others tested t h e results h a v e p r o v e d u n s a t i s f a c t o r y o r e r r a t i c . I n s p i t e o f t h i s ,
m o s t Hypodermella
(sensu lato) species, i n d e e d m o s t o f t h e species t r e a t e d u n d e r
the first n i n e genera b e l o w , a p p e a r t o be m o r e o r less s t r o n g l y p a r a s i t i c a n d
m a t u r e o n y o u n g needles in situ o n t h e t w i g s , often o c c u r r i n g o n p a r t i a l l y green
needles o r o n d e a d needles s c a t t e r e d a m o n g o t h e r green needles w h i c h n o r m a l l y
c o n t i n u e t o r e m a i n a l i v e for o n e o r s e v e r a l seasons m o r e . T y p i c a l o f a n o t h e r
g r o u p of H y p o d e r m a t a c e a e a r e t h e c o n i f e r - i n h a b i t i n g species o f
Lophodermium
w h i c h a r e u s u a l l y present o n o l d e r needles a n d c o m p l e t e t h e i r m a t u r a t i o n o n t h e
g r o u n d after t h e c h a r a c t e r i s t i c n e e d l e - c a s t i n g , o n s l a s h , o r o n m e c h a n i c a l l y
i n j u r e d b r a n c h e s . F e w careful o b s e r v a t i o n s h a v e been m a d e o n t h e a g e o f t h e
needles i n v o l v e d o r o n t h e age of t h e needles a t t h e t i m e o f t h e i r n o r m a l f a l l . I t
is o b v i o u s , h o w e v e r , t h a t a m o n g c o n i f e r - i n h a b i t i n g species t h e r e a p p e a r t o be
t w o m a i n g r o u p s o f species, o n e o f w h i c h , c o m p r i s i n g m o s t o f t h e species o f t h e
first n i n e genera t r e a t e d b e l o w , is d e f i n i t e l y p a r a s i t i c o n y o u n g needles a n d
m a t u r e s after 1 o r 2 years o n needles in situ o n t h e t w i g s , a n d a n o t h e r w h i c h
a p p e a r s t o be b u t w e a k l y p a r a s i t i c a n d u s u a l l y requires 2 y e a r s t o d e v e l o p ,
o f t e n m a t u r i n g m o r e o r less s a p r o p h y t i c a l l y o n t h e g r o u n d after o v e r w i n t e r i n g
o n t h e f a l l e n needles o r o n needles w e a k e n e d b y s o m e e n v i r o n m e n t a l f a c r o r t o
b y mechanical injury.
O n e p h e n o m e n o n w h i c h s h o u l d b e c i t e d is t h e a s s o c i a t i o n o f c e r t a i n s e c o n d a r y
f u n g i s o m e t i m e s present o n needles p r i m a r i l y a t t a c k e d b y species o f H y p o -
1410
CANADIAN
JOURNAL
O F BOTANY.
V O L . 45.
1967
d e r m a t a c e a e . T h e s e fungi, a b o u t 30 i n n u m b e r , o c c u r o n l y o n host tissues k i l l e d
b y specific species of H y p o d e r m a t a c e a e a n d a p p a r e n t l y n o w h e r e else. T h e y
d e v e l o p o n l y after t h e host food s u p p l y has been p r e p a r e d for t h e m b y t h e
p r i m a r y fungus a n d a p p a r e n t l y are n o t p a r a s i t i c o n e i t h e r t h e h o s t p l a n t o r the
p r i m a r y p a r a s i t e . Naevia piniperda f o l l o w i n g after Lophodermium
macrosporum,
Nothophacidium
phyllophilum
after Hypodermella
nervata, a n d Pezizella
minuta
after Hypoderma lethale, are t y p i c a l e x a m p l e s . A few f u n g i f o l l o w after m o r e t h a n
one species of H y p o d e r m a t a c e a e , b u t for t h e m o s t p a r t the r e l a t i o n s h i p is r a t h e r
specific. Lophodermium
autumnale,
h o w e v e r , follows after Bifusella faullii a n d
Hypodermella
mirabilis
i n eastern N o r t h A m e r i c a a n d after
Hypoderma
robustum i n t h e west. S e v e r a l of the associated f u n g i m a y a c c o m p a n y t h e s a m e
species of H y p o d e r m a t a c e a e a l t h o u g h the s e c o n d a r y i n f e c t i o n i n a n y one a t t a c k
is l a r g e l y l i m i t e d to a single species. T h u s Lophodermium
autumnale,
Stegopezizella balsameae, o r Leptosphaeria
faullii
a n d i t s i m p e r f e c t state, Coniothyrium faullii,
m a y be f o u n d i n a s s o c i a t i o n w i t h Bifusella faullii.
F r o m the
e c o n o m i c v i e w p o i n t these s e c o n d a r y f u n g i are s o m e t i m e s e x t r e m e l y i m p o r t a n t .
B y r o b b i n g t h e p r i m a r y fungus o f i t s f o o d s u p p l y t h e y p r e v e n t the l a t t e r f r o m
f r u i t i n g a n d t h u s a c t as n a t u r a l b i o l o g i c a l c o n t r o l agents. A s a n e x a m p l e ,
d u r i n g t h e h e i g h t of i t s d e v e l o p m e n t i n 1961, a c o l l e c t i o n ( D A O M 88009) of
Hendersonia
pinicola W e h m . , s e c o n d a r y after Hypodermella
concolor, w a s so
a b u n d a n t t h a t t h e b l a c k masses of spores h u n g i n festoons o n the needles o f
Pinus banksiana a n d b l a c k e n e d the green leaves of v e g e t a t i o n b e l o w . D u r i n g
the f o l l o w i n g y e a r n o e v i d e n c e of e i t h e r the p r i m a r y o r the s e c o n d a r y species
c o u l d be f o u n d a t t h e site of t h e e a r l i e r i n f e c t i o n near N o r t h O n s l o w , P o n t i a c
C o u n t y , Q u e b e c . A p p a r e n t l y the H. concolor i n f e s t a t i o n i n t h i s s m a l l a r e a w a s
c o m p l e t e l y w i p e d o u t . A r e l a t e d species was d e s c r i b e d o n Pinus strobus u n d e r
the n a m e Hendersonula pinicola D e a r n . I t s p y c n i d i a o c c u r o n l y o n p i n e needles
k i l l e d b y Bifusella linearis ( P k . ) v . H o h n . w h e r e t h e y m a y b r e a k t h r o u g h the
s t r o m a t i c c r u s t of the Bifusella species o r t h r o u g h o t h e r p o i n t s i n the e p i d e r m i s
where the c r u s t is absent. T h e o r i g i n a l d e s c r i p t i o n of D e a r n e s s is o b v i o u s l y a
c o m p o s i t e one d e r i v e d i n p a r t f r o m a Hendersonia
species a n d i n p a r t f r o m
Bifusella
linearis. T h e i m m e r s e d n a t u r e of the p y c n i d i a a n d t h e absence of a
s t r o m a places t h i s fungus i n Hendersonia
and not i n Hendersonula. Since,
h o w e v e r , the specific e p i t h e t is p r e e m p t e d b y Hendersonia
pinicola
Wehm.,
Hendersonia dearnessii, n o m . n o v . is p r o p o s e d for Hendersonula
pinic
D e a r n . , M y c o l o g i a 20: 236. 1928 (non Hendersonia pinicola W e h m . , M y c o l o g i a
38: 3 1 2 . 1 9 4 6 ) .
Suggested Lines of Revision
T h e classification of the H y p o d e r m a t a c e a e d e v e l o p e d s l o w l y o v e r the y e a r s
a n d b e c a m e m o r e o r less fixed l a r g e l y a l o n g t h e lines of S a c c a r d o ' s a n d S p e g a z z i n i ' s general s c h e m e of classification o n the basis of spore shape, c o l o r , a n d
s e p t a t i o n . A l t h o u g h o t h e r c h a r a c t e r s h a v e been used i n a b r o a d sense t o
segregate Coccomyces a n d t h e genera of the Rhytisma
c o m p l e x f r o m t h e rem a i n d e r of the H y p o d e r m a t a c e a e , i t has r a r e l y h a p p e n e d t h a t spore c h a r a c t e r s
h a v e p l a y e d s u c h a p r i m a r y role i n the classification of a f a m i l y . U s u a l l y these
characters h a v e been used s e c o n d a r i l y a n d i n large families m a y h a v e been used
o v e r a n d o v e r a g a i n . N e v e r t h e l e s s , as l o n g as the n u m b e r of species r e m a i n e d
s m a l l , spore characters, e s p e c i a l l y shape, s e r v e d f a i r l y w e l l to d e l i m i t c e r t a i n
D A R K E R : REVISION O F T H E H Y P O D E R M A T A C E A E
1411
era. U n f o r t u n a t e l y , as t h e n u m b e r of species m o u n t e d , t h e d i s t i n c t i o n s
between spore shapes b r o k e d o w n a n d t e n d e d t o i n t e r g r a d e . M o r e o v e r , w h e n
considered alone, spore shapes t h r e w m a n y u n r e l a t e d species t o g e t h e r a n d
sometimes s e p a r a t e d o b v i o u s l y r e l a t e d species r a t h e r w i d e l y . R e c e n t l y , h o w ever, c e r t a i n f u n d a m e n t a l c h a r a c t e r s a n d r e l a t i o n s h i p s h a v e c o m e t o l i g h t a n d
i t is n o w felt t h a t i n a t least s e v e r a l g r o u p s c e r t a i n f a i r l y c l e a r p r i m a r y d i s t i n c tions m a y n o w be m a d e a t the generic l e v e l o n t h e basis of t h e m o r p h o l o g y of the
h y s t e r o t h e c i a , a s c i , a n d p y c n i d i a , o r a s y n t h e s i s of t h e i r c h a r a c t e r s . S p o r e shape,
s e p t a t i o n , a n d c o l o r m a y t h e n be relegated t o the p o s i t i o n of s e c o n d a r y g e n e r i c
characters.
I n the classification p r o p o s e d b e l o w , the genera Bifusella, Hypodermella,
and
Hypoderma h a v e b e e n c o n s i d e r a b l y r e d u c e d a n d m o s t of t h e i r c o n i f e r - i n h a b i t i n g
species h a v e been r e m o v e d to n e w genera. I t is felt t h a t t h i s r e a l i g n m e n t shows
m o r e a c c u r a t e l y the r e l a t i o n s h i p s of t h e v a r i o u s species a n d genera t o one
a n o t h e r a n d t o t h e o t h e r genera of t h e H y p o d e r m a t a c e a e .
T h e H y p o d e r m a t a c e a e , as t r e a t e d b e l o w , fall i n t o s e v e n g r o u p s o r c o m p l e x e s
of r e l a t e d genera w h i c h m a y be d e s i g n a t e d as f o l l o w s : A, t h e " n e r v i s e q u i a " o r
Lirula c o m p l e x of 3 g e n e r a ; B, t h e Hypodermella
c o m p l e x of 5 g e n e r a ; C, the
Bifusella
c o m p l e x of 3 g e n e r a ; D, the Lophodermium
c o m p l e x of 4 g e n e r a ;
E, the genus Coccomyces; F, t h e Colpoma c o m p l e x of 3 g e n e r a ; a n d G, the
Rhytisma c o m p l e x of 3 genera.
T h e first t h r e e g r o u p s are for the m o s t p a r t s t r o n g l y p a r a s i t i c a n d m a t u r e o n
leaves w h i c h are u s u a l l y in situ o n t w i g s a t the t i m e of spore m a t u r i t y . I n
c o n t r a s t , the Lophodermium
a n d Coccomyces c o m p l e x e s a r e c o m p o s e d l a r g e l y of
w e a k l y p a r a s i t i c species w h i c h i n general d e v e l o p o n i n j u r e d o r w e a k e n e d p l a n t s
m o s t often t o w a r d s t h e e n d of the g r o w i n g season o r t o w a r d s the e n d of the
n o r m a l life s p a n of the l e a v e s . A m o n g t h e species of these g r o u p s are t h e
t y p i c a l needle-casts a n d l e a f - i n h a b i t i n g species i n w h i c h the diseased needles or
leaves u s u a l l y fall t o the g r o u n d a n d after a s o r t of s a p r o p h y t i c existence
t h r o u g h o u t the w i n t e r p r o d u c e t h e i r p y c n i d i a a n d ascocarps d u r i n g t h e
f o l l o w i n g s p r i n g o r s u m m e r . T h e Colpoma c o m p l e x ranges f r o m w e a k l y p a r a s i t i c
species l i k e Colpoma quercinum t o s a p r o p h y t i c species w h i c h d e v e l o p o n d e a d
d e c o r t i c a t e d w o o d . T h e Rhytisma
c o m p l e x is s t r o n g l y p a r a s i t i c , a l t h o u g h the
d a r k s t r o m a t a w h i c h d e v e l o p o n green, d e c i d u o u s leaves d u r i n g t h e g r o w i n g
season of the h o s t p r o d u c e m a t u r e ascocarps o n l y after o v e r w i n t e r i n g o n the
g r o u n d w h i l e o n evergreen leaves t h e y m a y be p r o d u c e d w h i l e s t i l l in situ o n the
branch.
A. T h e " n e r v i s e q u i a " o r Lirula
c o m p l e x (genera 1-3). P e r h a p s the m o s t
p r e s s i n g p r o b l e m i n t h e t a x o n o m y of t h e H y p o d e r m a t a c e a e concerns the
d i s p o s i t i o n of the n e r v i s e q u i o u s species o c c u r r i n g o n Abies a n d Picea. T h e y
f o r m a h o m o g e n e o u s g r o u p of c l o s e l y r e l a t e d species w h i c h are so e a s i l y
r e c o g n i z e d t h a t t h e y c a n be segregated f r o m a l l o t h e r H y p o d e r m a t a c e a e b y the
u n a i d e d eye. Y e t , i n the o l d e r classifications, t h e y are d i s t r i b u t e d i n a n a n o m a lous f a s h i o n a m o n g t h e t h r e e genera Bifusella, Hypoderma,
and
Hypodermella,
a n d a t v a r i o u s t i m e s c e r t a i n species h a v e e v e n been p l a c e d i n t h e genus
Lophodermium.
I n these genera t h e y h a v e c o m e t o rest a m o n g g r o u p s of v e r y
d i v e r s e c o m p o s i t i o n c o m p o s e d of species e x h i b i t i n g a w i d e v a r i e t y o f h y s t e r o t h e c i a l c h a r a c t e r s a n d p a r a s i t i z i n g a b r o a d range of host genera. O n Abies a n d
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C A N A D I A N J O U R N A L O F B O T A N Y . V O L . 45,
1967
Picea the g r o u p is c h a r a c t e r i z e d b y l o n g , b l a c k t o b r o w n h y s t e r o t h e c i a often
e x t e n d i n g the w h o l e l e n g t h of the needles, w h i c h are a t least 2 y e a r s i n age. O n
Abies o n t h e u p p e r surface of 1-year-old needles t h e p y c n i d i a l s t r u c t u r e w h i c h
falls i n the f o r m genus Hypodermina
v . H o h n . extends i n a c o n t i n u o u s o r b r o k e n
l i n e a l o n g the sulcus o r o c c a s i o n a l l y i n t w o lines, one a l o n g e a c h w i n g of t h e
needle. T h e p r i n c i p a l d e v e l o p m e n t of t h i s c o m p l e x is o n Abies, b u t there are
t w o species o n Picea, Lophodermium
macrosporum
( H a r t . ) R e h m a n d Bifusella
crepidiformis
D a r k e r , w h o s e affinities a p p e a r t o be w i t h the n e r v i s e q u i o u s
species o n Abies r a t h e r t h a n w i t h a n y g r o u p of the r e m a i n d e r of the H y p o d e r m a t a c e a e . T h e m o s t l o g i c a l s o l u t i o n is to c o n s i d e r a l l o f these n e r v i s e q u i o u s
species as f o r m i n g a s p e c i a l i z e d c o m p l e x w i t h i n t h e H y p o d e r m a t a c e a e . I n
c o n t r a s t , t h e v a s t m a j o r i t y of the r e m a i n d e r h a v e m o r e i r r e g u l a r l y s c a t t e r e d
h y s t e r o t h e c i a a n d p y c n i d i a . S e c o n d a r i l y one m a y d i v i d e t h i s n e r v i s e q u i o u s
c o m p l e x i n t o three n e w genera o n the basis o f spore shape, as f o l l o w s : (a) a
s h o r t - s p o r e d genus based o n Hypoderma robustum, (b) a b i f u s i f o r m s p o r e d genus
to a c c o m m o d a t e Bifusella abietis, B. faullii,
a n d B. crepidiformis,
a n d (c) a
m u c h larger, f i l i f o r m - c l a v a t e s p o r e d genus b a s e d o n Hypodermella
nervisequia.
B. T h e Hypodermella
c o m p l e x (genera 4-8). T h i s g r o u p of genera is c h a r a c t e r i z e d b y r a t h e r s t r o n g l y p a r a s i t i c species w h i c h f r u i t o n needles in situ o n t h e
tree, s o m e t i m e s o n p a r t i a l l y green needles. T h e y possess for the m o s t p a r t
c o n s p i c u o u s , u s u a l l y e l l i p t i c a l , d a r k or c o n c o l o r o u s f r u i t i n g bodies, w i t h r a t h e r
b r o a d , c l a v a t e - s a c c a t e a s c i a n d r o d - l i k e t o d i s t i n c t l y c l a v a t e or l o n g f u s i f o r m o r
e v e n a l m o s t f i l i f o r m spores. T h e m o s t s t r i k i n g feature o f t h e t y p e species,
Hypodermella
laricis v . T u b . is the s h a p e of i t s ascospores, often referred t o as
i n v e r t e d tear-shaped, t h a t i s , b r o a d l y c l a v a t e a p i c a l l y a n d t a p e r e d t o w a r d s t h e
base. T h e genus s u b s e q u e n t l y c a m e t o be the r e s t i n g place for a l a r g e a n d
heterogeneous g r o u p o f species o n Pinus a n d Abies, w i t h a few o n p h a n e r o g a m i c
hosts. D a r k e r ( 1 9 3 2 , 1 9 3 5 ) , f o l l o w i n g t h i s o l d c o n c e p t o f spore shape, r e c o g n i z e d
19 species o n conifer needles w h i c h f a l l a c c o r d i n g t o his k e y i n t o four w e l l
defined g r o u p s of species. L a g e r b e r g (1949) suggested t h a t the genus s h o u l d be
r e g a r d e d as m o n o s p e c i f i c w i t h t h e n a m e Hypodermella
reserved for t h e t y p e
species o n l y a n d t h a t t h e o t h e r species s h o u l d be relegated t o the genus
Lophodermium.
T h i s , however, i n no w a y appears to solve our d i l e m m a .
P e r h a p s n o one c h a r a c t e r t a k e n a l o n e is sufficiently s i g n i f i c a n t , b u t a s y n thesis of s e v e r a l enables one t o define t h e a b o v e m e n t i o n e d four g r o u p s c l e a r l y .
T h e s e g r o u p s , t h e n , m a y be c i r c u m s c r i b e d as d i s t i n c t genera t y p i f i e d b y the
first species d e s c r i b e d i n e a c h . I n a d d i t i o n t o t h e Hypodermella
nervisequia
g r o u p t r e a t e d u n d e r the " n e r v i s e q u i a " c o m p l e x a b o v e , t h e r e r e m a i n t h r e e o t h e r
generic aggregates of species: Hypodermella v . T u b . , w i t h the type-species o n l y ,
subcuticular, lacking a n opening mechanism, w i t h simple, elliptic, dark colored
p y c n i d i a , o n Larix i n E u r a s i a a n d N o r t h A m e r i c a ; t h e Hypodermella
ampla
g r o u p w i t h s u b e p i d e r m a l , d a r k h y s t e r o t h e c i a , f a i r l y deep-seated, a t m o s t w i t h a
s i m p l e i m m e r s e d core o f p a r e n c h y m a cells f o r e s h a d o w i n g t h e s l i t as i n t h e t y p e
species, p y c n i d i a w h e n present, s i m p l e , o n Pinus, p r i n c i p a l l y i n N o r t h A m e r i c a
b u t w i t h one species f r o m J a p a n ; a n d the Hypodermella
sulcigena g r o u p for
w h i c h v o n H o h n e l e s t a b l i s h e d a n e w genus, Lophodermella,
w i t h deep-seated,
s u b h y p o d e r m a l , m o r e o r less c o n c o l o r o u s h y s t e r o t h e c i a , p y c n i d i a u n r e p o r t e d , o n
Pinus, i n E u r o p e a n d N o r t h A m e r i c a .
D A R K E R : REVISION O F T H E H Y P O D E R M A T A C E A E
1413
I n a d d i t i o n t o these, former Hypodermella
species, a n d w i t h affinities t o t h e
Hypodermella
ampla g r o u p a r e three Hypoderma
species, H. hedgcockii,
H.
lethale, a n d H. pedatum, w i t h s u b h y p o d e r m a l h y s t e r o t h e c i a , a n d s h o r t r o d - l i k e
spores, o n Pinus, i n N o r t h A m e r i c a , w h i c h m a y be d e s i g n a t e d the H. hedgcockii
group- F i n a l l y , Elytroderma
D a r k e r , w i t h t w o - c e l l e d ascospores a n d s u b e p i d e r m a l h y s t e r o t h e c i a o n Pinus i n N o r t h A m e r i c a a n d E u r o p e , c o m p l e t e s t h e
genera of this c o m p l e x .
C. T h e Bifusella c o m p l e x (genera 9 - 1 1 ) . T h e r e m o v a l of t h e " n e r v i s e q u i o u s "
species f r o m Bifusella, as p r o p o s e d a b o v e , leaves a m o n g t h e c o n i f e r - i n h a b i t i n g
species o n l y the type-species, B. linearis ( P k . ) v . H o h n . , a n d a r a t h e r a b e r r a n t
species, B. striiformis
D a r k e r , w h i c h , because of i t s s t r i k i n g d i s s i m i l a r i t i e s i n
characters o t h e r t h a n spore f o r m , is p l a c e d i n a n e w genus. B. linearis, w h i c h
o c c u r s o n soft p i n e s , i s s t r i c t l y s u b c u t i c u l a r a n d is c h a r a c t e r i z e d b y a n e x t e n s i v e ,
effused p y c n i d i a l s t a t e m a d e u p of n u m e r o u s m o r e o r less coalesced p y c n i d i a
a n d a h y s t e r o t h e c i a l s t a t e i n w h i c h t h e p a r a p h y s e s d i s a p p e a r so e a r l y i n t h e
d e v e l o p m e n t t h a t t h e species has c o m e t o be r e g a r d e d as a p a r a p h y s a t e . I t s
closest r e l a t i v e s are t w o species of Hypoderma
w h i c h a l s o o c c u r o n soft p i n e s .
T h e s e species are b o t h s u b c u t i c u l a r , possess large c o n s p i c u o u s p y c n i d i a , s h o w a
t e n d e n c y t o w a r d s t h e b i f u s i f o r m c o n d i t i o n , a n d possess p a r a p h y s e s w h i c h
d i s a p p e a r o r t e n d t o d o so before m a t u r i t y of the a s c o c a r p . I n Hypoderma
pini
( D e a r n . ) D a r k e r , t h e p a r a p h y s e s d i s a p p e a r so e a r l y t h a t i t t o o h a s often been
r e g a r d e d as a p a r a p h y s a t e . I n H. saccatum D a r k e r , t h e p a r a p h y s e s t h o u g h m o r e
persistent f i n a l l y u n d e r g o l y s i s a n d a t m a t u r i t y m a y be l a c k i n g . T h e p y c n i d i a o f
B. linearis, H. pini, a n d H. saccatum possess a definite t h i n p s e u d o p a r e n c h y m a tous c o v e r i n g a b o v e t h e s p o r u l a t i n g c u s h i o n a n d i n t h i s respect r e s e m b l e
s o m e w h a t the p y c n i d i a l s t a t e of Lophodermium
berberidis (Schleich.) R e h m
d e s c r i b e d a n d i l l u s t r a t e d b y N a n n f e l d t (1932). I n Bifusella linearis t h e c o v e r i n g
l a y e r o f t h e p y c n i d i u m a p p e a r s t o c o n s i s t of a s i n g l e l a y e r of d a r k p s e u d o p a r e n c h y m a i m m e d i a t e l y b e l o w a n d a t t a c h e d t o the c u t i c l e . T h e s p e r m a t i a , t o o ,
are v e r y s t r i k i n g , a t t a i n i n g a l e n g t h o f 20 y. o r m o r e . I n t h i s species t h e effused
p y c n i d i a l s t r u c t u r e after d i s c h a r g e of t h e s p e r m a t i a d e v e l o p s i n t o a s t r o m a - l i k e
c r u s t n e a r o r t h r o u g h w h i c h t h e h y s t e r o t h e c i a often e r u p t . T h e fact t h a t
Hemiphacidium
planum
( D a v i s ) K o r f , w h i c h d e v e l o p s s e c o n d a r i l y after B.
linearis, m a y also o c c u r after H. saccatum o n Pinus flexilis suggests a f u r t h e r
r e l a t i o n s h i p b e t w e e n t h e t w o H y p o d e r m a t a c e a e as the s e c o n d a r y f u n g i are i n
general q u i t e specific i n t h e i r h o s t r e l a t i o n s h i p s as w e l l as i n t h e i r a s s o c i a t i o n s
w i t h t h e p r i m a r y parasites. A c c o r d i n g l y , for t h e reasons s t a t e d , these t w o
Hypoderma species h a v e been t r a n s f e r r e d b e l o w t o t h e genus
Bifusella.
T h e o l d e r genus Duplicaria
F c k l . , w h i c h is a l l i e d t o Bifusella v . H o h n . , also
has b i f u s i f o r m spores, b u t a m u c h t h i c k e r c o v e r i n g l a y e r a n d a m o d e r a t e l y w e l l
developed s u b h y m e n i a l layer, a b u n d a n t persistent paraphyses, a n d small,
rather inconspicuous pycnidia.
A n a b e r r a n t species, Bifusella striiformis D a r k e r , w i t h s u b e p i d e r m a l h y s t e r o t h e c i a , s o m e w h a t b i - , o c c a s i o n a l l y t r i f u s i f o r m spores, a n d w i t h p y c n i d i a
u n k n o w n , is p l a c e d i n a n e w genus n e a r Duplicaria
a n d Bifusella i n t h e k e y t o
genera g i v e n b e l o w . I t is n o t p a r t i c u l a r l y c l o s e l y a l l i e d t o e i t h e r o r t o a n y o t h e r
m e m b e r of the H y p o d e r m a t a c e a e b u t is p l a c e d here because of i t s spore shape.
D. T h e Lophodermium
c o m p l e x (genera 1 2 - 1 5 ) . I n c o n t r a s t t o the foregoing
1414
CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
r a t h e r s t r o n g l y p a r a s i t i c g r o u p s is a v e r y d i v e r s e a n d large c o m p l e x of species
w h i c h are p r o b a b l y o n l y w e a k l y p a r a s i t i c . T h e y u s u a l l y c o m p l e t e t h e i r life
cycles m o r e o r less s a p r o p h y t i c a l l y o n fallen conifer needles, o r o n fallen leaves,
o r o n t w i g s o r o n d e a d areas o n evergreen leaves of v a r i o u s p h a n e r o g a m i c
p l a n t s . T h e g r o u p i n c l u d e s t h e s o - c a l l e d " n e e d l e c a s t s " of conifer needles. T h e
ascocarps i n t h e g e n e r a of t h i s g r o u p a r e t y p i c a l l y s h o r t , e l l i p t i c a l , d a r k c o l o r e d ,
a n d r a t h e r s h a r p l y d e l i m i t e d . T h e i r a s c i t e n d to be n a r r o w , v a r y i n g f r o m c y l i n d r i c a l t o m o r e o r less c l a v a t e a n d s o m e t i m e s l o n g - s t a l k e d b u t n e v e r b r o a d l y
s a c c a t e as i n m a n y of t h e foregoing m o r e p a r a s i t i c g r o u p s . T h e i r p a r a p h y s e s are
a b u n d a n t , s i m p l e , s o m e t i m e s c o i l e d o r c l u b - s h a p e d a t the t i p . A s c o s p o r e s v a r y
f r o m s h o r t fusiform o r r o d - s h a p e d t o needle-shaped. P y c n i d i a are s m a l l , c i r c u l a r
to e l l i p t i c a l a n d f a l l i n general i n t o the f o r m genus Leptostroma. F o u r genera are
assigned here.
(J
Hypoderma
D C . ex S a i n t A m a n s e m e n d . D e N o t . , w h i c h has been a " c a t c h a l l " genus for a n y s h o r t - s p o r e d species, s t i l l r e m a i n s one o f the l a r g e r genera
after r e m o v i n g s e v e r a l species elsewhere as p r o p o s e d a b o v e . T h e genus s t i l l
r e m a i n s i n need of s t u d y . I n general i t i n c l u d e s a m i x t u r e of species l a r g e l y
characterized b y clavate asci w h i c h become long-stalked at m a t u r i t y . A s
c o n s t i t u t e d a t present i t i n c l u d e s species whose ascospores v a r y f r o m r o d - l i k e to
fusiform a n d m a y be one-celled, pseudoseptate, o r d e f i n i t e l y u n i s e p t a t e . I t s
ascocarps t e n d t o be r a t h e r t h i n a n d fiat. S u c h a d e f i n i t i o n r e m o v e s from
Hypoderma a l l of t h e c o n i f e r - i n h a b i t i n g species except H. thujae D u r r i e u a n d
H. cunninghamiae
Teng.
A n e w genus is p r o p o s e d b e l o w for Hypoderma
desmazierii
Duby. This
species w i t h i t s u n s t a l k e d , s u b c y l i n d r i c a l a s c i resembles m o r e closely Lophodermium pinastri t h a n t h e o t h e r species of Hypoderma.
Its s t r o n g l y d e v e l o p e d
o p e n i n g m e c h a n i s m a n d t h e m a n n e r i n w h i c h i t s d e v e l o p i n g a s c o c a r p pushes a
g r o u p of e p i d e r m a l h o s t cells t o the b o t t o m o f t h e f r u i t i n g b o d y a r e c e r t a i n l y
r e m i n i s c e n t of L.
pinastri.
Lophodermium
C h e v . , the largest a n d m o s t c o m p l e x genus, t h o u g h a d m i t t e d l y
c o n t a i n i n g a w i d e l y d i v e r g e n t g r o u p of o r g a n i s m s , is the m o s t difficult to
r e o r g a n i z e i n a n y s a t i s f a c t o r y m a n n e r . T h e genus has been t r e a t e d i n a m o n o g r a p h i c s t u d y b y T e h o n (1935) b u t t h e generic l i m i t s a p p l i e d b y v o n H o h n e l
(1917a, d) a n d T e h o n are so p a t e n t l y a r t i f i c i a l t h a t t h e y h a v e r a r e l y been g i v e n
r e c o g n i t i o n b y o t h e r m y c o l o g i s t s . A l t h o u g h c e r t a i n b r o a d d i v i s i o n s m i g h t be
suggested i t seems p r e m a t u r e t o a t t e m p t to m a k e a n y w o r t h w h i l e r e o r g a n i z a tion at this time.
Lophomerum
O u e l l e t t e a n d M a g a s i , h o w e v e r , m a k e s use of p h r a g m o s p o r o u s
s e p t a t i o n as a c h a r a c t e r to segregate c e r t a i n species f r o m the large mass of
Lophodermium
species. C a r e m u s t be exercised i n a p p l y i n g t h i s c r i t e r i o n since
u n d e r e x t r e m e l y m o i s t c o n d i t i o n s m a n y of t h e species o f t h i s genus c a n be
forced t o f o r m a c e n t r a l s e p t u m or e v e n to g e r m i n a t e w i t h i n t h e ascus.
E. T h e genus Coccomyces (genus 16). Coccomyces s t a n d s s o m e w h a t a p a r t
f r o m o t h e r genera o f t h e H y p o d e r m a t a c e a e b u t i t a p p a r e n t l y has r a t h e r definite
affinities to the genus Lophodermium.
I t i n c l u d e s s u b c i r c u l a r to p o l y g o n a l f o r m s
w h i c h tear o p e n l a c e r a t e l y o r o p e n b y s h a r p l y defined r a d i a t e l y a r r a n g e d s l i t s
provided w i t h organized opening mechanisms.
F. T h e Colpoma c o m p l e x (genera 1 7 - 1 9 ) . T h i s g r o u p centers a r o u n d the
D A R K E R : REVISION O F T H E H Y P O D E R M A T A C E A E
1415
Colpoma a n d i n c l u d e s f o r m s w h i c h are e r u m p e n t t h r o u g h the b a r k o r bare
w o o d . T h e y possess a r a t h e r w e l l d e v e l o p e d , s o m e t i m e s m o r e or less d a r k e n e d
basal l a y e r b e n e a t h the s u b h y m e n i a l l a y e r . Colpoma is t h e best k n o w n genus
a n d consists of a n u m b e r of species w i t h scolecosporous ascospores. I n c l u d e d
here at present are a n u m b e r of species, s e v e r a l o f w h i c h s h o u l d p r o b a b l y be
reduced t o s y n o n y m y . Xyloschizon
S y d . , a s o m e w h a t d o u b t f u l genus o n t h e
basis of the d e s c r i p t i o n of its t y p e species, X. weirianum S y d . , a p p e a r s to be a
p h r a g m o s p o r o u s segregate f r o m Colpoma. T h e s e c o n d species, X. stratum S y d . ,
o n Crataegus, does n o t a p p e a r t o differ m a r k e d l y f r o m the w i d e s p r e a d Colpoma
crispum ( P . ex F r . ) S a c c , c o m m o n o n b a r k a n d b a r e w o o d of conifers. T h e t h i r d
genus of t h i s g r o u p is Bifusepta
D a r k e r , a p h r a g m o s p o r o u s genus w i t h b i f u s i form spores.
G. T h e Rhytisma
c o m p l e x (genera 2 0 - 2 2 ) . F i n a l l y , t h e s e v e n t h g r o u p i n g
includes the genera Rhytisma, Placuntium,
a n d Nymanomyces
a n d c o m p r i s e s the
large s t r o m a t i c t y p e s w h i c h m a y h a v e s e v e r a l ascocarps i n each s t r o m a as i n
most Rhytisma species a n d i n Nymanomyces.
O f t e n i n Placuntium
andromedae
(P.) E h r e n b g . ex v . H o h n . , the n u m b e r of ascocarps i n a s t r o m a is r e d u c e d to
one large c i r c u l a r a s c o c a r p s u r r o u n d i n g a sterile a r e a i n e a c h s t r o m a . T h e
characters of Placuntium
are those of a r a t h e r r o b u s t Rhytisma
w i t h large,
s l i g h t l y g r e e n i s h - y e l l o w c l a v a t e ascospores i n c o n t r a s t t o t h e m o r e h y a l i n e ,
s m a l l e r c l a v u l a t e ascospores of Rhytisma.
I n t h e genus Nymanomyces
the
ascospores are b r o w n a n d o v o i d .
s
T h e present s t u d y , t h e n , d i v i d e s t h e H y p o d e r m a t a c e a e a m o n g 22 genera,
i n c l u d i n g 15 h i t h e r t o v a l i d l y p u b l i s h e d genera c o m p o s e d o f t h e 11 genera
recognized b y v o n A r x a n d M u l l e r (1954) t o g e t h e r -with Lophodermella v . H o h n . ,
the s o m e w h a t d o u b t f u l Xyloschizon
S y d . , Bifusepta
Darker, and
Lophomerum
O u e l l e t t e a n d M a g a s i . I n a d d i t i o n s e v e n n e w genera are p r o p o s e d b e l o w t o g i v e
a more m e a n i n g f u l c l a s s i f i c a t i o n , e s p e c i a l l y for the c o n i f e r - i n h a b i t i n g species.
Concerning the Names of the Genera Hypoderma and Lophodermium
T h e n o m e n c l a t u r e o f the genus Lophodermium
is b o u n d i n t i m a t e l y w i t h t h a t
of Hypoderma,
a genus e s t a b l i s h e d b y D e C a n d o l l e ( i n L a m a r c k a n d D e
C a n d o l l e , 1805) for 5 species segregated f r o m Hysterium P e r s . a n d Xyloma P e r s .
a n d t o w h i c h D e C a n d o l l e (1815) l a t e r a d d e d 7 m o r e species a n d a n u m b e r of
v a r i e t i e s . S a i n t A m a n s (1821) e s t a b l i s h e d p r i o r i t y for Hypoderma
and placed
t h e r e i n 5 o f D e C a n d o l l e ' s species i n c l u d i n g H. arundinaceum
(Schrad.) D C .
a n d H. virgultorum D C , a n a v o w e d s u b s t i t u t e for Hysterium rubi P e r s . , b o t h of
w h i c h were d e s t i n e d t o p l a y l e a d i n g p a r t s i n the c o m p l i c a t e d i n t e r c h a n g e s
b e t w e e n Hypoderma
a n d Lophodermium.
C h e v a l l i e r (1822) segregated f r o m
Hypoderma
o n r a t h e r q u e s t i o n a b l e g r o u n d s a n e w genus Lophoderma
and
i n c l u d e d H. arundinaceum
i n the f o r m e r a n d Lophoderma rubi (Pers.) ex C h e v .
i n the l a t t e r . L a t e r , C h e v a l l i e r (1826) a b a n d o n e d b o t h g e n e r a a n d u n i t e d t h e i r
species i n a n e w g6nus Lophodermium.
C l e a r - c u t c i r c u m s c r i p t i o n s were n o t
d r a w n u p for Hypoderma
D C . ex S a i n t A m a n s a n d Lophodermium
Chev. until
D e N o t a r i s (1847) r e v i s e d b o t h genera a n d i n c l u d e d Hypoderma
rubi (Pers. ex
C h e v . ) D e N o t . i n the f o r m e r a n d Lophodermium
arundinaceum
( S c h r a d . ex
S a i n t A m a n s ) C h e v . i n the l a t t e r . U n f o r t u n a t e l y , t h i s t r e a t m e n t r e v e r s e d the
p o s i t i o n t a k e n b y C h e v a l l i e r i n 1822. O t t o K u n t z e (1898) a r r i v e d a t e s s e n t i a l l y
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CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
t h e s a m e c o n c l u s i o n s as C h e v a l l i e r (1822) b y a r g u i n g t h a t , s i n c e 3 of t h e 5
species o r i g i n a l l y n a m e d b y D e C a n d o l l e i n 1805 were c o n g e n e r i c , the m a j o r
g r o u p of species w h i c h i n c l u d e d H. arundinaceum,
s h o u l d h a v e been assigned
l a t e r to Hypoderma
i n s t e a d of Lophodermium.
A c c o r d i n g l y , he transferred a l l
Lophodermium
species l i s t e d i n t h e first 11 v o l u m e s of S a c c a r d o ' s S y l l o g e
F u n g o r u m b a c k t o Hypoderma
a n d a l l of the Hypoderma
species i n the s a m e
v o l u m e s t o a n e w genus
Hypodermopsis.
C h e v a l l i e r (1822) i n d i v i d i n g the genus Hypoderma a n d p l a c i n g H.
arundinaceum i n t h e Hypoderma e n t i t y t h e r e b y a u t o m a t i c a l l y fixed t h i s species as t y p e .
'Lophodermium
C h e v . (1826) i n i t s o r i g i n a l c i r c u m s c r i p t i o n a n d i n the r e v i s e d ,
r e s t r i c t e d c i r c u m s c r i p t i o n of D e N o t a r i s (1847) is superfluous a n d therefore
i l l e g i t i m a t e ( A r t . 63) because i t i n c l u d e d L. arundinaceum,
the type-species of
Hypoderma.
Hypoderma
as t r e a t e d b y O . K u n t z e (1898) i n c l u d e d H.
arundinaceum a n d is i n a c c o r d w i t h C h e v a l l i e r ' s t r e a t m e n t of 1822.
I n the Lophoderma
e n t i t y C h e v a l l i e r (1822) p l a c e d Lophoderma
rubi, L.
scirpinum,
a n d X . nervisequum, b u t l a t e r he (1826) a b a n d o n e d Lophoderma
and
t r a n s f e r r e d i t s species t o g e t h e r w i t h a n u m b e r of Hypoderma species i n c l u d i n g
H. arundinaceum
t o his n e w genus Lophodermium,
a n a v o w e d s u b s t i t u t e for
Hypoderma.
D e N o t a r i s (1847) r e v i s e d Hypoderma
w i t h the type-species, H.
arundinaceum,
r e m o v e d . W i t h t h i s r e v i s i o n the n a m e w o u l d h a v e t o be a s c r i b e d
t o D e N o t a r i s a n d w o u l d be a l a t e r h o m o n y m of Hypoderma
D C . ex S a i n t
A m a n s a n d Hypoderma as r e v i s e d b y C h e v a l l i e r , a n d a s y n o n y m of Lophoderma
C h e v . Hypodermopsis
0 . K u n t z e is superfluous because i t i n c l u d e d i n i t s
c i r c u m s c r i p t i o n a l l three species of Lophoderma C h e v .
T h e c u r r e n t usage of Hypoderma a n d Lophodermium
follows the t r e a t m e n t of
D e N o t a r i s (1847) a n d is illegal for b o t h genera b u t the r e v i s i o n of D e N o t a r i s
has since his t i m e been f o l l o w e d b y p r a c t i c a l l y a l l m y c o l o g i s t s w i t h the except i o n of O . K u n t z e (1898) a n d t o r e p o r t those w h o h a v e a d o p t e d t h i s usage
results i n a l i s t of a l l persons k n o w l e d g e a b l e i n the H y p o d e r m a t a c e a e i n c l u d i n g
D u b y (1861), F u c k e l (1870), K a r s t e n (1873), S a c c a r d o (1883), R e h m (1887),
E l l i s a n d E v e r h a r t (1892), S c h r o e t e r (1893), a n d a m o n g m o r e recent w o r k e r s ,
v o n H o h n e l (1917a), H i l i t z e r (1929), N a n n f e l d t (1932), T e h o n (1935), T e r r i e r
(1942), a n d others.
I n k e e p i n g w i t h the s p i r i t of A r t i c l e 14 of the I n t e r n a t i o n a l C o d e of B o t a n i c a l
N o m e n c l a t u r e ( L a n j o u w et al. 1961) i t is h i g h l y d e s i r a b l e t h a t these t w o genera
be m a i n t a i n e d w i t h the c i r c u m s c r i p t i o n s of D e N o t a r i s t o a v o i d the n u m e r o u s
c o n f u s i n g changes t h a t a s t r i c t a p p l i c a t i o n of the rules w o u l d e n t a i l .
T h e s i m p l e s t m e t h o d of l e g a l i z i n g t h e present usage requires first, as perm i t t e d b y A r t i c l e 7, N o t e 8, of t h e " C o d e , " the s e l e c t i o n a n d c o n s e r v a t i o n of a
n e w type-species for Hypoderma
D C . ex S a i n t A m a n s , a v a l i d n a m e a t present
l e g a l l y t y p i f i e d b y H. arundinaceum.
If, h o w e v e r , H. virgultorum D C . ex S a i n t
A m a n s , a n o b l i g a t e s y n o n y m of H. rubi ( P e r s . ex C h e v . ) D e N o t . , were c o n s e r v e d as t h e type-species of Hypoderma,
the s t a t u s o f t h e genus w o u l d t h e n be
fixed w i t h a c i r c u m s c r i p t i o n i n k e e p i n g w i t h t h a t o f D e N o t a r i s . Lophoderma
C h e v . ( i n c l u d i n g L. rubi) w o u l d t h e n b e c o m e superfluous ( A r t . 63). Hypoderma
i n t h e sense of C h e v a l l i e r w i t h the type-species (H. rubi) r e m o v e d w o u l d h a v e
t o be a s c r i b e d t o C h e v a l l i e r ( A r t . 48) a n d w o u l d b e c o m e a l a t e r h o m o n y m .
Lophodermium
C h e v . , because i t i n c l u d e s H. rubi, w o u l d s t i l l r e m a i n superfluous
D A R K E R : REVISION OF T H E
HYPODERMATACEAE
1417
d Lophodermium
i n t h e sense of D e N o t a r i s w i t h the t y p e of
Lophodermium
C h e v r e m o v e d w o u l d h a v e t o be a s c r i b e d t o D e N o t a r i s a n d w o u l d b e c o m e a
later h o m o n y m o f Lophodermium
C h e v . Hypoderma
D C . e x S a i n t A m a n s , as
treated b y D e N o t a r i s , w o u l d t h e n b e c o m e l e g a l a n d i t s usage w o u l d c o r r e s p o n d t o t h a t of S a i n t A m a n s a n d w o u l d agree w i t h c u r r e n t usage. Hypoderma i n
the sense of 0 . K u n t z e , h o w e v e r , w o u l d b e c o m e s y n o n y m o u s w i t h
Hypoderma
C h e v . , w h i l e Hypodermopsis
O . K u n t z e w o u l d b e c o m e i l l e g i t i m a t e as superfluous
for Hypoderma
D C . ex S a i n t A m a n s . U n f o r t u n a t e l y , as p o i n t e d o u t a b o v e ,
Lophodermium
w o u l d s t i l l r e m a i n superfluous. T o l e g a l i z e t h i s n a m e i t is
therefore necessary t o f a l l b a c k o n A r t i c l e 14 of the " C o d e " a n d c o n s e r v e
Lophodermium
C h e v . w i t h the type-species L. arundinaceum
( S c h r a d . ex S a i n t
Amans) Chev.
T h e f o r m a l p r o p o s a l s for c o n s e r v a t i o n are m a d e b e l o w for s u b m i s s i o n t o the
G e n e r a l C o m m i t t e e for B o t a n i c a l N o m e n c l a t u r e . P e n d i n g the r e c o m m e n d a tions of the C o m m i t t e e o n the p r o p o s a l s , the e x i s t i n g usage of Hypoderma
and
Lophodermium
is b e i n g f o l l o w e d .
PROPOSAL 1
P r o p o s a l for the c o n s e r v a t i o n of a type-species for Hypoderma
A m a n s (1821) ( H y p o d e r m a t a c e a e ) .
D C . ex S a i n t
L E C T O T Y P U S : Hypoderma
rubi (Pers. ex C h e v . ) D e N o t . G i o r n . B o t . I t a l .
2(2): (37). 1847. (typ. cons. prop.).
[ s s Hysterium
rubi P e r s . , O b s . M y c . 1: 84. 1 7 9 6 ; = Hypoderma
virgultorum D C , F l . F r a n c . 3 ed. 6: 165. 1815 (an a v o w e d s u b s t i t u t e ) ; = H.
virgultorum
D C . ex S a i n t A m a n s , F l . A g e n . 515. 1 8 2 1 ; =s
Lophoderma
rubi (Pers.) ex C h e v . , J . P h y s . 94: 3 1 . 1 8 2 2 ; = Hysterium
rubi P e r s . ex
F r . , S y s t . M y c . 2: 5 8 7 . 1 8 2 3 ] .
V E R S U S L E C T O T Y P U S : Hypoderma
A m a n s , F l . A g e n . 5 1 6 . 1 8 2 1 . (typ. rej.
arundinaceum
prop.).
(Schrad.)
D C . ex S a i n t
DISCUSSION
W h e n C h e v a l l i e r (1822) d i v i d e d Hypoderma
D C . ex S a i n t A m a n s i n t o t w o
g r o u p s , he r e t a i n e d H. arundinaceum
i n Hypoderma,
i n effect s e l e c t i n g a
l e c t o t y p e for the n a m e , a n d erected Lophoderma to i n c l u d e L. rubi ( a n o b l i g a t e
s y n o n y m of H. virgultorum).
D e N o t a r i s (1847) r e v i s e d t h e genus
Hypoderma
w i t h the e x c l u s i o n of H. arundinaceum,
b u t the i n c l u s i o n of H. rubi. T h i s l a t e r
usage h a s been f o l l o w e d i n a l l m a j o r studies w i t h t h e e x c e p t i o n of t h a t of
0 . K u n t z e (1898). T h e p r o p o s a l seeks to legalize the n o m e n c l a t u r e of m o r e t h a n
140 years of reference to the genus Hypoderma a n d i n c i d e n t a l l y to e l i m i n a t e a n y
c l a i m the r a r e l y a n d a l w a y s i n c o r r e c t l y used Lophoderma m i g h t h a v e to species
currently placed i n
Hypoderma.
PROPOSAL 2
P r o p o s a l for the c o n s e r v a t i o n of the generic n a m e Lophodermium
Chevallier
(1826) ( H y p o d e r m a t a c e a e ) .
Lophodermium
C h e v a l l i e r , F l . P a r i s 1: 4 3 5 . 1 8 2 6 . (nom. cons. prop.).
L E C T O T Y P U S : L. arundinaceum
prop.).
( S c h r a d . ex S a i n t A m a n s ) C h e v . (typ.
cons,
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CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
DISCUSSION
W h e t h e r Hypoderma
D C . ex S a i n t A m a n s r e m a i n s t y p i f i e d b y H.
arundinaceum i n a c c o r d a n c e w i t h the c h o i c e of C h e v a l l i e r (1822) o r , t h r o u g h the
a c c e p t a n c e of p r o p o s a l 1, becomes t y p i f i e d b y H. rubi ( u n d e r the n a m e of H.
virgultorum),
the n a m e Lophodermium
C h e v . is superfluous a n d therefore
i l l e g i t i m a t e , because i t s o r i g i n a l c i r c u m s c r i p t i o n i n c l u d e d b o t h species. P r o p o s a l
2 serves a t w o - f o l d p u r p o s e : t o conserve the n a m e of a genus t h a t has been i n
use for 140 years, a n d , to c o n s e r v e i t i n the sense of D e N o t a r i s . T h u s , the
r e v i s i o n b y D e N o t a r i s of b o t h Hypoderma a n d Lophodermium,
w h i c h is w h a t
e v e r y one is f o l l o w i n g , c a n be m a d e l e g a l .
REASONS FOR CONSERVATION
A c c o r d i n g to a s t r i c t a p p l i c a t i o n of the rules of n o m e n c l a t u r e , w i t h o u t the
a c c e p t a n c e of the a b o v e - m e n t i o n e d p r o p o s a l s , a l l Lophodermium
species w o u l d
h a v e to be p l a c e d i n the genus Hypoderma
a n d a l l Hypoderma
species i n the
genus Lophoderma. T h i s is e s s e n t i a l l y w h a t w a s p r o p o s e d b y O . K u n t z e (1898)
e x c e p t t h a t he erected Hypodermopsis
O. K u n t z e , an inadvertent but avowed
s u b s t i t u t e for Lophoderma. M o r e t h a n 100 species of Lophodermium
and about
50 species of Hypoderma,
m o r e t h a n one-half of the k n o w n species of the
H y p o d e r m a t a c e a e , w o u l d be i n v o l v e d i n changes if the t w o genera,
Hypoderma
a n d Lophodermium,
were n o t r e t a i n e d w i t h the c i r c u m s c r i p t i o n s of D e N o t a r i s .
W i t h r e t e n t i o n i n t h i s sense there w o u l d be n o a m b i g u i t y a t t a c h e d to e i t h e r
n a m e . W h i l e Hypoderma
is v a l i d l y p u b l i s h e d a n d l e g i t i m a t e there is n o l e g i t i m a t e n a m e for Lophodermium
i n the sense of D e N o t a r i s . A c c e p t a n c e of the
proposals w o u l d n o t i n v o l v e the c r e a t i o n of n e w c o m b i n a t i o n s or a u t h o r
c i t a t i o n s b u t w o u l d m e r e l y legalize the genus Lophodermium
a n d the usage o f
c o m b i n a t i o n s u n d e r b o t h genera i n the sense of D e N o t a r i s a n d of c u r r e n t
practice.
K E Y TO G E N E R A OF H Y P O D E R M A T A C E A E
A . S t r o m a t a simple, mono-ascocarpous
B.
B . B a s a l stromatic tissue w e a k l y developed
C.
C . Ascocarps elliptical to linear
D.
D . Ascocarps m o s t l y linear, often nervisequious, d a r k b r o w n
to b l a c k ; p y c n i d i a continuous or in b r o k e n lines
E.
E . Ascospores rod-shaped
I. V I R G E L L A
E . Ascospores bifusiform
II. I S T H M I E L L A
E . Ascospores clavate t o c l a v a t e - c y l i n d r i c a l
III. L I R U L A
D . Ascocarps m o s t l y e l l i p t i c a l , concolorous to b l a c k ; p y c n i d i a scattered
or u n k n o w n
F.
F . A s c i more or less b r o a d l y saccate to clavate
G.
G . Ascospores not bifusiform
H.
H . Ascocarps subcuticular, ascospores c l a v a t e
IV. H Y P O D E R M E L L A
H . Ascocarps subepidermal, ascospores variable
I.
I. Ascospores rod-shaped, continuous
V. PLOIODERMA
I. Ascospores clavate, continuous
VI. DAVISOMYCELLA
I. Ascospores elongated, fusiform, 2-celled
VII. E L Y T R O D E R M A
H . Ascocarps subhypodermal, ascospores c l a v a t e . . . V I I I . L O P H O D E R M E L L A
G . Ascospores bifusiform
. . J.
j . Ascocarps subcuticular
K.
K . P y c n i d i a large, often effuse, conspicuous
IX. BIFUSELLA
K . P y c n i d i a s m a l l , discrete, inconspicuous
X. DUPLICARIA
J . Ascocarps subepidermal
XI. SOLEELLA
F . A s c i n a r r o w l y clavate or c y l i n d r i c a l
L.
L . A s c i clavate a n d t y p i c a l l y long-stalked, ascospores
rod-shaped
XII. HYPODERMA
L . A s c i more or less c y l i n d r i c a l
M.
D A R K E R : REVISION O F T H E H Y P O D E R M A T A C E A E
1419
M . Ascospores rod-shaped
XIII. M E L O D E R M A
M . Ascospores
filiform
N.
N . Ascospores continuous
XIV. LOPHODERMIUM
N . Ascospores phragmosporous
XV. LOPHOMERUM
C. Ascocarps subcircular to polyhedral
XVI. COCCOMYCES
B B a s a l stromatic tissue strongly developed
O.
O. Ascospores clavate to filiform, continuous
XVII. COLPOMA
o! Ascospores clavate, phragmosporous
XVIII. XYLOSCHIZON
o! Ascospores bifusiform, phragmosporous
XIX. BIFUSEPTA
A . S t r o m a t a complex, p o l y - a n d usually syncarpous
P.
P. Ascospores hyaline, clavate to
filiform
Q.
Q. Ascocarps numerous i n s t r o m a ; ascospores filiform to clavulate
X X . RHYTISMA
O Ascocarps few i n s t r o m a ; ascospores b r o a d l y clavate
XXI. PLACUNTIUM
P . Ascospores b r o w n , o v o i d
XXII. NYMANOMYCES
Genera of Hypodermataceae
Virgella gen. n o v . H y p o d e r m a t a c e a r u m
Hysterothecia h y p o p h y l l a , atrofusca, innata, nervisequia, longitudinali
i n c i s u r a a p e r i e n t i a , p r i m o r d i o i n c i s u r a e c o n s p i c u o ; s t r a t u m tegens a t r i pseudoparenchymatis; s t r a t u m basilare p l e c t e n c h y m a t i c u m a c h r o u m . A s c i saccatoc l a v a t i . P a r a p h y s e s s i m p l i c e s , filiformes. A s c o s p o r a e b a c i l l a r e s , m u c o i n v o l u t a e .
Pycnidia simplicia continua epiphylla; stratum conidiophororum applanatum;
spermatia minuta, bacillaria.
I.
E T Y M O L O G I A : virga, a s t r e a k o r s t r i p e of c o l o r o n clothes, et -ella, a d i m i n u t i v e
suffix; i n reference to the a p p e a r a n c e of the h y s t e r o t h e c i u m .
SPECIES TYPICA:
Virgella
robusta
(v. T u b . )
Darker
Hysterothecia hypophyllous, dark brown, innate, nervisequious, opening b y
a l o n g i t u d i n a l fissure; p r i m o r d i u m of s l i t c o n s p i c u o u s ; c o v e r i n g l a y e r of d a r k
p s e u d o p a r e n c h y m a ; b a s a l l a y e r p l e c t e n c h y m a t o u s , colorless. A s c i saccatec l a v a t e . P a r a p h y s e s s i m p l e , filiform. A s c o s p o r e s r o d - s h a p e d , enclosed i n
gelatinous sheath. P y c n i d i a simple, continuous, e p i p h y l l o u s ; conidiophore layer
flat; s p e r m a t i a m i n u t e , b a c i l l a r .
I. Virgella robusta (v. T u b . ) c o m b . nov.
= Lophodermium
infectans M a y r , D i e W a l d u n g e n v o n N o r d a m e r i k a .
336.1890 (nom. nud.).
= Hypoderma robustum v . T u b . , A r b . B i o l . A b t h . L a n d - u . F o r s t w . k a i s .
G e s u n d h . 2(1): 16.1901.
S P E C I M E N I S O T Y P I C U M : o n Abies concolor ( G o r d . ) P a r r y , S a n B e r n a r d i n o
M o u n t a i n s , C a l i f o r n i a , H. Mayr, i n F a r l o w H e r b a r i u m .
I n c o n v e r s a t i o n Professor C . v o n T u b e u f i n f o r m e d the w r i t e r a n u m b e r of
years ago t h a t M a y r ' s m a t e r i a l , t h r o u g h serious neglect, h a d b e c o m e w o r t h l e s s
a n d h a d been d i s c a r d e d . T h e i s o t y p e m a t e r i a l sent t o Professor W . G . F a r l o w
b y M a y r is a p a r t of the o r i g i n a l c o l l e c t i o n a n d was a p p a r e n t l y the basis also for
v o n T u b e u f ' s l a t e r d e s c r i p t i o n (see B o y c e , M y c o l o g i a 19: 284, 2 8 5 . 1 9 2 7 ) .
II.
Isthmiella gen. n o v . H y p o d e r m a t a c e a r u m
G e n u s Virgettae s u b s i m i l e , sed ascosporis b i f u s i f o r m i b u s .
E T Y M O L O G I A : isthmus, i s t h m u s , et -ella, a d i m i n u t i v e suffix, i n reference to the
spore shape.
S P E C I E S T Y P I C A : Isthmiella abietis ( D e a r n . ) D a r k e r
A genus s o m e w h a t s i m i l a r to Virgella, b u t w i t h b i f u s i f o r m ascospores.
1420
1.
C A N A D I A N J O U R N A L O F B O T A N Y . V O L . 45, 1967
Isthmiella abietis
B== Bifusella
(Dearn.) comb. nov.
abietis D e a r n . , M y c o l o g i a 18: 2 3 9 . 1 9 2 6 .
S P E C I M E N H O L O T Y P I C U M : i n D e a r n e s s H e r b a r i u m , o n Abies
( H o o k . ) N u t t . , C h a l l i s N a t . F o r . , B o n a n z a , I d a . , A u g . 7, 1911. G. G.
9395.
lasiocarpa
Hedgcock:
l^Isthmiella crepidiformis
^•Bifusella
3.
crepidiformis
(Darker) comb. nov.
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 2 2 . 1 9 3 2 .
Isthmiella faullii ( D a r k e r )
Bifusella
faullii
comb. nov.
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 1 9 . 1 9 3 2 .
I I I . L i r u l a gen. n o v . H y p o d e r m a t a c e a r u m
G e n u s Virgellae s u b s i m i l e , sed ascosporis c l a v a t o - f i l i f o r m i b u s .
E T Y M O L O G I A : lira, a f u r r o w , e t -ula, a d i m i n u t i v e suffix; i n reference to t h e
a p p e a r a n c e of the h y s t e r o t h e c i u m , e s p e c i a l l y w h e n o p e n .
S P E C I E S T Y P I C A : L. nervisequia
( D C . ex F r . )
A genus s o m e w h a t s i m i l a r to Virgella,
1.
Darker
b u t w i t h c l a v a t e - f i l i f o r m ascospores.
Lirula nervisequia
( D C . ex F r . ) c o m b . n o v .
^Hypoderma
nervisequum D C , F l . F r a n c . 6: 1 6 7 . 1 8 1 5 .
^Lophoderma
nervisequum ( D C . ) ex C h e v . , J . P h y s . 94: 3 1 . 1822.
= Hysterium nervisequium
( D C ) ex F r . , S y s t . M y c . 2 : 5 8 7 . 1 8 2 3 .
s= Lophodermium
nervisequium
( D C . ex F r . ) C h e v . , F l . P a r i s 1: 435.
1826.
^Lophodermium
nervisequium
( D C . ex F r . ) R e h m , i n R a b h . , K r y p t . - F l .
I I . 1(3): 4 4 . 1 8 8 7 .
Hypodermopsis
nervisequa ( D C ex F r . ) O . K u n t z e , R e v . G e n . P I . 3 ( 2 ) :
487. 1898.
a s Hypodermella
nervisequia
( D C . ex F r . ) L a g e r b e r g , M e d d . S t a t .
SkogsforsQksanst. 7: 1 4 8 . 1 9 1 0 .
= Hypodermella
lirelliformis
D a r k e r , C o n t r i b . A r n o l d A r b o r . 1: 4 5 . 1 9 3 2 .
T h e i s o t y p e of the F r i e s i a n h o l o t y p e is p r e s e n t i n M o u g e o t ' s a m p l e c o l l e c t i o n
w h i c h is p r e s e r v e d i n t h e h e r b a r i u m o f t h e C o n s e r v a t o i r e et J a r d i n B o t a n i q u e s
( G - D C ) , G e n e v a , S w i t z e r l a n d . T h i s s p e c i m e n has o n l y the s m a l l , i n c o n s p i c u o u s , s c a t t e r e d p y c n i d i a a n d n o t the b r o a d , c o n s p i c u o u s k i n d i l l u s t r a t e d b y
H a r t i g , v o n T u b e u f , a n d others. N o t h a v i n g seen the t y p e of
Hysterium
nervisequium
a t the t i m e of p u b l i c a t i o n , D a r k e r (1932) a s s u m e d the H a r t i g
i l l u s t r a t i o n s to represent the p y c n i d i a l s t a t e of H. nervisequium.
Unfortunately
t h i s w a s n o t s o a n d c o n s e q u e n t l y H. lirelliformis
D a r k e r must become a
s y n o n y m of L. nervisequia. T h e species w r o n g l y d e t e r m i n e d a n d d e s c r i b e d as
H. nervisequia b y D a r k e r is n o w p r o p o s e d as a n e w v a r i e t y .
2. Lirula
nervisequia
( D C ex F r . ) D a r k e r v a r .
conspicua v a r .
nov.
V a r i e t a s Lirellae
nervisequiae
s i m i l i s , sed p y c n i d i i s c o n s p i c u i s , e p i p h y l l i s ,
c o n t i n u i s s e c u n d u m s i n u m acus, subeffusis, 2 8 0 - 6 3 0 n l a t i s , a t r o b r u n n e i s p o s t
dissipationem spermatiorum.
E T Y M O L O G I A : conspicua,
conspicuous.
DARKER: REVISION OF T H E HYPODERMATACEAE
1421
S P E C I M E N T Y P I C U M : D . S a c c a r d o , M y c o t h . I t a l . N o . 504. 1900. "Lophodermium nervisequum ( D C . ) R e h m — i n foliis Abietis pectinatae. M a i o 1898. A b . A .
Carestia."
V a r i e t y s i m i l a r to Lirula nervisequia b u t w i t h c o n s p i c u o u s p y c n i d i a , e p i p h y l lous, c o n t i n u o u s a l o n g the sinus of the needle, s o m e w h a t effuse, 2 8 0 - 6 3 0 /x w i d e ,
d a r k b r o w n after the d i s p e r s a l of the s p e r m a t i a .
S T A T U S P Y C N I D I C U S : Hypodermina
nervisequa ( L k . ) v . H o h n . , S i t z b . k . A k a d .
Wiss. W i e n , Math.-nat. Klasse. A b t . 1.125(1/2): 55.1916.
= Hypodermium
nervisequum L k . , S p . P I . 6 ( 2 ) : 8 9 . 1 8 2 4 .
=sSchizoderma nervisequum ( L k . ) D u b y , B o t . G a l l . 2 : 8 8 5 . 1 8 3 0 .
— Daedala nervicola H a z s l i n s z k y , V e r h . Z o o l . - b o t . G e s . W i e n 37: 1 5 4 . 1 8 8 7 .
F r o m v o n H o h n e l ' s d e s c r i p t i o n of the p y c n i d i a l s t a t e i t a p p e a r s t h a t h e w a s
d e s c r i b i n g the p y c n i d i a of the v a r i e t y conspicua w h e n h e m a d e the n e w c o m b i n a t i o n Hypodermina
nervisequa ( L k . ) v . H 6 h n . a n d i t seems q u i t e l i k e l y t h a t
b o t h L i n k a n d D u b y were d e s c r i b i n g t h e s a m e fungus. F o r v a r i o u s reasons v o n
H o h n e l rejected b o t h Hypodermium
L k . a n d Schizoderma D u b y b u t o v e r l o o k e d
the genus Daedala H a z s l . w i t h the species D. nervicola H a z s l . F r o m a n e x a m i n a t i o n of the t y p e s p e c i m e n , t h i s , too, a p p e a r s t o be the i m p e r f e c t s t a t e of t h e v a r .
conspicua.
If so, the genus has p r i o r i t y o v e r Hypodermina
v. Hohn., but
u n f o r t u n a t e l y the genus n a m e is so s i m i l a r to Daedalea P . ex F r . , w i t h w h i c h i t
has been confused b y the o n l y w r i t e r s w h o h a v e c i t e d i t ( H i l i t z e r , 1929, D a r k e r ,
1932, a n d T e h o n , 1935), t h a t i t seems b e s t t o i n v o k e A r t i c l e 75 of the I n t e r n a t i o n a l C o d e of B o t a n i c a l N o m e n c l a t u r e a n d a b a n d o n i t . I n t h i s e v e n t i t m a y
be t r e a t e d as a n o r t h o g r a p h i c v a r i a n t of Daedalea P e r s . ex F r . of w h i c h i t
becomes a later h o m o n y m .
T h e p y c n i d i a l s t a t e of L. nervisequia v a r . conspicua, u n t i l p r o o f is s h o w n t o
the c o n t r a r y , s h o u l d be c a l l e d Hypodermina
nervisequa ( L k . ) v . H o h n . O n the
other h a n d , the p y c n i d i a l s t a t e o f L. nervisequia does n o t a p p e a r to h a v e b e e n
n a m e d , a l t h o u g h i t w a s briefly d e s c r i b e d b y D a r k e r (1932) as the p y c n i d i a l
state of Hypodermella
lirelliformis.
3.
Lirula abietis-concoloris ( M a y r
ex D e a r n . ) c o m b . n o v .
= Lophodermium
abietis-concoloris
Mayr, Die Waldungen von Nordamerika, 336.1890 (nom. nud.).
s= Hypodermella
abietis-concoloris
( M a y r ) ex D e a r n . , M y c o l o g i a 16: 150.
1924.
S P E C I M E N N E O T Y P I C U M : i n D e a r n e s s H e r b a r i u m , o n Abies concolor
P a r r y , n e a r W h i t n e y , B a k e r C o . , O r e . O c t . 2 8 , 1 9 2 0 . J. S. Boyce: 732.
(Gord.)
V o n T u b e u f (1901) s t a t e d t h a t M a y r ' s m a t e r i a l w a s i m m a t u r e a n d i n c o n v e r s a t i o n i n 1932 t o l d the w r i t e r t h a t t h r o u g h l a c k of care M a y r ' s c o l l e c t i o n s
h a d b e c o m e worthless a n d h a d been d i s c a r d e d . D e a r n e s s r e v i v e d t h e specific
e p i t h e t for s e v e r a l s p e c i m e n s w h i c h he c o n s i d e r e d t o fit M a y r ' s d e s c r i p t i o n .
A l t h o u g h d o u b t w i l l a l w a y s r e m a i n c o n c e r n i n g M a y r ' s c o l l e c t i o n , the n e o t y p e
s p e c i m e n of D e a r n e s s is g o o d a n d defines a d i s t i n c t i v e species. I t represents,
h o w e v e r , b u t one of s e v e r a l closely related species o r v a r i e t i e s of a n aggregate
w h i c h extends t h r o u g h o u t the m o u n t a i n s of w e s t e r n N o r t h A m e r i c a .
CANADIAN JOURNAL OF BOTANY. VOL. 45,
1422
1967
Lirula macrospora
4.
(Hartig) comb. nov.
H = Hysterium
(Hypoderma)
macrosporum
Hartig, Wicht. Krankh.
W a l d b . 101.1874.
s= Lophodermium
macrosporum
(Hartig) R e h m , in Rabh., K r y p t . - F l . II.
1(3): 4 5 . 1 8 8 7 .
^Hypodermopsis
macrospora ( H a r t i g ) O . K u n t z e , R e v . G e n . P I . 3(2):
487.1898.
= Hypodermella
macrospora
(Hartig) Lagerberg. M e d d . Stat. Skogsforsoksanst. 7: 1 1 3 . 1 9 1 0 .
—Lophodermium
filiforme D a r k e r , C o n t r . A r n o l d A r b o r . 1: 8 5 . 1 9 3 2 .
S T A T U S P Y C N I D I C U S : Hypodermina
k o s l o v . A k a d . Z e m e d . 3: 5 7 . 1 9 2 9 .
hartigii
Hilitzer, V e d . Spisy V y d . Ces-
O n the basis of spore s h a p e t h i s species has a l w a y s p r e s e n t e d a p r o b l e m to
t a x o n o m i s t s , f a l l i n g as i t does m o r e o r less i n t e r m e d i a t e l y b e t w e e n
Lophodermium a n d Hypodermella.
I t s general m o r p h o l o g y , life c y c l e , a n d s t r o n g l y p a r a s i t i c
n a t u r e suggest r e l a t i o n s h i p s w i t h Lirula
(the Hypodermella
g r o u p o n Abies)
r a t h e r t h a n w i t h Lophodermium
w h i c h is i n a g r e e m e n t w i t h L a g e r b e r g ' s earlier
c o n t e n t i o n . T h e r e is o n Picea a n aggregate of c l o s e l y r e l a t e d species or v a r i e t i e s ,
a n d a l t h o u g h the w r i t e r a c c e p t s for the p r e s e n t T e r r i e r ' s p l a c e m e n t of Lophodermium filiforme i n the s y n o n y m y of L. macrospora, f u r t h e r s t u d y m a y necessitate
i t s r e e s t a b l i s h m e n t as a d i s t i n c t t a x o n .
Lirula mirabilis
5.
= Hypodermella
(Darker) comb. nov.
mirabilis D a r k e r , C o n t r . A r n o l d A r b o r . 1: 4 6 . 1 9 3 2 .
Lirula nervata ( D a r k e r )
6.
=
Hypodermella
nervata
comb. nov.
D a r k e r , C o n t r . A r n o l d A r b o r e t u m 1: 5 1 . 1 9 3 2 .
Lirula punctata ( D a r k e r )
7.
= Hypodermella
IV.
Hypodermella v .
punctata
comb. nov.
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 48. 1932.
T u b . , B o t . C e n t r a l b l . 6 1 : 49. 1895.
H y s t e r o t h e c i a b l a c k , e l l i p t i c a l , often i n a m o r e o r less c o n t i n u o u s r o w ,
subcuticular; covering layer dark without evident opening mechanism;
h y m e n i u m flat, a r i s i n g f r o m a t h i n p a r e n c h y m a t o u s s u b h y m e n i a l l a y e r seated
on a t h i n b a s a l l a y e r of l i g h t b r o w n p s e u d o p a r e n c h y m a d a r k e n i n g b e l o w a n d
c o n t i n u o u s w i t h a c o m p a c t d a r k p s e u d o p a r e n c h y m a o c c u p y i n g the e p i d e r m i s
a n d to a lesser e x t e n t t h e h y p o d e r m i s . A s c i c l a v a t e , 4-spored a t m a t u r i t y .
Ascospores clavate. Paraphyses short, simple. P y c n i d i a numerous, black.
T Y P E - S P E C I E S : Hypodermella
laricis
v. Tub.
1. Hypodermella
laricis v . T u b . , B o t . C e n t r a l b l . 6 1 : 4 9 . 1 8 9 5 .
= Hypodermella
l a r i c i s v . T u b . v a r . octospora D e a r n . , M y c o l o g i a 18: 2 4 1 .
1926.
P Y C N I D I A L S T A T E : ? Leptothyrella
laricis
D e a r n . , M y c o l o g i a 2 0 : 240. 1928.
T h e t y p e of H. laricis v a r . octospora D e a r n . is q u i t e i m m a t u r e a n d i t is q u i t e
p r o b a b l e t h a t four of the e i g h t spores w o u l d h a v e a b o r t e d before r e a c h i n g
maturity.
DARKER: REVISION OF T H E HYPODERMATACEAE
DISPOSITION O F S Y N O N Y M O U S E P I T H E T S
1423
(HYPODERMELLA)
abietis-concoloris
( M a y r ) D e a r n . =Lirula
abietis-concoloris
ampla ( D a v i s ) D e a r n . = Davisomycella
ampla
arcuata D a r k e r = Lophodermella
arcuata
cerina D a r k e r = Lophodermella
cerina
concolor ( D e a r n . ) D a r k e r = Lophodermella
concolor
conjuncta D a r k e r ^Lophodermella
conjuncta
hiratsukae D a r k e r = Davisomycella
hiratsukae
lacrimiformis
D a r k e r = Davisomycella
lacrimiformis
laricis v . octospora D e a r n . = Hypodermella
laricis
limitata D a r k e r = Davisomycella
limitata
lirelliformis
D a r k e r = L i r u l a nervisequia
macrospora ( H a r t i g ) L a g e r b . = Lirula
macrospora
medusa D e a r n . = Davisomycella
medusa
mirabilis D a r k e r = Lirula
mirabilis
montana D a r k e r = Davisomycella
montana
montivaga ( P e t r . ) D e a r n . = Lophodermella
montivaga
montivaga v . concolor D e a r n . = Lophodermella
concolor
nervata D a r k e r = L i r u l a nervata
nervisequia ( D C . ex F r . ) L a g e r b . =Lirula
nervisequia
punctata D a r k e r ^Lirula
punctata
sulcigena ( R o s t r . ) v . T u b . = Lophodermella
sulcigena
V . D a v i s o m y c e l l a gen. n o v .
Hypodermatacearum
H y s t e r o t h e c i a i n n a t a , externe a t r a , p l e r u m q u e b r e v i a e l l i p t i c a q u e u s q u e
oblonga, aut elongata, subepidermalia, primordio Iongitudinalis incisurae
simplici parenchymatoso aut ignoto. H y m e n i u m s u b c u p u l a t u m ; s u b h y m e n i u m
tenue, h y a l i n u m . A s c i c l a v a t i v e l s a c c a t i . P a r a p h y s e s n u m e r o s a e
filiformes
simplices. Ascosporae clavatae a d basin attenuatae, i n m a t u r i t a t e non fascicuIatae u s q u e fasciculatae. P y c n i d i a , u b i n o t a , p a r v a a p p l a n a t a , superficiei acus
concoloria.
E T Y M O L O G I A : A D r . J . J . D a v i s q u i p r i m a m s p e c i e m h u j u s generis d e s c r i p s i t ,
mykes, fungus, et -ella, a d i m i n u t i v e suffix.
S P E C I E S T Y P I C A : Davisomycella
ampla
(Davis) Darker
H y s t e r o t h e c i a i n n a t e , e x t e r n a l l y b l a c k , for t h e m o s t p a r t s h o r t e l l i p t i c a l to
o b l o n g , o r elongate, s u b e p i d e r m a l , w i t h p r i m o r d i u m of t h e l o n g i t u d i n a l s l i t
simple, parenchymatous, or u n k n o w n . H y m e n i u m somewhat cupulate; subh y m e n i u m t h i n , hyaline. A s c i clavate or saccate. Paraphyses
numerous,
filiform,
s i m p l e . A s c o s p o r e s c l a v a t e , a t base a t t e n u a t e d , a t m a t u r i t y n o t
fasciculate to fasciculate. P y c n i d i a , where k n o w n , s m a l l flat, c o n c o l o r o u s w i t h
surface of t h e needle.
1.
Davisomycella ampla ( D a v i s ) c o m b .
nov.
= Lophodermium
pinastri ( S c h r a d . ex H o o k . ) C h e v . v a r . amplum D a v i s ,
T r a n s . W i s e . A c a d . S c i . A r t s & L e t t . 18: 2 5 2 . 1 9 1 8 .
=Lophodermium
amplum ( D a v i s ) D a v i s , I.e. 19: 6 9 5 . 1 9 1 9 .
= Hypodermella
ampla ( D a v i s ) D e a r n . , M y c o l o g i a 16: 1 5 2 . 1 9 2 4 .
CANADIAN JOURNAL OF BOTANY. VOL. 45,
1424
1967
Davisotnycelta hiratsukae ( D a r k e r )
2.
== Hypodermella
hiratsukae
comb. nov.
D a r k e r , J . A r n o l d A r b o r . 16: 3 6 4 . 1 9 3 5 .
Davisomycella lacrimiformis ( D a r k e r )
3.
= Hypodermella
comb. nov.
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 4 0 . 1 9 3 2 .
lacrimiformis
Davisomycella limitata ( D a r k e r )
4.
^= Hypodermella
limitata
comb. nov.
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 3 9 . 1 9 3 2 .
Davisomycella medusa ( D e a r n . )
5.
comb. nov.
medusa D e a r n . , M y c o l o g i a 16: 1 5 2 . 1 9 2 4 .
ass Hypodermella
Davisomycella montana ( D a r k e r )
6.
= Hypodermella
VI.
Ploioderma
montana
gen. n o v .
comb. nov.
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 4 4 . 1 9 3 2 .
Hypodermatacearum
G e n u s Davisomycellae
s i m i l e sed ascosporis b a c i l l a r i b u s . H y s t e r o t h e c i a a t r a
aut paene achroa, elliptica, subepidermalia. H y m e n i u m s u b c o n c a v u m ; subh y m e n i u m tenue, a c h r o u m . A s c i c l a v a t o - s a c c a t i . P a r a p h y s e s s i m p l i c e s ,
n l i f o r m e s . A s c o s p o r a e breves b a c i l l a r e s . P y c n i d i a s i m p l i c i a a p p l a n a t a , a u t
ignota.
E T Y M O L O G I A : ploion,
b o a t , et derma,
skin.
S P E C I E S T Y P I C A : Ploioderma
hedgcockii ( D e a r n . ) D a r k e r
G e n u s s i m i l a r to Davisomycella
b u t w i t h ascospores r o d - l i k e . H y s t e r o t h e c i a
b l a c k o r a l m o s t colorless, e l l i p t i c a l , s u b e p i d e r m a l . H y m e n i u m s o m e w h a t c o n c a v e ; s u b h y m e n i u m t h i n , colorless. A s c i c l a v a t e - s a c c a t e . P a r a p h y s e s s i m p l e ,
filiform. A s c o s p o r e s s h o r t , r o d - l i k e . P y c n i d i a s i m p l e , flat, o r u n k n o w n .
1.
Ploioderma hedgcockii ( D e a r n . )
^Hypoderma
S T A T U S P Y C N I D I C U S : Leptostroma
2.
hedgcockii D e a r n . , M y c o l o g i a 20: 240. 1928.
Ploioderma lethale ( D e a r n . )
ssHypoderma
3.
comb. nov.
hedgcockii D e a r n . , M y c o l o g i a 18: 240. 1926.
comb. nov.
lethale D e a r n . , M y c o l o g i a 18: 2 4 1 . 1 9 2 6 .
Ploioderma pedatum ( D a r k e r )
= Hypoderma
comb. nov.
pedatum D a r k e r , C o n t r . A r n o l d A r b o r . 1: 3 0 . 1 9 3 2 .
V I I . E L Y T R O D E R M A D a r k e r , C o n t r . A r n o l d A r b o r . 1: 6 2 . 1 9 3 2 .
H y s t e r o t h e c i a b l a c k , s h o r t e l l i p t i c a l to e l o n g a t e d , s u b e p i d e r m a l ; h y m e n i u m
c o n c a v e ; s u b h y m e n i a l l a y e r c o m p a r a t i v e l y t h i n , colorless. A s c i f u s i f o r m c l a v a t e . P a r a p h y s e s s i m p l e , filiform. A s c o s p o r e s large, f u s i f o r m , fasciculate,
2-celled. P y c n i d i a s i m p l e , c o n c o l o r o u s .
T Y P E - S P E C I E S : Elytroderma
deformans ( W e i r ) D a r k e r
1. Elytroderma deformans ( W e i r ) D a r k e r , C o n t r . A r n o l d A r b o r . 1: 6 3 . 1 9 3 2 .
= Hypoderma deformans W e i r , J . A g r i c . R e s . 6: 2 7 7 . 1 9 1 6 .
2.
Elytroderma hispanicum ( T o r r e s
Juan) comb. nov.
s= Hypoderma
hispanicum
Torres Juan, in Benito M a r t i n e z & Torres
J u a n , I n s t . F o r e s t a l I n v e s t . E x p e r . B o l . 88: 6 7 . 1 9 6 5 .
DARKER: REVISION OF T H E HYPODERMATACEAE
1425
V I I I - L O P H O D E R M E L L A v . H o h n . , B e r . deutsch. bot. Ges. 35: 247.1917.
H y s t e r o t h e c i a m o r e o r less c o n c o l o r o u s w i t h needle surface, s h o r t t o e l o n gated s u b h y p o d e r m a l , c i r c u l a r t o e l l i p t i c a l i n cross-section. H y m e n i u m c u p u late; s u b h y m e n i u m t h i n . A s c i clavate to subcylindric. Paraphyses simple,
filiform. A s c o s p o r e s c l a v a t e , s u b b i s e r i a t e t o f a s c i c u l a t e . P y c n i d i a s m a l l , flasklike, or u n k n o w n .
T Y P E - S P E C I E S : Lophodermella
1. Lophodermella
sulcigena
sulcigena
(Rostr.) v . H o h n .
( R o s t r . ) v . H o h n . , B e r . d e u t s c h . b o t . G e s . 3 5 : 247.
1917.
=
? Hypodermium
sulcigenum L k . , S p . P I . 6 ( 2 ) : 8 9 . 1 8 2 4 .
= ? Schizoderma sulcigenum
(Lk.) D u b y , in D e Candolle & D u b y , Bot.
G a l l . 2: 885.1830.
= Hypoderma sulcigenum R o s t r . , T i d s s k r . S k o v b r . 6: 2 8 4 . 1 8 8 3 .
= Hypodermella
sulcigena ( R o s t r . ) v . T u b . , B o t . C e n t r a l b l . 6 1 : 4 9 . 1 8 9 5 .
= Hypoderma pinicola B r u n c h . , B e r g e n s M u s . A a r b o g 8: 6 . 1 8 9 2 .
= Hypodermopsis
pinicola ( B r u n c h . ) 0 . K u n t z e , R e v . G e n . P I . 3 ( 2 ) : 4 8 7 .
1898.
T h e e p i t h e t Hypodermium
sulcigenum L k . w a s a p p l i e d t o a species g e n e r a l l y
t h o u g h t t o represent a p y c n i d i a l state. I n h i s t r e a t m e n t of the genus Hypodermium, v o n H o h n e l (1916) d i s m i s s e d the species sulcigenum
w i t h t h e brief
s t a t e m e n t t h a t i t w a s c o n s i d e r e d to be a n a s c o m y c e t e . A s t h e existence of a
c o n i d i a l s t a t e i n Lophodermella
sulcigena has n e v e r been d e m o n s t r a t e d i t seems
q u i t e possible t h a t v o n H o h n e l was c o r r e c t a n d t h a t L i n k a n d D u b y h a d the
ascocarp s t a t e before t h e m a l t h o u g h t h e d e s c r i p t i o n of the perfect s t a t e s t e m s
only from Rostrup's later account.
Hendersonia acicola v . T u b . w a s once c o n j e c t u r e d to be the i m p e r f e c t s t a t e of
Lophodermella
sulcigena, b u t the Hendersonia
is o n l y a n o t h e r of t h e n u m e r o u s
s e c o n d a r y f u n g i w h i c h a t t a c k needles i n f e c t e d b y h y p o d e r m a t a c e o u s species
a n d serve i n the n a t u r a l b i o l o g i c a l c o n t r o l of t h e l a t t e r . I n t h e r e l a t e d Hypodermella concolor, h o w e v e r , a careful search w i l l often r e v e a l here a n d t h e r e a s m a l l
flask-like
s t r u c t u r e b e a r i n g m i n u t e s p e r m a t i a - l i k e spores s e a t e d i n a s u b s t o m a t a l c h a m b e r . T h e s e m i n u t e , r u s t - l i k e s p e r m a t i a are n o t i n c o n s i s t e n t w i t h
w h a t one m i g h t e x p e c t t o f i n d i n t h i s deep-seated s u b h y p o d e r m a l genus. T h e
s t o m a w o u l d p r o v i d e a n a t u r a l e x i t for the s p e r m a t i a t h r o u g h the t h i c k
epidermis a n d hypodermis.
2.
Lophodermella arcuata ( D a r k e r )
^Hypodermella
3.
Lophodermella cerina ( D a r k e r )
= Hypodermella
4.
comb. nov.
arcuata D a r k e r , C o n t r . A r n o l d A r b o r . 1: 5 7 . 1 9 3 2 .
comb. nov.
cerina D a r k e r , C o n t r . A r n o l d A r b o r . 1: 6 1 . 1 9 3 2 .
Lophodermella concolor ( D e a r n . )
= Hypodermella
18: 242. 1926.
= Hypodermella
1932.
montivaga
comb. nov.
(Petr.) D e a r n . f. concolor D e a r n . M y c o l o g i a
concolor ( D e a r n . ) D a r k e r , C o n t r . A r n o l d A r b o r . 1: 59.
CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
1426
5.
Lophodermella conjuncta ( D a r k e r )
= Hypodermella
conjuncta
comb. nov.
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 6 0 . 1 9 3 2 .
6. Lophodermella montivaga P e t r . , i n P e t r . & S y d . , A n n . M y c . 2 0 : 1 9 1 . 1 9 2 2 .
= Hypodermella
montivaga ( P e t r . ) D e a r n . , M y c o l o g i a 16: 1 5 1 . 1 9 2 4 .
I X . B I F U S E L L A v . H o h n . , A n n . M y c . 15: 3 1 8 . 1 9 1 7 .
H y s t e r o t h e c i a b l a c k , e l l i p t i c a l , s u b c u t i c u l a r ; h y m e n i u m flat, s u b h y m e n i a l
layer thin. A s c i clavate. Paraphyses wanting or tending to disappear a t maturi t y . A s c o s p o r e s b i f u s i f o r m o r m o r e o r less r o d - l i k e a n d t e n d i n g t o w a r d s a
b i f u s i f o r m c o n d i t i o n . P y c n i d i a large or, i f s m a l l , t e n d i n g t o coalesce i n t o large
areas, c o n s p i c u o u s , c o v e r e d b y a t h i n p s e u d o p a r e n c h y m a t o u s l a y e r a t t a c h e d t o
the c u t i c l e ; s p e r m a t i a r e l a t i v e l y large, b a c i l l a r .
T Y P E - S P E C I E S : Bifusella
linearis
(Pk.) v. H o h n .
1. Bifusella linearis ( P k . ) v . H o h n . , A n n . M y c . 15: 3 1 8 . 1 9 1 7 .
s= Rhytisma lineare P k . , N . Y . S t . M u s . A n n . R e p t . 25 (1871): 1 0 0 . 1 8 7 3 .
^•Hypodermalineare
(Pk.) T h i i m . , Diagn. M y c o t h . U n i v . C e n t . X - X I I .
12.. 1878.
= Lophodermium
lineare ( P k . ) E l l . & E v . , N . A m e r . P y r e n . 7 2 1 . 1 8 9 2 .
= Hypodermopsis
linearis ( P k . ) 0 . K u n t z e , R e v . G e n . P l a n t . 3(2): 487.
1898.
PYCNIDIAL STATE:
2.
Bifusella pini ( D e a r n . )
= Hypoderma
1924.
^Hypoderma
3.
IHypodermium
robustum
pini
effusum
S c h w . , S y n . A m . B o r . 297. 1 8 2 2 .
comb. nov.
v . T u b . v a r . p i n i D e a r n . , M y c o l o g i a 16: 1 4 9 .
( D e a r n . ) D a r k e r , C o n t r . A r n o l d A r b o r . 1: 35. 1932.
Bifusella saccata ( D a r k e r )
= Hypoderma
comb. nov.
saccatum D a r k e r , C o n t r . A r n o l d A r b o r . 1: 3 3 . 1 9 3 2 .
DISPOSITION O F SYNONYMOUS E P I T H E T S
(BIFUSELLA)
abietis D e a r n . = Isthmiella
abietis
crepidiformis
D a r k e r = Isthmiella
crepidiformis
faullii D a r k e r = Isthmiella
faullii
striiformis D a r k e r = Soleella
striiformis
X.
D U P L I C A R I A F c k l . , S y m b . M y c . 265. 1870.
Hysterothecia black, elliptical, subcuticular, opening irregularly; h y m e n i u m
c u p u l a t e , a t m a t u r i t y flat, s u b h y m e n i a l l a y e r colorless, g r a d i n g i n t o a t h i n b a s a l
l a y e r of large b r o w n i s h cells i n c o n t a c t w i t h b r o w n d i s c o l o r e d e p i d e r m a l cells
occupied b y d a r k b r o w n hyphae. A s c i clavate-saccate. Ascospores bifusiform,
continuous, the lower section tending t o taper basally. Paraphyses simple,
f i l i f o r m , as l o n g as the a s c i . P y c n i d i a s m a l l , b l a c k .
T Y P E - S P E C I E S : Duplicaria
1. Duplicaria
empelri ( F r . ex F r . ) F c k l .
empelri ( F r . ex F r . ) F c k l . , S y m b . M y c . 2 6 5 . 1 8 7 0 .
s=XylomaempetriFr.,
Obs. M y c . 2: 363.1818.
^Rhytisma
empetri ( F r . ) F r . , V e t e n s k . A k a d . H a n d l . 1 0 5 . 1 8 1 9 .
^Rhytisma
empetri ( F r . ) e x F r . , E l e n c h . F u n g . 2: 1 2 7 . 1 8 2 8 .
DARKER: REVISION OF T H E HYPODERMATACEAE
P Y C N I D I A L S T A T E : Melasmia
1427
empetri M a g n u s , B e r . D e u t s c h . B o t . G e s . 4 : 104.
1886.
XI.
S o l e e l l a gen. nov. H y p o d e r m a t a c e a r u m
Hysterothecia nigra, elliptica usque elongato-linearia, secundum lineas
s t o m a t u m , s u b e p i d e r m a l i a ; h y m e n i u m a p p l a n a t u m ; s u b h y m e n i u m tenue.
A s c i s u b c l a v a t i ; p a r a p h y s e s s i m p l i c e s ; ascosporae bifusiformes, r a r o t r i f u s i formes. P y c n i d i a i g n o t a .
E T Y M O L O G I A : solea, s a n d a l , et -ella, a d i m i n u t i v e suffix, c
S P E C I E S T Y P I C A : Soleella striiformis ( D a r k e r ) D a r k e r
H y s t e r o t h e c i a b l a c k , e l l i p t i c a l t o e l o n g a t e d l i n e a r , a l o n g s t o m a t a l lines,
s u b e p i d e r m a l ; h y m e n i u m flat; s u b h y m e n i u m t h i n . A s c i s o m e w h a t c l a v a t e ;
p a r a p h y s e s s i m p l e , f i l i f o r m ; ascospores b i f u s i f o r m , o c c a s i o n a l l y t r i f u s i f o r m .
Pycnidia unknown.
1.
Soleella striiformis ( D a r k e r )
= Bifusella
striiformis
comb. nov.
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 2 3 . 1 9 3 2 .
X I I . H Y P O D E R M A D C . ex S a i n t A m a n s e m e n d . D e N o t . , G i o r n . B o t . I t a l .
2(2): (35). 1847.
= Hypoderma
D C , in L a m . & D C , F l . F r a n c . 3 e d . 2: 3 0 4 . 1 8 0 5 ; 3 e d . 6:
165. 1815.
= Hypoderma
D C . ex S a i n t A m a n s , F l . A g e n . 5 1 5 . 1 8 2 1 .
^Lophoderma
C h e v . , J o u r n . P h y s . 94: 3 1 . 1 8 2 2 .
= Hypoderma
D C . ex F c k l . , J a h r b . N a s s a u . V e r . N a t u r k . 2 3 - 2 4 : 257.
1870.
= Hypodermopsis
0 . K u n t z e , R e v . G e n . P l a n t . 3(2): 487.1898.
= EpidermellaTehon,
I l l i n o i s B i o l . M o n o g r . 13(4): 1 1 9 . 1 9 3 5 .
= ?Locelliderma T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4): 1 2 2 . 1 9 3 5 .
Hysterothecia elliptical, b l a c k ; asci clavate w i t h well developed s t a l k ;
ascospores r o d - s h a p e d o r f u s i f o r m , often w i t h a false s e p t u m o r b e c o m i n g oneseptate a t m a t u r i t y .
T Y P E - S P E C I E S : Hypoderma
rubi (Pers. ex C h e v . ) D e N o t . (typus cons.
prop.).
1. Hypoderma
rubi ( P e r s . ex C h e v . ) D e N o t . , G i o r n . B o t . I t a l . 2(2): (37). 1847.
^Hysterium
rubi P e r s . , O b s . M y c . 1: 84. 1796; S y n . F u n g . 1: 100. 1801.
^Hypoderma
virgultorum D C , F l . F r a n c . 3 e d . 6: 1 6 5 . 1 8 1 5 (an a v o w e d
substitute).
= Hypoderma
virgultorum
D C . ex S a i n t A m a n s , F l . A g e n . 515. 1821 [ut
virgultarum].
^Lophoderma
rubi (Pers.) ex C h e v . , J . P h y s . 94: 3 1 . 1 8 2 2 .
= Hypoderma rubi (Pers.) ex C h e v . , I.e. 3 1 . pro syn.
= Hysterium rubi (Pers.) ex F r . , S y s t . M y c . 2 : 5 8 7 . 1 8 2 3 .
=Lophodermium
rubi (Pers. ex C h e v . ) C h e v . , F l . P a r i s 1: 4 3 6 . 1 8 2 6 .
= Hypoderma
virgultorum
D C . f. rubi ( P e r s . ex C h e v . ) R e h m , i n R a b h .
K r y p t . - F l . I I . 1(3): 3 3 . 1 8 8 7 [ut (Pers.) D C . ] .
= Hypodermopsis
rubi ( P e r s . ex C h e v . ) O . K u n t z e , R e v . G e n . P l a n t .
3 ( 2 ) : 4 8 7 . 1 8 9 8 [ut (Pers.) O K . ] .
CANADIAN JOURNAL OF BOTANY. VOL. 45.
1428
1967
— Hypoderma
commune ( F r . ) D u b y f. rubi (Pers. ex C h e v . ) R e h m ,
B a y e r . B o t . G e s . M i i n c h e n 13: 112. 1912 [ut (Pers.) D C . ] .
2. Hypoderma
conigenum
(Pers.) D C . ex S. F . G r a y , N a t . A r r a n g m . B r i t .
P l a n t s , 5 0 9 . 1 8 2 1 [ut conigerum].
= Hysterium conigenum P e r s . , O b s . M y c . 1: 3 0 . 1 7 9 6 [ u t c o n i g e r u m ] ; S y n .
F u n g . 1: 102. 1801.
^Hypoderma
conigenum (Pers.) D C , F l . F r a n c . 3 e d . 2: 3 0 5 . 1 8 0 5 .
s= Hypodermopsis
conigena ( P e r s . ex S . F . G r a y ) O . K u n t z e , R e v . G e n .
P l a n t . 3 ( 2 ) : 487. 1898 [ut [P.] ( D C , C o o k e ) ] .
— Hysterium conigenum F r . , S y s t . M y c . 2: 5 8 6 . 1 8 2 3 .
== Hysterium
(Hypoderma)
conigenum ( F r . ) C k e . , H a n d b . B r i t . F u n g i 2:
762.1871.
s= Hypoderma
conigenum ( F r . ) C k e . , i n M a s s e e , B r i t . F u n g u s - F l o r a 4 :
3 5 . 1 8 9 5 [ut (Pers.) C k e . ] .
A s o r i g i n a l l y d e s c r i b e d Hysterium
conigenum
P e r s . w a s s a i d to o c c u r o n
Pinus abies, n o w k n o w n as Picea abies, whereas Hysterium
conigenum F r . w a s
r e p o r t e d o n Pinus silvestris. A l t h o u g h F r i e s d i d n o t refer to a n y p r e v i o u s use of
the e p i t h e t , R e h m , S a c c a r d o , a n d M a s s e e a l l a s c r i b e d i t t o P e r s o o n , b u t l i k e
F r i e s a n d C o o k e t h e y d e f i n i t e l y s t a t e d t h e host to be Pinus silvestris. U n t i l i t is
d e t e r m i n e d w h e t h e r we a r e d e a l i n g w i t h a g o o d species o r p e r h a p s w i t h t w o
species, the p r o b l e m of n o m e n c l a t u r e m u s t r e m a i n i n a b e y a n c e . M e a n w h i l e , as
s t a t e d b y R e h m (1887), the species r e m a i n s d o u b t f u l .
3. Hypoderma cunninghamiae
( K e i s s l . ) T e n g , S i n e n s i a 7: 2 6 1 . 1 9 3 6 .
= Hypoderma
strobicola v . T u b . f. cunninghamiae
Keissl., A k a d . Wiss.
W i e n , M a t h . - n a t . C I . A n z e i g e r 6 1 ( 2 ) : 13(4). 1942.
^HypodermahandeliiPetr.,
S y d o w i a 1: 3 7 1 . 1 9 4 7 .
4. Hypoderma
thujae D u r r i e u , S y d o w i a B e i h . 1: 3 5 6 . 1 9 5 7 .
DISPOSITION OF SYNONYMOUS E P I T H E T S
(HYPODERMA)
arundinaceum
( S c h r a d . ) D C . ex S a i n t A m a n s ^Lophodermium
arundinaceum
brachysporum
( R o s t r . ) v . T u b . = Meloderma
desmazierii
brachysporum
( R o s t r . ) O . K u n t z e = Meloderma
desmazierii
commune f. rubi (Pers. ex C h e v . ) R e h m = Hypoderma
rubi
conigenum ( F r . ) C k e . == Hypoderma conigenum (Pers.) D C . ex S. F . G r a y
deformans W e i r = Elytroderma
deformans
desmazierii D u b y = Meloderma
desmazierii
handelii P e t r . = Hypoderma
cunninghamiae
hedgcockii D e a r n . = Ploioderma
hedgcockii
hispanicum T o r r e s J u a n s = Elytroderma
hispanicum
juniperinum
( F r . ) O . K u n t z e = Lophodermium
juniperi
laricinum
( D u b y ) O . K u n t z e = Lophodermium
laricinum
lethale D e a r n . = Ploioderma
lethale
lineare ( P k . ) T h i i m . == Bifusella
linearis
namyslowskii
B ir u l a et a l . = Meloderma desmazierii nervisequum D C . = Lirula
nervisequia
pedatum D a r k e r = Ploioderma
pedatum
D A R K E R : REVISION O F T H E H Y P O D E R M A T A C E A E
1429
pinastri (Schrad.) D C . ^Lophodermium
pinastri
pini ( D e a r n . ) D a r k e r = Bifusella
pini
pinicola B r u n c h . == Lophodermella
sulcigena
robustum v . T u b . = Virgella robusta
robustum v a r . pini D e a r n . = Bifusella
pini
sabinae ( F a u t r . ) 0 . K u n t z e = Lophodermium
juniperi
saccatum D a r k e r = Bifusella
saccata
strobicola v . T u b . = Meloderma
desmazierii
strobicola f. cunninghamiae
K e i s s l . = Hypoderma
cunninghamiae
sulcigenum ( L k . ) R o s t r . = Lophodermella
sulcigena
virgultorum D C . = Hypoderma
rubi
virgultorum D C . ex S a i n t A m a n s ^Hypoderma
rubi
virgultorum f. rubi ( P e r s . ex C h e v . ) R e h m = Hypoderma
rubi
XIII.
Meloderma
gen. nov. H y p o d e r m a t a c e a r u m
H y s t e r o t h e c i a e l l i p t i c a , saepe i n o r d i n e , a t r a sed t y p i c e a c a n a z o n a c i r c u m data, s u b e p i d e r m a l i a sed s u b c u t i c u l a r i a p r o p e l a b i a s ; Iabiae o c h r o a e o r d i n e
externarum t u m i d a r u m c e l l u l a r u m ; tegmen a t r u m pseudoparenchymaticum,
tenue i n m a r g i n e ; s t r a t u m basilare p l e c t e n c h y m a t i c u m ; h y m e n i u m a p p l a n a t u m ; asci s u b c y l i n d r i c i ; p a r a p h y s e s p r i m o s i m p l i c e s filiformes d e n i q u e u n c i natae, afflatae, s u b r a m o s a e i n a p i c e ; ascosporae b a c i l l a r e s , s u b c l a v a t a e usque
fusiformes, m u c o i n v o l u t a e . P y c n i d i a s i m p l i c i a , a p p l a n a t a .
ETYMOLOGIA:
melas, b l a c k , et derma,
skin.
S P E C I E S T Y P I C A : Meloderma desmazierii ( D u b y ) D a r k e r
H y s t e r o t h e c i a e l l i p t i c a l , often i n a r o w , b l a c k b u t t y p i c a l l y s u r r o u n d e d b y a
g r a y i s h zone, s u b e p i d e r m a l , b u t s u b c u t i c u l a r n e a r the l i p s ; l i p s colorless w i t h a
row of o u t e r s w o l l e n c e l l s ; d a r k c o v e r i n g l a y e r p s e u d o p a r e n c h y m a t o u s , t h i n a t
margins; basal layer plectenchymatous; h y m e n i u m flat; asci s u b c y l i n d r i c a l ;
paraphyses a t first s i m p l e , f i l i f o r m , a t l e n g t h h o o k e d , s w o l l e n , s o m e w h a t
b r a n c h e d a t t i p ; ascospores r o d - l i k e , s o m e w h a t c l a v a t e t o f u s i f o r m , w i t h
gelatinous e n v e l o p e . P y c n i d i a s i m p l e , flat.
1.
Meloderma desmazierii ( D u b y ) c o m b . n o v .
s s Hypoderma desmazierii D u b y , M e m . S o c . P h y s . H i s t . N a t . G e n e v e 16:
54. 1861.
= Hypodermopsis
desmazierii
( D u b y ) O . K u n t z e , R e v . G e n . P I . 3(2):
4 8 7 . 1 8 9 8 [ut desmazieri].
=Lophodermium
brachysporum
R o s t r . , T i d s s k r . S k o v b r . 6: 2 8 1 . 1883.
= Hypoderma
brachysporum
v. T u b . Pflanzenkrankheiten durch krypto g a m e P a r a s i t e n v e r u r s a c h t , 247. 1895 n o n H. brachysporum Speg., B o l .
A c a d . N a c . C i e n c . C o r d o b a 11: 116 [repr. pag.]. 1887.
= Hypoderma
brachysporum
(Rostr.) O . K u n t z e , R e v . G e n . PI. 3(2):
487.1898.
= Hypoderma
strobicola v . T u b . , i n v . T u b . & S m i t h , Diseases of P l a n t s
Induced b y C r y p t o g a m i c Parasites, 233.1897.
= Lophodermium
lineatum S m i t h & R a m s b o t t o m , T r a n s . B r i t . M y c . S o c .
6: 3 6 5 . 1 9 2 0 .
1430
CANADIAN JOURNAL OF BOTANY. VOL. 45,
= Hypoderma
35. 1928.
namyslowskii
B i r u l a et a l . , R o c z n . N a u k R o l n . L e s n y c h , 20-
S T A T U S P Y C N I D I C U S : Leptostroma
strobicola
s l o v . A k a d . Z e m e d . 3: 99, 149. 1929.
XIV.
1967
Hilitzer, V e d . Spisy V y d . Cesko-
L O P H O D E R M I U M C h e v . , F l . P a r i s 1: 4 3 5 . 1 8 2 6 (nom. cons, prop.)
= Hypoderma
C h e v . , J o u r n . P h y s . 94: 3 1 . 1 8 2 2 .
= Aporia D u b y , M e m . S o c . P h y s . H i s t . N a t . G e n e v e 16: 5 1 . 1 8 6 1 .
= Lophodermina
v . H o h n . , B e r . Deutsch. B o t . Ges. 35: 418.1917.
= Lophodermellinav.
H o h n . , Ber. Deutsch. B o t . Ges. 35: 419.1917.
= Dermascia T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4): 6 0 . 1 9 3 5 .
H y s t e r o t h e c i a i n n a t e e r u m p e n t , e l l i p t i c a l , d u l l to s h i n i n g b l a c k , o p e n i n g b y a
longitudinal slit, opening mechanism well developed or w a n t i n g ; h y m e n i u m
m o r e o r less flat. A s c i n a r r o w l y c y l i n d r i c to c y l i n d r i c c l a v a t e , 8-spored, o c c a s i o n a l l y 4-spored w i t h 4 a b o r t e d spores present, m o s t l y s u b a c u t e a t the a p e x .
P a r a p h y s e s s i m p l e f i l i f o r m . A s c o s p o r e s s i m p l e , m o r e o r less u n i f o r m l y f i l i f o r m ,
h y a l i n e , fasciculate, u n i c e l l u l a r , s u r r o u n d e d b y a t h i n g e l a t i n o u s s h e a t h .
P y c n i d i a s i m p l e , r o u n d o r e l l i p t i c a l , c o n c o l o r o u s w i t h host o r c o l o r e d , o r b l a c k ,
or u n k n o w n . Spermatia simple, bacillar.
T Y P E - S P E C I E S : Lophodermium
(typus cons. prop.).
arundinaceum
( S c h r a d . ex S a i n t A m a n s ) C h e v .
1. Lophodermium
arundinaceum
( S c h r a d . ex S a i n t A m a n s ) C h e v . , F l . P a r i s 1:
435.1826.
= Hysterium
arundinaceum
S c h r a d . , S c h r a d . J . B o t . 2: 6 8 . 1 7 9 9 .
= Xyloma arundinaceum
(Schrad.) R e b . , P r o d . F l o r . N e o m a r c h . , 342.
1804.
= Hypoderma
arundinaceum
( S c h r a d . ) D C , F l . F r a n c . , 3 ed. 2 : 305.
1805.
= Hypoderma
arundinaceum
(Schrad.) D C . ex S a i n t A m a n s , F l . A g e n .
516.1821.
2.
Lophodermium aucupariae
(Schleich.) c o m b . n o v .
= Hysterium
aucupariae S c h l e i c h . , P l a n t a e C r y p t . H e l v e t i a e , E x s . n . 6 3 ,
1805; Catalogus P l a n t . H e l v e t i a , 58.1821.
= Hypoderma
xylomoides D C . f aucupariae
(Schleich.) D C , i n L a m . &
D C , F l . F r a n c . 3 e d . , 6: 1 6 5 . 1 8 1 5 .
= Lophodermium
tumidum R e h m ex N a n n f . , N o v a A c t a R e g . S o c . S c i e n t .
U p s a l . I V . 8 ( 2 ) : 2 3 7 . 1 9 3 2 . N o n L . tumidum ( F r . ) R e h m , i n R a b h . , K r y p t . F l . I I . 1(3): 4 0 . 1 8 8 7 , = Hysterium
tumidum F r . , S y s t . M y c . 2: 591. 1823.
= Coccomyces coronatus ( S c h u m . ex F r . ) D e N o t . , E r b a r . C r i t t . I t a l . I .
N o . 236.1859.
S T A T U S P Y C N I D I C U S : Leptostroma
A k a d . Z e m e d . 3: 8 0 , 1 4 6 . 1 9 2 9 .
sorbicolum
Hilitzer, V e d . Spisy Ceskosl.
U n d e r Hysterium
tumidum F r i e s is d e s c r i b e d a fungus n o w g e n e r a l l y c o n s i d e r e d s y n o n y m o u s w i t h Coccomyces coronatus ( S c h u m . ex F r . ) D e N o t . w h i c h
DARKER: REVISION OF T H E HYPODERMATACEAE
1431
D e N o t a r i s h a d earlier c a l l e d Coccomyces tumida ( F r . ) D e N o t . F u c k e l (1870)
d e t e r m i n e d m a t e r i a l o n Sorbus as C. tumida a n d issued i t i n F . r h e n . 746 u n d e r
t h a t n a m e . R e h m a c c e p t e d F u c k e l ' s d e t e r m i n a t i o n b u t h a v i n g n o t e d t h a t the
specimen w a s a Lophodermium
m a d e the c o m b i n a t i o n L. tumidum ( F r . ) R e h m .
N a n n f e l d t p o i n t e d o u t t h a t R e h m h a d n o t been d e a l i n g w i t h the s a m e species as
F r i e s a n d d r o p p e d the n a m e o f F r i e s f r o m the e p i t h e t m a k i n g i t r e a d L. tumidum
R e h m . U n f o r t u n a t e l y the n a m e s u p p l i e d b y N a n n f e l d t is a l a t e r h o m o n y m of
L. tumidum ( F r . ) R e h m a n d is b a s e d o n a different t y p e s p e c i m e n . T h e r e i s ,
however, a n e a r l i e r v a l i d e p i t h e t for t h i s species o n Sorbus. S c h l e i c h e r c a l l e d i t
Hysterium
aucupariae
S c h l e i c h . i n 1805 a n d t h e n a m e w a s also u s e d as a
v a r i e t a l e p i t h e t b y D e C a n d o l l e as Hypoderma
xylomoides
DC.f
aucupariae
(Schleich.) D C . A s Lophodermium
aucupariae (Schleich.) i t a p p e a r s t o p r o v i d e
a n a c c e p t a b l e a n d v e r y a p p r o p r i a t e n a m e for t h i s w e l l k n o w n species.
3. Lophodermium
cedrinum
M a i r e , B u l l . S o c . H i s t . N a t . A f r . N o r d 8: 174. 1917.
P Y C N I D I A L S T A T E : Labrella cedrina D u r . & M o n t . , L ' E x p l o r a t i o n s c i e n t i f i q u e
de 1'Algerie, C r y p t o g a m i e , 1: 5 9 9 . 1 8 4 9 .
4. Lophodermium
1911.
chamaecyparisii
5. Lophodermium
conigenum
A k a d . Z e m e d . 3: 7 6 . 1 9 2 9 .
S h a r a i & H a r a , B o t . M a g . T o k y o 2 5 : 69.
(Brunaud) Hilitzer, V e d . Spisy V y d . Ceskosl.
=Lophodermium
pinastri ( S c h r a d . ex H o o k . ) C h e v . f. conigena B r u n a u d ,
A c t . Soc. L i n n . B o r d e a u x 42: 97.1888.
^Lophodermina
conigena ( B r u n a u d ) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4):
92.1935.
6. Lophodermium
consociatum D a r k e r , C o n t r . A r n o l d A r b o r . 1: 7 9 . 1 9 3 2 .
== D e r m a s c i a c o n s o c i a t a ( D a r k e r ) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4):
63.1935.
7. Lophodermium
crassum D a r k e r , C o n t r . A r n o l d A r b o r . 1: 8 8 . 1 9 3 2 .
8. Lophodermium
durilabrum
9.
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 8 7 . 1 9 3 2 .
Lophodermium juniperi ( G r e v . ) c o m b . n o v .
s= Hysterium juniperi G r e v . , S c o t t . C r y p t . F l . 1: t a b . 2 6 . 1 8 2 2 .
= Hysterium juniperinum
F r . , O b s . M y c . 2: 3 5 5 . 1 8 1 8 .
s^Hysterium
pinastri [3 H. juniperinum
F r . , S y s t . M y c . 2: 588. 1823.
== Lophodermium
juniperinum
(Fr.) D e N o t . , G i o r n . B o t . I t a l . 2(2):
(46). 18-47.
= Hypoderma juniperinum
( F r . ) O . K u n t z e , R e v . G e n . P I . 3 ( 2 ) : 487.
1898.
= Lophodermina
juniperina
( F r . ) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4):
96.1935.
== Lophodermium
juniperinum
v a r . sabinae F c k l . , J a h r b . N a s s a u . V e r .
N a t u r k . 2 3 - 2 4 : 256.1870.
= Lophodermium
juniperinum
f. cupressi-thyoidis
S a c c , M i c h . 2 : 570.
1882.
1432
CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
=; Lophodermina
cupressi-thyoidis
(Sacc.) T e h o n , I l l i n o i s B i o l . M o n g r
13(4): 94. 1935.
=Lophodermium
sabinae F a u t r e y , in R o u r a . , R e v . M y c . 13: 1 6 9 . 1 8 9 1 .
s= Hypoderma
sabinae ( F a u t r e y ) 0 . K u n t z e , R e v . G e n . P I . 3 ( 2 ) : 487.
1898.
10. Lophodermium
lacerum D a r k e r , C o n t r . A r n o l d A r b o r . 1: 8 0 . 1 9 3 2 .
= Dermascia
lacera ( D a r k e r ) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4): 68.
1935.
11. Lophodermium
laricinum
D u b y , M e m . S o c . P h y s . H i s t . N a t . G e n e v e 16:
58.1861.
= Hypoderma
laricinum
( D u b y ) O . K u n t z e , R e v . G e n . P L 3 ( 2 ) : 487.
1898.
Lophodermina
laricina ( D u b y ) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4): 98.
1935.
12. Lophodermium
nitens D a r k e r , C o n t r . A r n o l d A r b o r . 1: 7 4 . 1 9 3 2 .
^Lophodermina
nitens ( D a r k e r ) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4):
102.1935.
13. Lophodermiumpetrakii
D u r r i e u , S y d o w i a B e i h . 1: 3 5 5 . 1 9 5 7 .
T h i s is p r o b a b l y t h e s a m e species d e s c r i b e d o r c i t e d earlier as f o l l o w s :
T e n g , C o n t r . B i o l . L a b . S c i . S o c . C h i n a , B o t . 8: 7, 1932 [ut
Lophodermium
pinastri]; T e n g , S i n e n s i a 4 : 137, 1933 [ut Lophodermium
uncinatum];
T e n g , I.e.,
7: 2 6 2 , 1 9 3 6 , a n d P e t r a k , S y d o w i a 1: 3 7 3 , 1 9 4 7 [ u t L o p h o d e r m i u m sp.].
14. Lophodermium
piceae ( F c k l . ) v . H o h n . , S i t z b . A k a d . W i s s . W i e n , M a t h . n a t u r w . K l . 126(1): 2 9 6 . 1 9 1 7 .
= Phacidium piceae F c k l . , J a h r b . N a s s a u . V e r . N a t u r k . 2 7 - 2 8 : 5 1 . 1 8 7 3 .
s= Coccomyces piceae ( F c k l . ) R e h m , i n R a b h . , K r y p t . - F l . I I . 1(3): 80.
1888.
= Coccomyces piceae ( F c k l . ) S a c c , S y l l . F u n g . 8: 7 4 6 . 1 8 8 9 .
= Lophodermium
abietis R o s t r . , T i d s s k r . S k o v b r . 12: 2 0 1 . 1 8 9 1 .
P Y C N I D I A L S T A T E : Leptostroma
abietis D e a r n . , N . Y . S t . M u s . B u l l . 266: 6 5 .
1925.
= Hypodermina
abietis ( D e a r n . ) H i l i t z e r , V e d . S p i s y V y d . C e s k o s l .
A k a d . Z e m e d . 3: 6 0 , 1 4 6 . 1 9 2 9 .
15. Lophodermium
pinastri
( S c h r a d . ex H o o k . ) C h e v . , F l . P a r i s 1: 436. 1826.
^Hysterium
pinastri S c h r a d . , S c h r a d . J o u r n . B o t . 2: 69. 1799.
^Hypodermapinastri
(Schrad.) D C , F l . F r a n c . 3 ed. 2: 305. 1805.
^Hysterium
pinastri S c h r a d . ex H o o k . , F l . S c o t i a 2: 8. 1821.
= Hysterium pinastri S c h r a d . ex F r . , S y s t . M y c . 2: 5 8 7 . 1 8 2 3 .
^Lophodermellina
pinastri ( S c h r a d . ex H o o k . ) v . H o h n . , A n n . M y c . 15:
311.1917.
= Lophodermium
pinicolum
T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4): 55.
1935.
— HysteriumUmitatumWibel,
P r i m . F l . W e r t h . 329.1799.
DARKER: REVISION OF T H E HYPODERMATACEAE
1433
= Aporia obscura D u b y , M e m . S o c . P h y s . H i s t . N a t . G e n e v e 16: 5 1 . 1 8 6 1 .
== Schizothyrium
obscurum ( D u b y ) S a c c , S y l l . F u n g . 2: 7 2 5 . 1 8 8 3 .
= Lophodermium
baculiferum
Mayr, Die Waldungen von Nordamerika,
313.1890 (nom. nud.).
= Scolecodothispinicola
M i l e s , M y c o l o g i a 18: 165. 1926.
== Lophodermiumaustrale
D e a r n . , M y c o l o g i a 18: 2 4 2 . 1 9 2 6 .
=Lophodermium
laricis D e a r n . , M y c o l o g i a 18: 2 4 3 . 1 9 2 6 .
P Y C N I D I A L S T A T E : Leptostroma
pinastri
D e s m . , A n n . S c . N a t . I I . 1 9 : 138.
1843.
= Depazea linearis R o s t r . , T i d s s k r . S k o v b r . 6: 2 6 0 . 1 8 8 3 .
^GloeosporiumpiniOud.,
N e d e r l . K r u i d k . A r c h . I I I . 2: 7 5 4 . 1 9 0 2 .
16. Lophodermium
1954.
pini-excelsae
17. Lophodermium
pini-pumilae
T o k y o , 53: 151.1952.
A h m a d , i n P e t r . & A h m a d , S y d o w i a 8: 172.
Sawada, B u l l . G o v t . F o r . E x p t . Stat., M e g u r o ,
18. Lophodermium sawadae nom. nov.
s= Lophodermium
thujae S a w a d a , R e p t . F o r . E x p t . S t a t . , T o k y o , 4 6 , 1 3 9 .
1950 ( n o n i . thujae D a v i s , T r a n s . W i s e A c a d . 2 0 , 4 2 4 . 1 9 2 2 ) .
19. Lophodermium
thujae D a v i s , T r a n s . W i s e . A c a d . 20, 4 2 4 . 1922 [ut thuyae].
= Lophodermina
thujae ( D a v i s ) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4):
109. 1935.
20. Lophodermium
1950.
thujopsidis
S a w a d a , R e p . F o r . E x p t . Stat., T o k y o , 46: 141.
21. Lophodermium
uncinatum
D a r k e r , C o n t r . A r n o l d A r b o r . 1: 7 6 . 1 9 3 2 .
= Lophodermina
uncinata ( D a r k e r ) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4):
110. 1935.
DISPOSITION O F S Y N O N Y M O U S E P I T H E T S
(LOPHODERMIUM)
abietis R o s t r u p =Lophodermium
piceae
abietis-concoloris
M a y r = Lirula
abietis-concoloris
agharkarii T i l a k = Lophomerum
agharkarii
amplum D a v i s = Davisomycella
ampla
australe D e a r n . =Lophodermium
pinastri
autumnale D a r k e r = Lophomerum
autumnale
baculiferum M a y r = Lophodermium
pinastri
brachysporum
R o s t r . = Meloderma
desmazierii
clavuligerum S p e g . ^Lophomerum
clavuligerum
filiforme D a r k e r = Lirula
macrospora
gilvum R o s t r . = Naemacyclus
niveus
infectans M a y r = Virgella robusta
juniperinum
I. cupressi-thyoidis
S a c c . —Lophodermium
juniperi
laricis D e a r n . ==Lophodermium pinastri
lineare ( P k . ) E . & E . = Bifusella
linearis
lineatum S m i t h & R a m s b o t t o m = Meloderma
desmazierii
macrosporum
( H a r t i g ) R e h m =Lirula
macrospora
CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
1434
nervisequium
( D C . ex F r . ) C h e v . =Lirula
nervisequia
nervisequium
( D C . ex F r . ) R e h m ^Lirula
nervisequia
phacidium
D e N o t . = Coccomyces
coronatus
pinastri v a r . amplum D a v i s = Davisomycella
ampla
pinicolum T e h o n = Lophodermium
pinastri
quercus B o s e & M u l l e r = Lophomerum
quercus
sabinae F a u t r e y = Lophodermium
juniperi
septatum T e h o n = Lophomerum
septatum
thujae S a w a d a = Lophodermium
sawadae
tumidum ( F r . ) R e h m = Coccomyces
coronatus
tumidum R e h m ex N a n n f . —Lophodermium
aucupariae
XV.
LOPHOMERUM
O u e l l e t t e & M a g a s i , M y c o l o g i a 58: 275. 1966.
G e n u s s i m i l a r t o Lophodermium
but w i t h phragmosporous
ascospores.
H y s t e r o t h e c i a b l a c k , e l l i p t i c , o p e n i n g b y a l o n g i t u d i n a l fissure, m o s t l y s u b c u t i c u l a r . C o v e r i n g l a y e r b l a c k , p s e u d o p a r e n c h y m a t o u s . H y m e n i u m flat;
s u b h y m e n i u m colorless, p l e c t e n c h y m a t o u s . A s c i s u b c y l i n d r i c , 8-spored.
P a r a p h y s e s s i m p l e , f i l i f o r m . A s c o s p o r e s f i l i f o r m , b e c o m i n g 3 to several-septate
a t m a t u r i t y . P y c n i d i a s i m p l e , flat; c o n i d i a b a c i l l a r .
T Y P E - S P E C I E S : Lophomerum
autumnale
(Darker) Magasi
1. Lophomerum
autumnale ( D a r k e r ) M a g a s i , i n O u e l l e t t e & M a g a s i , M y c o l o g i a
58: 2 7 5 . 1 9 6 6 .
=s Lophodermium
autumnale D a r k e r , C o n t r . A r n o l d A r b o r . 1: 77. 1932.
= Lophodermina
autumnalis
(Darker) T e h o n , Illinois B i o l . M o n o g r .
13(4): 8 9 . 1 9 3 5 .
2.
Lophomerum agharkarii ( T i l a k )
^Lophodermium
3.
agharkariiTilak,Sydowia
comb. nov.
13: 2 8 . 1 9 5 9 .
Lophomerum clavuligerum (Speg.)
comb. nov.
S 3 Lophodermium
clavuligerum
Speg., B o l . A c a d . N a c . Cienc. C o r d o b a
11: 118 ( r e p r i n t p a g . ) . 1887.
= Lophodermina
clavuligera (Speg.) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4):
91. 1935.
4. Lophomerum
1966.
darkeri
O u e l l e t t e , i n O u e l l e t t e & M a g a s i , M y c o l o g i a 58: 276.
5. Lophomerum
septatum
( T e h o n ) ex O u e l l e t t e , i n O u e l l e t t e & M a g a s i ,
M y c o l o g i a 58: 2 7 9 . 1 9 6 6 .
= Lophodermina
septata T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4): 109. 1935.
= Lophodermium
septatum ( T e h o n ) T e r r i e r , M a t e r . F l o r e C r y p t . Suisse
9(2): 32.1942.
6.
Lophomerum quercus ( B o s e & E . M u l l e r )
comb. nov.
= Lophodermium
quercus [ut querci] B o s e & E . M u l l e r , I n d i a n P h y t o p a t h . 18: 3 4 0 . 1 9 6 5 .
DARKER: REVISION OF T H E HYPODERMATACEAE
XVI
1435
C O C C O M Y C E S D e N o t . , G i o r n . B o t . I t a l . 2 ( 2 ) : (38). 1847.
^= Coccomycellav.
H o h n . , A n n . M y c . 15: 3 2 3 . 1 9 1 7 .
A s c o c a r p s o n leaves a n d t w i g s , s u b c i r c u l a r to p o l y g o n a l , i n n a t e , c o n n a t e w i t h
host tissue, b l a c k t o m o r e o r less c o n c o l o r o u s ; o p e n i n g l o b a t e l y o r r a d i a l l y ;
opening m e c h a n i s m w e a k l y o r f a i r l y w e l l d e v e l o p e d ; h y m e n i u m flat, colorless
to y e l l o w i s h - o r a n g e ; s u b h y m e n i u m t h i n ; a s c i s u b c l a v a t e , s o m e t i m e s s o m e w h a t
s t a l k e d ; p a r a p h y s e s u s u a l l y exceeding the a s c i , c o i l e d o r u n c i n a t e t o c l u b shaped a t the t i p ; ascospores c o n t i n u o u s , c y l i n d r i c a l t o s u b c l a v a t e , m o r e o r less
fasciculate, s u r r o u n d e d b y a definite n a r r o w s h e a t h .
T Y P E - S P E C I E S : Coccomyces coronatus
1. Coccomyces coronatus
( S c h u m . ex F r . ) D e N o t .
( S c h u m . ex F r . ) D e N o t . , E r b a r . C r i t t . I t a l . I . N o . 236.
1859.
=
=
=
=
=
= Ascobolus coronatus S c h u m . , E n u m . P l a n t S a e l l . 2: 4 3 7 . 1 8 0 3 .
=2 Phacidium
coronatum ( S c h u m . ) F r . , O b s . M y c . 1: 1 6 7 . 1 8 1 5 .
= Phacidium
coronatum ( S c h u m . ) ex F r . , S y s t . M y c . 2: 5 7 7 . 1 8 2 3 .
= Lophodermium
phacidium
de N o t . , G i o r n . B o t . I t a l . 2 ( 2 ) : (42). 1847
(an a v o w e d s u b s t i t u t e ) .
= Coccomyces coronatus ( S c h u m . ex F r . ) K a r s t . , B i d r . K a n n . F i n l a n d s
N a t u r . F o l k l 9 : 256.1871.
Peziza comitialis B a t s c h , E l e n c h . F u n g . C o n t . 1: 2 1 7 . 1 7 8 6 .
= Coccomyces comitialis
( B a t s c h ) ex D e a r n . & H o u s e , N . Y . S t . M u s .
B u l l . 266: 6 5 . 1 9 2 5 .
Pezizaviridis
Bolt., H i s t . F u n g . Halifax, 109.1789.
Xyloma pezizoides P e r s . , S y n . fung. 1 0 5 . 1 8 0 1 .
Peziza connivens M a r t i u s , F l . C r y p t . E r l a n g . 4 6 3 . 1 8 1 7 .
Hysterium
tumidum F r . , S y s t . M y c . 2: 5 9 1 . 1 8 2 3 .
= Coccomyces tumidus ( F r . ) D e N o t . , G i o r n . B o t . I t a l . 2 ( 2 ) : (39). 1847.
= Lophodermium
tumidum ( F r . ) R e h m , i n R a b h . , K r y p t . - F l . I I . 1(3):
40. 1887.
T h e genus Coccomyces c o m p r i s e s a large n u m b e r of species some of w h i c h
s h o w affinities to the genus Lophodermium
a n d o t h e r s s u p e r f i c i a l l y a t least t o
the P h a c i d i a c e a e .
XVII.
C O L P O M A W a l l r . , F l . C r y p t . G e r m . 2: 4 2 2 . 1 8 3 3 .
= Cenangium F r . s u b g . [ut t r i b u s ] Cliihris F r . , S y s t . M y c . 2: 1 8 9 . 1 8 2 2 .
= Cliihris ( F r . ) E n d l . , G e n . P l a n t . 3 2 . 1 8 3 6 .
= Sporomega C o r d a , I c o n . F u n g . 5: 3 4 . 1 8 4 0 .
= Pragmoparopsis
v. H o h n . , A n n . M y c . 15: 3 2 0 . 1 9 1 7 .
F r u i t - b o d y e l o n g a t e d , often c u r v e d , o r s h o r t , u n d e r p e r i d e r m o r b r e a k i n g
through naked w o o d ; opening usually b y a long slit; covering layer well
d e v e l o p e d ; b a s a l l a y e r also w e l l d e v e l o p e d , t h i c k , often s o m e w h a t d a r k
colored; h y m e n i u m subcupulate; s u b h y m e n i u m conspicuous, h y a l i n e ; asci
c l a v a t e w i t h a n a r r o w e d b a s a l s t a l k ; p a r a p h y s e s a b u n d a n t , slender, often
c o i l e d o r b e n t a t t i p ; ascospores c o n t i n u o u s , m o r e o r less f a s c i c u l a t e , n a r r o w l y
filiform,
d e l i c a t e l y s h e a t h e d , o c c u p y i n g the u p p e r s w o l l e n p o r t i o n of the ascus.
T Y P E - S P E C I E S : Colpoma quercinum
( P e r s . ex S a i n t A m a n s ) W a l l r .
CANADIAN JOURNAL OF BOTANY. VOL. 45,
1436
1. Colpoma quercinum
1967
(Pers. ex S a i n t A m a n s ) W a l l r . , F l . C r y p t . G e r m . 2: 423.
1833.
== Hysterium
quercinum
P e r s . , O b s . M y c . 1: 8 3 . 1796; S y n . F u n g . 100.
1801.
== Hypoderma quercinum (Pers.) D C , F l . F r a n c . 3 e d . 2 : 3 0 6 . 1 8 0 5 .
== Hypoderma
quercinum
(Pers.) D C . ex S a i n t A m a n s , F l . A g e n . 515.
1821.
= Tryblidium
quercinum (Pers. ex S a i n t A m a n s ) P e r s . , M y c . E u r . 1: 333.
1822.
= Cenangium
s u b g . Clithris quercinum
(Pers. ex S a i n t A m a n s ) F r . ,
S y s t . M y c . 2: 1 8 9 . 1 8 2 2 [ut (Pers.)].
= Clithris quercina ( P e r s . ex S a i n t A m a n s ) K a r s t . , N o t . F a u n a F l o r a
F e n n . 11: 2 6 0 . 1 8 7 0 ; B i d r . K a n n . F i n l . N a t . F o l k , 19: 2 2 1 . 1 8 7 1 .
=
Clithris quercina
( P e r s . ex S a i n t A m a n s ) S c h r o t . , K r y p t . - F l o r a
Schles.3(2): 165.1893.
= Hysterium nigrum T o d e , F u n g i M e c k l . 2: 5 . 1 7 9 1 .
= Clithris nigra ( T o d e ) K e i s s l . , K r y p t . M u s . V i n d o b . 523.
= Colpoma nigra ( T o d e ) v . H o h n . , A n n . M y c . 15: 3 1 9 . 1 9 1 7 .
= Variolaria
corrugata B u l l . , H i s t . C h a m p . [ V e n t e n a t E d . ] 1: 187. p i . 432,
f. 4 . 1 8 0 9 .
= Colpoma corrugata ( B u l l . ) e x O . K u n t z e , R e v . G e n . P I . 3(2): 4 8 7 . 1 8 9 8 .
= Sphaeria collapsa S o w e r b . , E n g l . F u n g . t. 373, f. 3. 1 7 9 7 - 1 8 0 9 .
X V I I I . X Y L O S C H I Z O N S y d . , i n S y d . & Petr., A n n . M y c . 20: 192.1922.
H y s t e r o t h e c i a elliptical to elongated, scattered or crowded o n bare w o o d , at
first c o v e r e d , b e c o m i n g e r u m p e n t a n d finally a l m o s t s u p e r f i c i a l , o p e n i n g b y a
l o n g i t u d i n a l s l i t , c o v e r i n g a n d b a s a l l a y e r s w e l l d e v e l o p e d ; a s c i slender a n d
l o n g - c l a v a t e , n a r r o w e d d o w n w a r d s i n t o a l o n g , t h i c k s t a l k ; ascospores elongatec y l i n d r i c a l , h y a l i n e , w i t h 3 d e l i c a t e s e p t a ; p a r a p h y s e s s i m p l e , n u m e r o u s , often
h o o k e d a t the t i p .
T Y P E - S P E C I E S : Xyloschizon
1. Xyloschizon
weirianum
weirianum
Sydow.
S y d . , i n S y d . & Petr., A n n . M y c . 20: 192.1922.
A second species, X. atratum S y d . , I.e. p . 193, a p p e a r s n o t to differ m a r k e d l y
f r o m the w o o d i n h a b i t i n g forms of Colpoma
crispum.
XIX.
B I F U S E P T A D a r k e r , M y c o l o g i a 55: 816.1963.
H y s t e r o t h e c i a a p p e a r i n g g o l d e n b r o w n to g r a y i s h b l a c k , e l l i p t i c a l , s u b c u t i c u l a r ; c o v e r i n g l a y e r of d a r k p s e u d o p a r e n c h y m a a l t e r n a t i n g w i t h wedges of
p a r e n c h y m a ; h y m e n i u m flat; s u b h y m e n i u m t h i n , seated o n a d a r k e n e d b a s a l
layer. A s c i clavate. Paraphyses simple, filiform. Ascospores bifusiform, phragmosporous. P c y n i d i a small, black, circular. S p e r m a t i a bacillar.
T Y P E - S P E C I E S : Bifusepta
tehonii D a r k e r .
1. Bifusepta tehonii D a r k e r , M y c o l o g i a 5 5 : 8 1 6 . 1 9 6 3 .
= Bifusella vaccinii T e h o n , M y c o l o g i a 3 1 : 6 7 8 . 1 9 3 9 .
N o n 73. vaccinii (Schrot.) T e h o n , I l l i n o i s B i o l . M o n o g r . 13(4): 1 1 7 . 1 9 3 5 .
D A R K E R : REVISION O F T H E H Y P O D E R M A T A C E A E
X X
1437
R H Y T I S M A F r . , S y s t . M y c . 2: 5 6 5 . 1 8 2 3 .
== Melanosorus D e N o t . , G i o r n . B o t . I t a l . 2 ( 2 ) : (49). 1847.
= Xyloma P e r s . , T e n t . D i s p . F u n g . 5 . 1 7 9 7 ; S y n . F u n g . 103. 1801.
= Pachyrhytisma
v . H o h n . , A n n . M y c . 15: 3 1 7 . 1 9 1 7 .
Ascocarps embedded i n a round or elliptical s t r o m a ; stroma dark outside and
w h i t e w i t h i n ; ascocarps o p e n i n g l a c e r a t e l y o r b y elongated s t r a i g h t or c u r v e d
s l i t s ; o p e n i n g m e c h a n i s m p r e s e n t or a b s e n t . A s c i c l a v a t e , s o m e t i m e s s l i g h t l y
s t a l k e d ; ascospores fascicled, c l a v a t e t o f i l i f o r m - c l a v a t e , o r s l i g h t l y f u s i f o r m ,
gelatinous s h e a t h w h e n present n o t w e l l d e v e l o p e d ; p a r a p h y s e s f i l i f o r m ,
sometimes s w o l l e n o r b e n t a t the t i p .
T Y P E - S P E C I E S : Rhytisma
acerinum
(Pers. ex S a i n t A m a n s ) F r .
1. Rhytisma acerinum (Pers. ex S a i n t A m a n s ) F r . , S y s t . M y c . 2: 569. 1823.
= Xyloma acerinum P e r s . , T e n t . D i s p . F u n g . 6. 1 7 9 7 ; S y n . F u n g . 104.
1801.
= Melanosorus
acerinus (Pers. ex S a i n t A m a n s ) D e N o t . , G i o r n . B o t .
I t a l . 2 ( 2 ) : (50). 1847.
P Y C N I D I A L S T A T E : Melasmia
acerina
L e v . , A n n . S c . N a t . I I I . 5: 276.
1846.
T h e s y n o n y m y of this species is m u c h m o r e i n v o l v e d t h a n i n d i c a t e d a n d n o
a t t e m p t has b e e n m a d e to i n v e s t i g a t e i t t h o r o u g h l y .
XXI.
P L A C U N T I U M E h r e n b . ex v . H o h n . , A n n . M y c . 15: 317. 1917.
= Placuntium
Ehrenb.,
Sylv. M y c o l . Berol. 17.1818.
F e r t i l e a s c o m a t a e p i p h y l l o u s o r o n y o u n g stems, s u b c u t i c u l a r , e m b e d d e d i n a
h i g h l y d e v e l o p e d s t r o m a , s h i n i n g b l a c k e x t e r n a l l y , o p e n i n g t o expose the
h y m e n i a l surface, c o m m o n l y w i t h a single buff-colored h y m e n i a l surface
s u r r o u n d i n g a c e n t r a l sterile area o r w i t h s e v e r a l sterile areas i n l a r g e r a s c o m a t a .
A s c i c l a v a t e , l a r g e r t h a n i n Rhytisma,
u p to 1 8 8 - 2 0 0 X 2 5 - 2 8 p.; ascospores
b r o a d l y c l a v a t e , often b a s a l l y m o r e o r less filiform, s o m e t i m e s s l i g h t l y greenishy e l l o w , u p to 5 5 - 9 0 X 5-8 p, w i t h a c o n s p i c u o u s g e l a t i n o u s s h e a t h .
T Y P E - S P E C I E S : Placuntium
andromedae
(Pers. ex F r . ) v . H o h n .
1. Placuntium
andromedae (Pers. ex F r . ) v . H o h n . , A n n . M y c . 15: 317. 1917.
= Xyloma andromedae P e r s . , S y n . F u n g . 1 0 4 . 1 8 0 1 .
= Placuntium
andromedae (Pers.) E h r e n b . , S y l v . M y c o l . B e r o l . 1 7 . 1 8 1 8 .
s^Rhytisma
andromedae (Pers.) ex F r . , S y s t . M y c . 2 : 567. 1823.
P Y C N I D I A L S T A T E : A p y c n i d i a l state, a p p a r e n t l y u n d e s c r i b e d , w a s f o u n d o n
specimens c o l l e c t e d i n e a r l y J u l y , 1925, L a k e T i m a g a m i , N i p i s s i n g D i s t r i c t ,
O n t a r i o . I n t h i s g a t h e r i n g , D A O M 83512, the p y c n i d i a a r e e p i p h y l l o u s , d a r k
b r o w n to b l a c k , o c c a s i o n a l l y f o r m i n g a m o r e o r less c o n t i n u o u s z o n e b u t m o r e
f r e q u e n t l y a b r o k e n series s u r r o u n d i n g the ascocarps of P. andromedae
at a
d i s t a n c e a b o u t e q u a l to the w i d t h of the p y c n i d i a t h e m s e l v e s . P y c n i d i a l s t r o m a t a u p to 1.0-1.2 m m . i n w i d t h a n d often w i t h s e v e r a l p y c n i d i a l c a v i t i e s i n the
same s t r o m a , the c a v i t i e s s e p a r a t e d b y s t r o m a t i c c o l u m n s ; s t r o m a t a w i t h d a r k
c o v e r i n g l a y e r 2 - 3 cells t h i c k , u p to 4 - 6 p i n t h i c k n e s s ; s p e r m a t i o p h o r e s a r i s i n g
in v e r t i c a l r o w s f r o m a h o r i z o n t a l , often d a r k e n e d l a y e r of e l o n g a t e d t h i c k
w a l l e d s c l e r e n c h y m a tissue 4 - 5 cells t h i c k ; s p e r m a t i o p h o r e s 3 - 4 cells i n l e n g t h ,
1438
CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
1 2 - 1 6 X 2 - 4 fi, t e r m i n a l c e l l o f t e n m o r e or less elongated a n d a t t e n u a t e d
t e r m i n a l l y a n d b e a r i n g one o r t w o o r p e r h a p s m o r e 3 - 5 X 1 M s p e r m a t i a ;
b a s a l l y the s p e r m a t i o p h o r e s t e n d to d a r k e n , a n d together w i t h the b a s a l
s c l e r e n c h y n a whose c e l l c o n t e n t s t e n d to b e c o m e b r o w n i s h , g i v e the a p p e a r a n c e
of a d a r k b a s a l l a y e r i n t h i c k sections.
L i t t l e is k n o w n of t h e d e v e l o p m e n t of the p y c n i d i a a n d t h e i r s u b s e q u e n t fate
a l t h o u g h some m a t u r e p y c n i d i a were s t i l l present o n the s a m e leaves w i t h
m a t u r e ascocarps. I t is n o t k n o w n , therefore, w h e t h e r these p y c n i d i a m i g h t
l a t e r h a v e been c o n v e r t e d i n t o ascocarps as s o m e t i m e s h a p p e n s i n Rhytisma
(Jones, 1925). T h e c o v e r e d p y c n i d i a present i n t h i s m a t e r i a l a p p r o a c h r a t h e r
m o r e c l o s e l y Leptostroma berberidis ( T h i i m . & W i n t . ) N a n n f . , the p y c n i d i a l s t a t e
of Lophodermium
berberidis (Schleich.) R e h m as d e s c r i b e d b y N a n n f e l d t , t h a n
the genus Melasmia o r the s i m p l e Leptostroma t y p e s s u c h asZ,. pinastri.
XXII.
N Y M A N O M Y C E S P . H e n n . , M o n s u n i a 1: 2 8 . 1 8 9 9 .
T h e five genera of the Nymanomyces
c o m p l e x h a v e a l l a t some t i m e been
r e g a r d e d as h a v i n g affinities to Rhytisma
f r o m w h i c h t h e y differ t h r o u g h the
possession of r e l a t i v e l y s h o r t , e l l i p t i c a l ascospores w h i c h are s a i d to be h y a l i n e
a n d 2-celled i n Synglonium
P e n z . a n d S a c c . a n d 1-celled, b r o w n i s h to b r o w n i n
t h e o t h e r genera. E x c e p t i n the genus Nymanomyces
no i m p e r f e c t s t a t e is
k n o w n . I n Nymanomyces
aceris-laurini
( P a t . ) P . H e n n . , h o w e v e r , T e r r i e r (1942)
o b s e r v e d some s t r u c t u r e s w h i c h m i g h t c o n c e i v a b l y represent a n i m p e r f e c t state,
b u t c o u l d n o t , he b e l i e v e d , b e l o n g i n the genus Melasmia L e v .
C e r t a i n c o n f u s i o n has a r i s e n p a r t l y f r o m the fact t h a t Synglonium
Penz. and
S a c c , the first genus n a m e d , w a s d e s c r i b e d t w i c e b y the s a m e a u t h o r s , o r i g i n a l l y
i n 1897 a n d l a t e r o n 1904, b u t the earlier d e s c r i p t i o n has been l a r g e l y o v e r l o o k e d
or i g n o r e d b y o t h e r a u t h o r s . T h i s a n d l a t e r d i s r e g a r d for the p r i n c i p l e of p r i o r i t y
has r e s u l t e d i n a n i n v o l v e d s y n o n y m y for s u c h a s m a l l g r o u p of species.
Nymanomyces
P . H e n n . , the second genus n a m e d , w a s based o n a s p e c i m e n
c o l l e c t e d b y N y m a n o n the s a m e h o s t as the t y p e of Synglonium.
A later
h o m o n y m of the t y p e species of Nymanomyces
was based o n a M a s s a r t s p e c i m e n
n a m e d b y P a t o u i l l a r d a n d r a i s e d to specific r a n k i n a p u b l i c a t i o n b y R a c i b o r s k i .
T h e t h i r d genus, Phaeorhytisma
P . H e n n . , w a s d e s c r i b e d i n the s a m e p a p e r as
Nymanomyces
b u t o n the f o l l o w i n g page. I n the s e c o n d p a r t of the s a m e p a p e r ,
the t y p e a n d o n l y species of Phaeorhytisma
w a s transferred to Criella w h i c h u p
to t h i s t i m e h a d e x i s t e d o n l y as a subgenus of Rhytisma.
F i n a l l y Saccardo,
himself, raised Rhytisma
F r . s u b g . Criella S a c c . to generic r a n k as
Criella
(Sacc.) S a c c . w i t h the t y p e species of the subgenus as the t y p e species of t h e
genus t o w h i c h he a d d e d three o t h e r species i n c l u d i n g Criella lonicerae ( P . H e n n .
a n d N y m . ) P . H e n n . a n d N y m . w h i c h h a d been n a m e d three years p r e v i o u s l y .
W h e t h e r H e n n i n g s a n d N y m a n c r e a t e d the genus i n t e n t i o n a l l y o r i n a d v e r t e n t l y
is u n i m p o r t a n t since the genera Nymanomyces
a n d Phaeorhytisma
both have
p r i o r i t y o v e r Criella. O n the o t h e r h a n d t h e s t a t u s of Synglonium
requires
f u r t h e r s t u d y . V o n H o h n e l (1909) c o n s i d e r e d the t y p e of Synglonium
insigne
P e n z . a n d S a c c . t o be a d i s o r g a n i z e d s p e c i m e n of Criella aceris-laurini
(Pat.)
S a c c . a n d S y d . a n d l a t e r (1918) referred to i t as a n o l d , i n f e c t e d s p e c i m e n of
Nymanomyces.
I n a s t u d y of the species w h i c h S a c c a r d o t r e a t e d u n d e r Criella
(Sacc.) S a c c .
D A R K E R : REVISION O F T H E H Y P O D E R M A T A C E A E
1439
von H o h n e l (1917a) w a s u n a b l e to s t a t e w i t h c e r t a i n t y w h a t the t y p e species,
C austro-caledonica ( C r i 6 ) S a c c , r e a l l y represented. I n the s a m e p a p e r a n d i n
another (v. H o h n . , 1917e) he d i s p o s e d of the o t h e r t h r e e species of Criella as
follows: C. aceris-laurini
(Pat.) Sacc. a n d S y d . was transferred to
Nymanomyces
w h i c h he m a i n t a i n e d as a n i n t r a e p i d e r m a l genus of t h e D e r m o p e l t i n a c e a e i n the
P h a c i d i a l e s sensu v . H o h n . ; C. lonicerae ( P . H e n n . a n d N y m . ) P . H e n n . a n d
N y m . was r e t u r n e d t o Phaeorhytisma
P . H e n n . w h i c h has p r i o r i t y o v e r Criella
and m u s t be r e t a i n e d , a n d Phaeorhytisma
w a s r e m o v e d t o the D e r m a t e a c e a e
near Pseudopeziza a n d Fabraea; a n d C. rhododendri ( R a c . ) S a c c . a n d S y d . w a s
made the t y p e species of a n e w genus, Pseudotrochila
v. H o h n . , somewhat
resembling Trochila F r . T h i s n e w genus was also p l a c e d i n the D e r m o p e l t i n aceae.
A s u m m a r y o f the five genera a n d t h e i r species is g i v e n b e l o w as a n a i d t o
anyone w i s h i n g to s t u d y the c o m p l e x f u r t h e r . S t u d e n t s of the g r o u p are i n
agreement t h a t a t least one genus of H y p o d e r m a t a c e a e is p r e s e n t here a n d t h a t
it s h o u l d p r o b a b l y be c a l l e d Nymanomyces
a l t h o u g h f u r t h e r e l u c i d a t i o n of the
status of the w h o l e g r o u p is needed, b u t e s p e c i a l l y of the species
Synglonium
insigne a n d Criella
austro-caledonica.
1. Synglonium
P e n z . & S a c c , M a l p i g h i a 11: 5 2 6 . 1 8 9 7 .
T Y P E - S P E C I E S : Synglonium
Javan. 63.1904.
2. Nymanomyces
insigne
P e n z a n d S a c c , I.e. 5 2 7 ; Icones F u n g .
P . H e n n . , M o n s u n i a 1: 2 8 . 1 8 9 9 .
T Y P E - S P E C I E S : Nymanomyces
aceris-laurini
P . H e n n . , I.e. 28, 170. 1899
(based o n c o l l e c t i o n of N y m a n ) .
= Rhytisma
acerinum
F r . f. aceris-laurini
Pat., A n n . Jard. Buitenzorg
S u p p l . 1: 1 2 1 . 1 8 9 7 (based o n c o l l e c t i o n of M a s s a r t ) .
= Nymanomyces
aceris-laurini
(Pat.) P . H e n n . in R a c i b . , Parasit. A l g e n
u . P i l z e J a v a ' s 2: 2 3 . 1 9 0 0 .
= Rhytisma
(Criella) aceris-laurini
(Pat.) P . H e n n . , in R a c i b . , H e d w .
B e i b l . 3 9 : (111). 1900.
= Criella aceris-laurini
( P a t . ) S a c c . a n d S y d . , S y l l . F u n g . 16: 7 8 6 . 1 9 0 2 .
3. Phaeorhytisma
P . H e n n . , M o n s u n i a 1: 2 9 . 1 8 9 9 .
T Y P E - S P E C I E S : Phaeorhytisma
lonicerae P . H e n n . a n d N y m . , i n P . H e n n . ,
M o n s u n i a 1: 2 9 . 1 8 9 9 .
= Criella lonicerae ( P . H e n n . a n d N y m . ) P . H e n n . a n d N y m . , i n P . H e n n . ,
M o n s u n i a 1: 1 7 1 . 1 8 9 9 .
4. Criella (Sacc.) S a c c , i n S a c c . a n d S y d . , S y l l . F u n g . 16: 7 8 6 . 1 9 0 2 .
=Rhytisma
F r . , s u b g . Criella S a c c , S y l l . F u n g . 8: 7 5 6 . 1 8 8 9 .
T Y P E - S P E C I E S : Criella austro-caledonica
( C r i e ) S a c c . a n d S y d . S y l l . F u n g . 16:
786.1902.
^•Rhytisma
austro-caledonicum
Crie, B u l l . Soc. L i n n . Normandie, Caen,
8: 4 4 6 . 1 8 7 4 .
=Rhytisma
(Criella) austro-caledonicum
C r i e , i n S a c c , S y l l . F u n g 8: 756.
1889.
1440
5. Pseudotrochila
CANADIAN JOURNAL OF BOTANY. VOL. 45, 1967
v, H o h n . , B e r . Deutsch. B o t . Ges. 35: 419.1917.
T Y P E - S P E C I E S : Pseudotrochila rhododendri ( R a c i b . ) v . H o h n . , I.e. 422.
= Cryptomyces
(Criella)
rhododendri
Racib., Parasit. Algen u. Pilze
J a v a ' s 3: 1 8 . 1 9 0 0 .
== Criella rhododendri ( R a c i b . ) S a c c . a n d S y d . , S y l l . F u n g . 16: 7 8 7 . 1 9 0 2 .
Discussion
T h e t a x o n o m i c changes p r o p o s e d a b o v e i n v o l v e p r i n c i p a l l y species f o r m e r l y
assigned to the genera Hypoderma, Lophodermium,
Hypodermella,
a n d Bifusella.
T h e r e v i s i o n s h a v e been for the m o s t p a r t a m o n g the m o r e p a r a s i t i c species,
w h e r e , as one m i g h t expect, d i s t i n c t i v e c h a r a c t e r s are m o r e s h a r p l y d e l i n e a t e d .
I n the o l d e r classifications w h i c h were e x t r e m e l y a r t i f i c i a l m o s t of the genera
were f o u n d e d p r i m a r i l y o n spore shape. I n the present scheme, c h a r a c t e r s of the
ascocarps, a s c i , a n d spores as w e l l as p y c n i d i a were m o r e or less s y n t h e s i z e d to
a r r i v e a t m o r e m e a n i n g f u l r e l a t i o n s h i p s . A l t h o u g h no m o d i f i c a t i o n s o r a d d i t i o n s
h a v e been i n c o r p o r a t e d i n t o t h i s w o r k as the r e s u l t of recent c y t o l o g i c studies b y
G o r d o n (1966), i t is possible t h a t e v e n t u a l l y c e r t a i n c o m p r o m i s e s m a y h a v e to
be m a d e , a n d f u r t h e r w o r k b y G o r d o n m a y r e s u l t i n r e e v a l u a t i o n of c e r t a i n
proposals m a d e here. I n the m a i n , h o w e v e r , D r . G o r d o n ' s recent findings seem
to c o m p l e m e n t those of the w r i t e r w i t h a few m i n o r e x c e p t i o n s .
I n the present w o r k i t is clear, t h a t b y segregating the o b v i o u s l y c l o s e l y
r e l a t e d species of the " n e r v i s e q u i a " o r Lirula
c o m p l e x f r o m o t h e r coniferi n h a b i t i n g species, a d i s t i n c t i v e g r o u p i n g has been c r e a t e d . I n t y p i c a l species of
t h i s c o m p l e x o n Abies the e p i p h y l l o u s p y c n i d i a d e v e l o p after one y e a r a n d a r e
m o r e or less c o n t i n u o u s a l o n g the s u l c u s o r i n t w o lines, one a l o n g each w i n g of
t h e needle. T h e h y p o p h y l l o u s ascocarps f o l l o w the " n e r v e " of the needle a n d
m a t u r e a t the e n d of t w o y e a r s . T h e e r e c t i o n of three g e n e r a for the n e r v i s e q u i o u s species, i.e., Virgella, Isthmiella, a n d Lirula, b r i n g s t h e m o n a s e c o n d a r y
basis i n t o l i n e w i t h the o l d S a c c a r d o a n s y s t e m based o n spore f o r m . T h e rem o v a l of the n e r v i s e q u i o u s species f r o m Hypodermella,
Bifusella, a n d Hypoderma, w h e r e t h e y h a v e a l w a y s a p p e a r e d to be d i s c o r d a n t elements, simplifies the
f u r t h e r t a x o n o m i c t r e a t m e n t of the species left i n the c o m p l e x e s s u r r o u n d i n g
these genera.
T h e Hypodermella c o m p l e x is t h e n l i m i t e d i n general to species w i t h r e l a t i v e l y
s h o r t ascocarps a n d s c a t t e r e d p y c n i d i a w h i c h precede the a p p e a r a n c e of the
ascocarps b y o n l y a few m o n t h s r a t h e r t h a n a f u l l y e a r as i n the n e r v i s e q u i o u s
c o m p l e x . A f t e r r e m o v a l of Lirula
the r e m a i n i n g species o f t h e o l d genus
Hypodermella
f a l l i n t o three v e r y d i s t i n c t g r o u p s w h i c h m a y be segregated i n t o
genera as f o l l o w s : Hypodermella
( t y p e : H. laricis) w i t h s u b c u t i c u l a r ascocarps
and n u m e r o u s s m a l l , d a r k p y c n i d i a , Davisomycella
( t y p e : D. ampla)
with
p r o m i n e n t s u b e p i d e r m a l ascocarps a n d p y c n i d i a w h i c h are i n c o n s p i c u o u s o r
u n k n o w n , a n d Lophodermella
v . H o h n . ( t y p e : L. sulcigena) w i t h m o r e or less
c o n c o l o r o u s , s u b h y p o d e r m a l ascocarps. W i t h o b v i o u s affinities to these c l a v a t e s p o r e d genera are Ploioderma, segregated f r o m Hypoderma, w i t h s h o r t r o d - l i k e
ascospores, a n d p o s s i b l y also Elytroderma w i t h l a r g e 2-celled ascospores.
T h e Bifusella c o m p l e x w i t h three genera is a r a t h e r heterogeneous g r o u p . T h e
transfer of the n e r v i s e q u i o u s species to Isthmiella a n d the r e m o v a l of the m o r e
or less a b e r r a n t 73. striiformis w o u l d h a v e left the t y p e species, 73. linearis, as the
DARKER: REVISION OF T H E HYPODERMATACEAE
1441
sole c o n i f e r - i n h a b i t i n g r e p r e s e n t a t i v e of the genus. T o Bifusella, h o w e v e r , h a v e
been a d d e d t w o f o r m e r Hypoderma
species o c c u r r i n g also o n soft pines. A l though the spores of these t w o species m i g h t best be d e s c r i b e d as r o d - l i k e , t h e y
have a d i s t i n c t t e n d e n c y t o w a r d s a b i f u s i f o r m c o n d i t i o n a n d i n b o t h the
paraphyses t e n d t o d i s a p p e a r a t m a t u r i t y as i n the t y p e species, B.
linearis.
L i k e the t y p e species, t h e y are also s u b c u t i c u l a r , a n d h a v e l a r g e p y c n i d i a w i t h a
d i s t i n c t c o v e r i n g tissue a b o v e t h e c u s h i o n of s p e r m a t i o p h o r e s a n d r e l a t i v e l y
large s p e r m a t i a . I n Duplicaria
the ascocarps are a l s o s u b c u t i c u l a r b u t t h e
p y c n i d i a are s m a l l a n d i n c o n s p i c u o u s . Soleella is a n a b e r r a n t m o n o s p e c i f i c genus
w i t h a b u n d a n t s u b e p i d e r m a l ascocarps b u t l a c k i n g p y c n i d i a .
T h e Hypoderma
c o m p l e x w i t h t w o of the o l d e r genera, Hypoderma
and
Lophodermium,
c o n t a i n s b y far the largest n u m b e r of species. B o t h genera are i n
need of c a r e f u l m o n o g r a p h i c t r e a t m e n t as t h e y h a v e b e c o m e d e p o s i t o r i e s for
miscellanies of species. Lophodermium,
t h o u g h v a l i d l y p u b l i s h e d , is i l l e g i t i m a t e
a n d requires c o n s e r v a t i o n as p r o p o s e d a b o v e . V o n H o h n e l ' s a t t e m p t to set u p
new genera i n t h i s c o m p l e x has n e v e r m e t w i t h g e n e r a l a p p r o v a l . I t is q u i t e
possible t h a t his genera Lophodermina
a n d Lophodermellina
c o u l d be redefined
from t h e i r t y p e species o n a m o r e a c c e p t a b l e basis t o i n c l u d e a few species b u t
n o t for a g e n e r a l transfer of species as w a s d o n e b y T e h o n . T h e i n v a l i d l y
p u b l i s h e d genus Dermascia T e h o n c a n n o t be used w i t h o u t v a l i d a t i o n a n d
subsequent r e d e f i n i t i o n a n d s h o u l d p e r h a p s be d r o p p e d f r o m the l i t e r a t u r e . A s
a c c e p t e d a b o v e Hypoderma a n d Lophodermium
h a v e n o t been r e e v a l u a t e d t o a n y
m a r k e d degree except t o r e m o v e Hypoderma
desmazierii
to a n e w genus,
Meloderma,
nearer Lophodermium,
a n d to a c c e p t the transfer of c e r t a i n
p h r a g m o s p o r o u s Lophodermium
species t o Lophomerum
Ouellette and M a g a s i .
T h e genus Coccomyces is a p p a r e n t l y a m o r e o r less m i x e d c o m p l e x of species
w i t h i n itself. O n the one h a n d are species l i k e C. coronatus w h i c h tear o p e n i n a
lacerate, p h a c i d i a c e o u s m a n n e r a t m a t u r i t y w h i l e o t h e r s o p e n r a d i a t e l y b y
means of o r g a n i z e d o p e n i n g m e c h a n i s m s .
Colpoma, together w i t h Xyloschizon
a n d Bifusepta,
f o r m a s m a l l c o m p l e x of
species i n w h i c h t h e b a s a l s t r o m a t i c tissue is v e r y s t r o n g l y d e v e l o p e d . T h e i r
occurrence o n dead stems a n d branches a n d even on dead w o o d p r o b a b l y
a c c o u n t s for the fact t h a t the s t r o m a i n c e r t a i n species has a t e n d e n c y to
separate l a t e r a l l y f r o m the h o s t tissue a n d t o b e c o m e s o m e w h a t e r u m p e n t . I n
c e r t a i n species, n o t a b l y Colpoma crispum, the ascocarps m a y be e m b e d d e d i n
the b a r k b u t i n the s a m e s p e c i m e n m a y pass o v e r t o b a r e w o o d w h e r e t h e y
o r i g i n a t e v e r y near the surface of the w o o d a n d a p p e a r a l m o s t c o m p l e t e l y
e r u m p e n t . V o n H o h n e l has suggested t h e transfer of t h i s a n d o t h e r species to a
n e w genus Pragmoparopsis
w h i c h he w o u l d place i n the T r y b l i d i a c e a e . Colpoma
quercinum o n o a k is w e l l k n o w n b u t there are a n u m b e r of species o n conifers
w h i c h d o n o t seem to differ m u c h f r o m one a n o t h e r o r f r o m C. crispum.
Xyloschizon S y d . appears t o be d i s t i n c t i n t h a t i t possesses large, p h r a g m o s p o r o u s
ascospores a l t h o u g h u n f o r t u n a t e l y the t y p e species has n e v e r been l o c a t e d b y
t h e w r i t e r . T h e second species, X. atratum, a p p e a r s t o be m o r e closely r e l a t e d t o
C. crispum. Bifusepta,
w i t h a f a i r l y t h i c k b a s a l s t r o m a t i c l a y e r , has s o m e
affinities w i t h Colpoma. Its b i f u s i f o r m , s e p t a t e spores r e s e m b l e s o m e w h a t those
of Bifusella b u t the s t r u c t u r e of the f r u i t i n g b o d y is m o r e c l o s e l y r e l a t e d to t h a t
of C. quercinum.
T h e Rhytisma
c o m p l e x c o m p r i s e s three g e n e r a : Rhytisma,
Placuntium,
and
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CANAPI
A N
JOURNAL OF BOTANY. VOL. 45, 1967
Nymanomyces.
These genera a r e c h a r a c t e r i z e d b y m o r e h i g h l y d e v e l o p e d
s t r o m a t a w h i c h are m o r e o r less s c l e r o t i c , e x t e r n a l l y d a r k , a n d i n t e r n a l l y l i g h t ,
w i t h one t o several ascocarps i n e a c h s t r o m a . T h e ascocarps o p e n b y t h e
r u p t u r i n g of the c o v e r i n g l a y e r w i t h o r w i t h o u t a n o p e n i n g m e c h a n i s m . T h e
ascospores are a c i c u l a r - c l a v u l a t e i n Rhytisma,
quite robust a n d clavate i n
Placuntium,
and ovoid and brown in
Nymanomyces.
The
assignment of the Phacidiales to a position between certain of the
P y r e n o m y c e t e s a n d t h e t r u e D i s c o m y c e t e s as p r o p o s e d b y G a u m a n n (1964)
s t i l l seems t o be t h e m o s t a c c e p t a b l e d i s p o s i t i o n o f this order. T h i s leaves t h e
Hypodermataceae i n t h e same relative position occupied earlier b y the H y s t e r i aceae (treated as a n order) o f R e h m (1887) o r t h e H y s t e r i i n e a e (treated also as
a n order) of L i n d a u (1896). D u r i n g t h e l o n g i n t e r v a l b e t w e e n t h e classifications
of R e h m a n d L i n d a u a n d those of m o r e recent w o r k e r s t h e H y p o d e r m a t a c e a e
h a v e been v a r i o u s l y assigned, n o t a b l y b y v o n H o h n e l , N a n n f e l d t , T e r r i e r , a n d
others, i n o r n e a r t h e H e l o t i a l e s i n t h e D i s c o m y c e t e s . W i t h G a u m a n n ' s treatment the cycle is complete a n d the Hypodermataceae, n o w separated from the
H y s t e r i a c e a e , a r e b a c k v e r y close t o t h e p o s i t i o n w h i c h t h e y o c c u p i e d s e v e n t y
t o e i g h t y y e a r s ago.
W h i l e c l a r i f y i n g some of t h e p r o b l e m s e n c o u n t e r e d i n t h e H y p o d e r m a t a c e a e ,
the present w o r k has m e r e l y t o u c h e d u p o n c e r t a i n o f t h e genera a n d references
t o t h e species of these genera h a v e been o m i t t e d i n large p a r t . S e v e r a l of t h e
l a r g e r genera are s t i l l i n need of m o n o g r a p h i c t r e a t m e n t . I t is h o p e d t h a t t h i s
r e d e f i n i t i o n o f genera m a y s t i m u l a t e f u r t h e r research e s p e c i a l l y i n the areas
least stressed here.
Acknowledgments
I t h a n k e s p e c i a l l y D r . J . W a l t o n G r o v e s w h o h a s followed t h e d e v e l o p m e n t o f
t h i s s t u d y w i t h i n t e r e s t a n d h a s k i n d l y r e v i e w e d t h e m a n u s c r i p t . I a m also
d e e p l y i n d e b t e d t o D r . L u e l l a K . W e r e s u b f o r a d v i c e o n some of t h e m o r e
k n o t t y p r o b l e m s of n o m e n c l a t u r e . M i s s D o r o t h y H a l l has s m o o t h e d m a n y
difficulties b y l o c a t i n g o l d , rare p u b l i c a t i o n s needed t o c o m p l e t e t h e w o r k .
Finally, I thank D r . I. M a c k e n z i e L a m b , Curator, a n d M i s s Ursula V . Metzger
of t h e F a r l o w H e r b a r i u m , C a m b r i d g e , M a s s a c h u s e t t s , a n d D r . C . R . B e n j a m i n ,
C u r a t o r o f t h e N a t i o n a l F u n g u s C o l l e c t i o n s , B e l t s v i l l e , M a r y l a n d , for recent
assistance, a n d t h e c u r a t o r s o f m a n y E u r o p e a n h e r b a r i a f o r t h e p r i v i l e g e s
a c c o r d e d t o m e a n u m b e r of years a g o . A l l o f these a n d m a n y others h a v e
contributed i n various ways to m a k e the s t u d y more complete.
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