Studies on Puccinia polysora Underw. I. The world distribution of forms of Puccinia polysora


Cammack, R.H.

Transactions of the British Mycological Society 41(1): 89-94

1958


An examination of herbarium material of Puccinia polysora from all areas of the world revealed two main size groups of uredospores. The smaller group was encountered in South East Asia and neighbouring islands with the exception of Borneo, and the larger group in the West Indies, Africa and the South Indian Ocean. The two main groups are consistent with the two directional spread, westwards and eastwards, from the origin of the rust, the Caribbean area. As data on cross-inoculation studies are lacking it is uncertain if distinct forms of P. polysora exist.

[
89
]
Trans.
Brit.
mycol.
Soc.
41
(1),
8
9
-
94
(1958).
STUDIES
ON
PUCCINIA
POLYSORA
UNDERW.
I.
THE
WORLD
DISTRIBUTION
OF
FORMS
OF
P.
POLYSORA
By
R.
H.
CAMMACK
West
African
Maize
Research
Unit,
Ibadan,
Nigeria
(With
2
Text
-figures)
An
examination
of
herbarium
material
of
Puccinia
polysora
from
all
areas
of
the
world
revealed
two
main
size
groups
of
uredospores.
The
smaller
group
was
encountered
in
South
East
Asia
and
neighbouring
islands
with
the
exception
of
Borneo,
and
the
larger
group
in
the
West
Indies,
Africa
and
the
South
Indian
Ocean.
The
two
main
groups
are
consistent
with
the
two
directional
spread,
westwards
and
eastwards,
from
the
origin
of
the
rust,
the
Caribbean
area.
As
data
on
cross
-inoculation
studies
are
lacking
it
is
uncertain
if
distinct
forms
of
P.
polysora
exist.
Until
the
appearance
of
maize
rust
caused
by
Puccinia
polysora
Underw.
in
West
Africa
in
1940,
knowledge
of
the
world
distribution
of
this
rust
was
confined
to
reports
from
Central
and
South
America
and
the
West
Indies.
The
fi
rst
record
was
established
by
Underwood
(1897),
who
described
it
as
new
from
Tripsacum
laxum
in
Alabama.
Arthur
(192o)
recorded
it
on
T.
laxum
in
Florida,
New
Jersey
and
San
Domingo
and
on
other
species
on
Tripsacum
in
Mexico
and
Cuba.
Cummins
(1941)
identified
it
on
Zea
mays
from
Peru
and
later
found
it
to
be
of
general
occurrence
on
maize
in
Central
and
South
America.
Other
records
are
given
by
Wood
&
Lipscomb
(1956)
.
THE
WORLD
SPREAD
OF
PUCCINIA
POLYSORA
ON
ZEA
MAYS
Central
and
South
America,
West
Indies
In
1945
the
rust
was
observed
on
Zea
mays
in
Trinidad
and
in
1947
in
British
Honduras.
In
1948,
it
was
recorded
in
Jamaica.
Information
concerning
the
severity
of
the
disease
in
these
areas
is
almost
lacking,
but
it
was
observed
that
'
negligible
'
damage
was
caused
in
the
West
Indies.
South
-East
Asia
and
adjacent
islands
Specimens
of
maize
rust
were
collected
in
Malaya
in
195o
and
identified
as
P.
polysora.
In
1952
maize
in
British
North
Borneo
was
affected
and
in
1956
Siam,
the
Philippines
and
Christmas
Island
(Indian
Ocean).
Again
reports
of
damage
are
scanty
but,
in
some
of
the
affected
areas
the
attack
was
widespread
and
locally
severe.
9
0
Transactions
British
Mycological
Society
The
African
Continent
Deighton
(1951)
reported
that
maize
rust
during
the
1949
growing
season
was
widespread
and
severe
in
Sierra
Leone.
The
rust
was
at
fi
rst
thought
to
be
P.
sorghi,
previously
only
once
recorded
in
Sierra
Leone
(in
1943),
but
the
virulence
of
the
disease
aroused
suspicions,
and
a
critical
examination
was
made
by
G.
R.
Bisby,
of
Sierra
Leone
specimens
col-
lected
in
1949
and
of
collections
made•in
1951
from
Ghana
and
Nigeria,
as
a
result
of
which
he
concluded
that
they
belonged
to
P.
polysora,
and
G.
B.
Cummins
confirmed
these
identifications.
After
further
examinations
of
all
maize
rust
specimens
from
the
African
Continent,
it
was
determined
with
reasonable
certainty
that
P.
polysora
did
not
occur
in
Africa
before
1
949.
In
195o
the
Department
of
Agriculture,
Ghana,
reported
a
generally
distributed
outbreak
of
maize
rust
in
that
colony,
and
it
was
observed
that
the
highest
incidence
of
rust
was
in
the
coastal
rain
-forest
areas,
where
rainfall
and
humidity
are
greatest
and
was
less
severe
in
the
drier
areas
of
the
interior.
All
specimens
collected
in
Ghana
during
195o
were
identified
as
P.
polysora.
In
mid
-195o
the
Department
of
Agriculture
of
the
French
Territories
of
Ivory
Coast
and
Dahomey
reported
outbreaks
of
rust,
and
mentioned
that
rust
was
most
severe
on
maize
planted
late
in
the
season
Table
I.
The
world
distribution
of
Puccinia
polysora
(Dates
of
fi
rst
appearance
are
listed
in
chronological
order.)
Year
fi
rst
C.M.I.
Territory
reported
Herbarium
no.
Host
Alabama
1897
Det.
Underwood
Tripsacum
laxum
Florida
1920
Det.
Arthur
T.
laxum
T.
latifolium
Peru
1
94
1
Det.
Cummins
Zea
mays
Trinidad
1
945
1
944
Z.
map
Honduras
1
947
Z•
mays
Jamaica
1
94
8
Z.
mays
Sierra
Leone
Ivory
Coast
1
949
1950
Z.
mays
Z.
mays
Ghana
1950
44
33
4
858
7
19664
4
84
37
2
580
8
1
Z.
mays
Dahomey
1950
47944
Z.
mays
Nigeria
Malaya
1950
t950
6
4
2
59
16
2
4
6
Z.
mays
Z.
mays
French
Guinea
1
95
1
53
0
57
Z.
mays
Belgian
Congo
1
95
2
52565
Z.
mays
Sudan
1952
51198
Z.
mays
Kenya
1952
50121
Z.
mays
Tanganyika
1952
50581
Z.
mays
North
Borneo
1
95
2
62556
Z.
mays
Nyasaland
1
953
5
2
59
0
Z.
mays
Northern
Rhodesia
1953
5
2
4
8
9
Z.
mays
Southern
Rhodesia
1953
56016
Z.
mays
Portuguese
S.
Africa
1
953
Det.
Carvalho
Z.
mays
Madagascar
1
953
Z•
mays
Mauritius
1
953
51927
Z.
mays
Reunion
1953
52909
Z.
mays
Agalega
Island
1
955
60590
Z.
mays
Rodriguez
Island
1955
60203
Z.
map
Christmas
Island
(Indian
Ocean)
1956
62093
Z.
mays
Siam
1956
Det.
Cummins
Z.
mays
Philippines
1
95
6
Det.
Cummins
Z.
mays
Puccinia
polysora.
R.
H.
Cammack
91
and
that
early
plantings
escaped
to
some
extent.
In
late
195o
the
rust
appeared
in
the
south-west
corner
of
Nigeria
(Rhind,
Waterston
&
Deighton,
1952).
The
following
year
the
rust
was
widespread
throughout
western
and
eastern
Nigeria
and
had
appeared
in
Cameroons
under
British
Trusteeship.
Nattrass
(
1952)
reported
P.
polysora
in
Kenya
in
1952.
The
previous
year,
having
been
warned
of
the
advent
of
P.
polysora
in
West
Africa,
an
examination
of
maize
rust
in
the
colony
had
revealed
only
P.
sorghi.
Also
in
1952
it
was
reported
in
the
Belgian
Congo,
Tanganyika
and
the
Sudan.
Early
in
1953
the
rust
appeared
in
Nyasaland
and
Southern
Rhodesia,
and
later
the
same
year
in
Madagascar,
Mauritius
and
Reunion
in
the
South
Indian
Ocean.
In
1955
further
reports
were
received
from
the
Islands
of
Agalega
and
Rodriguez
in
the
Indian
Ocean.
The
present
known
world
distribution
of
P.
polysora
is
given
in
Table
(see
also
C.M.I.
distribution
map
237)
and
when
there
is
a
specimen
at
the
Commonwealth
Mycological
Institute
a
herbarium
reference
number
is
given.
VARIATION
IN
UREDOSPORE
SIZE
In
order
to
determine
if
uniformity
existed
in
the
dimensions
of
uredo-
spores
an
analysis
was
made
of
collections
from
different
areas
in
West
Africa.
Samples
of
a
hundred
spores
from
herbarium
specimens
from
each
of
the
following
locations
were
measured
:
Territory
No.
of
samples
Territory
No.
of
samples
Sierra
Leone
3
Nigeria
1
4
Ivory
Coast
2
Cameroons
under
3
Ghana
11
British
Trusteeship
Dahomey
2
Frequency
distribution
curves
were
constructed
from
50o
measurements
of
lengths
and
breadths
of
uredospores
collected
at
Ibadan,
Nigeria,
during
1953,
and
all
other
West
African
samples
listed
above
were
subsequently
compared
with
these
curves,
and
it
was
found
that
none
departed
signifi-
cantly
from
the
size
distribution
of
the
Ibadan
sample.
An
analysis
of
uredospore
measurements
of
West
African
specimens
is
given
in
Table
2.
In
1956
F.
C.
Deighton
sent
the
writer
specimens
of
maize
rust
from
South
East
Asia
and
adjacent
islands,
and
mentioned
that
uredospores
from
that
area
were
apparently
smaller
than
in
collections
from
Africa.
Six
samples,
each
of
too
spores,
were
measured
from
Malayan
material,
Table
2.
An
analysis
of
the
measurements
of
uredospores
of
P.
polysora
from
collections
made
in
Vliest
Africa
and
the
Eastern
Pacific
West
Africa
Malaya,
Christmas
Is.
Mean
length
(µ)
32.70±0.14
28.67±0.23
Mean
breadth
(p.)
2
4'79
±
0'24
22.59
4-
o.31
Correlation
coefficient
of
—o.28
length
to
breadth
—0.31
Relative
volume
of
spores
1.0
0.78
(Smith,
1953)
Mean
difference
of
lengths
(p.)
4.03
±
0.38
Mean
difference
of
breadths
(µ)
2.20
±
0'24
92
Transactions
British
Mycological
Society
and
two
from
Christmas
Island
(Indian
Ocean).
The
analysis
of
these
samples
is
also
given
in
Table
2,
and
the
results
show
a
highly
significant
difference
(P
=
0.
I
°A
)
)
between
West
African,
and
Malayan
and
Christmas
Islands
(Indian
Ocean)
spores
both
in
lengths
and
breadths.
With
the
assistance
of
the
Commonwealth
Mycological
Institute
and
the
Arthur
Herbarium,
duplicates
of
all
available
specimens
of
P.
polysora
on
Zea
mays
and
on
the
alternate
hosts
Tripsacum
laxum
and
Euchlaena
mexicana
were
obtained.
Samples
of
loo
spores
from
a
total
of
sixty-five
specimens,
representative
of
all
areas
of
the
world
affected
by
the
rust,
were
measured.
28
27
26
25
E
24
23
22
Malaya,
Philippines,
Christmas
Islands
A
.
Borneo
•A ••
d",,
0
.•
AA
.1
••••
••••
••
sire
4,
Africa,
West
Indies
U.S.A.,
Central
America
1
29 30
31
32
Mean
length
CO
33
34
35
Fig.
s.
The
distribution
of
mean
measurements
of
uredospores
of
Puccinia
polysora.
Host
key:
•,
Zea
mays;
A,
Tripsacum
laxum;
Euchlaena
mexicana.
Uredospores
from
herbarium
specimens
which
have
been
identified
as
P.
polysora
fall
on
a
size
basis
into
two
main
groups.
This
grouping
is
illustrated
in
Fig.
r
in
which
the
mean
size
of
samples
is
plotted.
Samples
obtained
from
different
hosts
are
indicated
by
different
symbols.
This
fi
gure
shows
that
there
are
two
main
forms
of
P.
polysora
to
be
found
on
Z.
mays
when
distinguished
on
a
size
basis.
The
smaller
of
these
is
centred
on
the
Pacific
area
and
is
consistent
with
the
eastward
spread
of
the
rust
from
its
fi
rst
recorded
appearance
in
South
-East
Asia,
i.e.
from
Malaya.
Presumably
the
origin
of
the
rust
was
once
more
America.
Records
from
Borneo,
however,
are
not
consistent
with
this,
the
spores
Puccinia
polysora.
R.
H.
Cammack
93
being
much
larger
(Figs.
i,
2).
The
larger
spore
form
is
found
in
the
Caribbean
area,
Africa
and
islands
in
the
South
Indian
Ocean
and
is
consistent
with
the
observed
eastward
spread
(Table
1).
Fig.
2
illustrates
the
variation
in
morphology
with
location.
The
histo-
grams
were
each
constructed
from
200
measurements
from
samples
of
rust
chosen
at
random
from
the
areas
mentioned.
The
mean
size
of
each
sample
is
marked
by
a
vertical
line.
Kuala
Lumpur,
Malaya
Arlington,
Virginia
Trinidad,
West
Indies
Ibadan,
Nigeria
-
2
z
U
Jessleton,
North
Borneo
—100
Length
Breadth
20
I
i
1
30
40
14
20
30
Class
values
p
A
-111111----imilL
ik_mal._
Fig.
2.
Class
-frequency
histograms
of
the
measurements
of
fi
ve
samples
of
uredospores
of
Puccinia
polysora.
DISCUSSION
Data
are
lacking
on
cross
-inoculation
studies
of
cultures
of
Puccinia
polysora
from
areas
outside
Africa
and
it
is
not
possible
at
present
to
determine
if
biotypes
exist.
Attempts
to
infect
Tripsacum
laxum
at
Ibadan
with
West
94
Transactions
British
Mycological
Society
African
uredospore
material
obtained
from
Zea
mays
have
failed.
The
Tripsacum
used
was
clonal
material
from
the
West
Indies
in
which
area
natural
infection
of
this
host
by
P.
polysora
occurs
readily.
However,
it
has
not
been
established
whether
the
forms
of
P.
polysora
on
Zea
and
Tripsacum
in
the
West
Indies
are
identical.
Further
investi-
gations
are
required
to
determine
this
point,
using
cross
-inoculation
and
host
material
of
known
genetic
identity.
Until
this
has
been
done
there
is
no
justification
for
further
assumptions
on
the
relationship
of
the
groupings
in
Fig.
1.
The
effect
of
environment
on
uredospore
formation
may
be
a
contri-
butory
factor
to
the
variations
in
spore
size
that
have
been
found.
Levine
(1923)
and
Stakman
&
Levine
(1919)
have
shown
that
environmental
factors
influence
uredospore
size
in
P.
graminis,
and
Bailey
(1925)
observed
that
environmental
conditions
result
in
changes
in
uredospore
size
as
great
as,
or
even
greater
than,
the
differences
between
spores
of
different
races
of
P.
g.
avenae
produced
under
similar
conditions.
Hingorani
(1952)
showed
that
significant
fl
uctuations
were
induced
in
uredospore
size
of
races
2,
7
and
8
of
P.
g.
avenae
by
temperature,
light
intensity
and
the
degree
of
resistance
of
the
host.
This
last
factor,
especially,
may
be
applicable
to
P.
polysora.
Levine
(1923)
and
Waterhouse
(1930),
and
Manners
(1950)
have
shown
uredospore
size
variation
in
different
physiologic
races
of
P.
graminis
and
P.
glumarum,
respectively.
A
pleiotropic
effect
of
this
type
is
not
unexpected
in
P.
polysora
on
theoretical
grounds.
REFERENCES
ARTHUR,
J.
C.
(1920).
N.
Amer.
Flora,
7,
279.
BAILEY,
D.
L.
{1925}.
Physiologic
specialisation
in
Puccinia
graminis
avenae
Erikss.
&
Henn.
Tech.
Bull.
Minn.
Agric.
Exp.
Sta.
35.
CUMMINS,
G.
B.
(1941).
Identity
and
distribution
of
three
rusts
of
corn.
Phytopathology,
31,
856-857.
DEIGHTON,
F.
C.
(1951).
Ann.
Rep.
Dep.
Agric.
S.
Leone
1949,
p.
15.
HINGORANI,
M.
K.
(1952).
Morphological
and
pathological
studies
of
races,
2,
7
and
8
of
oat
stem
rust.
Phytopathology,
42,
486-488.
LEVINE,
M.
N.
(1923).
A
statistical
study
of
the
comparative
morphology
of
biologic
forms
of
Puccinia
graminis.
J.
agric.
Res.
2
4,
539-567.
MANNERS,
J.
G.
(I
95o).
Studies
on
the
physiologic
specialisation
of
yellow
rust
(Puccinia
glumarum
(Schm.)
Erikss.
&
Henn.)
in
Great
Britain.
Ann.
appl.
Biol.
37,
187-214.
NATTRASS,
R.
M.
(1952).
Preliminary
notice
of
the
occurrence
in
Kenya
of
a
rust
Puccinia
polysora
on
maize.
E.
Afr.
agric.
3.
18,
39.
RHIND,
D.,
WATERSTON,
J.
M.
&
DEIGHTON,
F.
C.
(1952).
Occurrence
of
Puccinia
polysora
Underwood
in
West
Africa.
Nature,
Lond.,
169,
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