A comparative study of the floral morphology in the genus Disperis (Orchidaceae)


Kurzweil, H.; Linder, H.P.

Beitrage zur Biologie der Pflanzen 66(3): 433-477

1991


The floral morphology of selected species of the genus Disperis (Orchidaceae-Orchidoideae-Coryciinae) is surveyed. The study concentrates on the little known tropical species of the genus, supplementing previous studies on the South African species, which are structurally rather different from most tropical species. Minute teeth adpressed to the rostellum just above the lip insertion, interpreted as vestiges of the lateral carpels (lateral stigma lobes), were observed in many tropical and some South African species, and appear to be the basic condition in Disperis. The congenital fusion of the lip to the gynostemium, which is often quoted as being one of the main characters of the Coryciinae, is shown to be completely or almost completely absent in a few species. The lateral gynostemium appendages are distinctly bi-partite in many species, with a prominent, smooth, erect portion and a small, sculptured, horizontal portion. This is suggestive of their derivation from basal bulges (staminodes) and auricles (filament appendages), which appears to be the general situation in the subfamily Orchidoideae.

Beitr.
Biol.
Pflanzen
66,
(1991),
433
-
477
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
(Orchidaceae)
(Vergleichende
Untersuchungen
zur
Bliitenmorphologie
der
Gattung
Disperis,
Orchidaceae)
By
H.
KURZWEIL
1
and
H.
P.
LINDER
(With
27
figures)
Received
October
16,
1991
Keywords
:
Orchidaceae
Disperis
comparative
flower
morphology
Summary
The
floral
morphology
of
selected
species
of
the
genus
Disperis,
(Orchidaceae-
Orchidoideae-Coryciinae)
is
surveyed.
The
study
concentrates
on
the
little
known
tropical
species
of
the
genus,
supplementing
the
careful
study
of
VOGEL
(1959)
on
the
South
African
species,
which
are
structurally
rather
different
from
most
tropical
species.
Minute
teeth
adpressed
to
the
rostellum
just
above
the
lip
insertion
interpreted
as
vestiges
of
the
lateral
carpels
(lateral
stigma
lobes)
were
observed
in
many
tropical
and
some
South
African
species,
and
appear
to
be
the
basic
condition
in
Disperis.
The
congenital
fusion
of
the
lip
to
the
gynostemium,
which
is
often
quoted
as
being
one
of
the
main
characters
of
the
Coryciinae,
is
shown
to
be
completely
or
almost
completely
absent
in
a
few
species.
The
lateral
gynostemium
appendages
are
distinctly
bi-partite
in
many
species,
with
a
prominent
smooth
erect
portion
and
a
small
sculptured
hori-
zontal
portion.
This
is
suggestive
of
their
derivation
from
basal
bulges
(staminodes)
and
auricles
(filament
appendages),
which
appears
to
be
the
general
situation
in
the
subfamily
Orchidoideae.
Zusammenfassung
Die
Bliitenmorphologie
der
Gattung
Disperis
(Orchidaceae-Orchidoideae-Corycii-
nae)
wurde
fiir
mehrere
Arten
vergleichend
untersucht.
Besonderes
Interesse
gilt
den
tropischen
Vertretern
der
Gattung,
wodurch
die
vorliegende
Arbeit
zu
einer
Ergan-
zung
der
ausfiihrlichen
Arbeit
von
VOGEL
(1959)
fiber
die
morphologisch
weitgehend
andersartigen
sudafrikanischen
Arten
wird.
In
der
Struktur
des
Gynostemiums
besitzen
viele
tropische
and
auch
einige
siidafri-
kanische
Arten
paarige
Fortsatze
am
Rostellum
oberhalb
der
Insertionsstelle
der
434
H.
Kurzweil
and
H.
P.
Linder
Lippe,
die
als
Rudimente
der
lateralen
Narbenlappen
gedeutet
wurden.
Die
Verwach-
sung
Lippe/
Gynostemium,
die
oft
als
eines
der
Hauptmerkmale
der
Coryciinae
ange-
sehen
wird,
fehlt
bei
einigen
Arten
vollig
oder
fast
vollig.
Laterale
Gynostemium-
Anhange
kommen
fast
allgemein
vor,
und
sind
oft
deutlich
zweiteilig:
wahrend
der
aufrechte,
langliche
Anteil
unstrukturiert
ist
und
der
Lippe
angewachsen
ist,
ist
der
kiirzere
Anteil
horizontal
und
stark
skulpturiert.
Dies
deutet
darauf
hin,
daB
an
der
Bildung
dieser
lateralen
Anhange
wahrscheinlich
wie
bei
den
meisten
Orchidoideae
sowohl
„basal
bulges"
(Staminodien)
als
auch
„auricles"
(Filament-Auswiichse)
beteiligt
sind.
Introduction
The
flower
and
especially
gynostemium
and
lip
morphology
of
the
genus
Disperis
(Orchidaceae-Orchidoideae-Coryciinae;
DRESSLER
1981)
is
among
the
most
complicated
to
be
found
in
orchids.
Detailed
floral
morphological
data
are
available
for
some
South
African
species
only
(VOGEL
1959).
Although
adding
invaluable
information
to
our
understanding
of
the
com-
plicated
flower
structures
of
the
genus
Disperis
in
South
Africa,
VOGEL'S
exhaustive
study
provided
very
little
information
with
regard
to
non-South
African
species.
Information
on
the
flower
morphology
is
also
given
in
the
species
descriptions
in
taxonomic
and
floristic
treatments
of
BOLUS
(1888,
1893
-
1896,
1911,
1913),
KRAENZLIN
(1897),
SCHLECHTER
(1898,
1914,
1925),
ROLFE
(1898,
1912
-
1913),
SMITH
(1905,
1908
-
1912),
VERDCOURT
(1968a,
1968b,
1975,
1978,
1988),
WILLIAMSON
(1977),
SEIDENFADEN
(1977),
JAYAWE-
ERA
(1981),
GEERINCK
(1984)
and
CRIBB
&
STEWART
(1985).
However,
these
are
often
not
very
accurate,
and
are
clearly
insufficient
for
a
phylogenetic
anal-
ysis.
There
is
no
detailed
and
comparative
study
of
the
flower
morphology
of
the
genus
available.
With
approximately
90
species
Disperis
is
the
largest
genus
of
the
Coryciinae,
and
is
also
the
only
one
to
extend
beyond
the
African
continent.
The
overall
distribution
area
of
the
genus
with
approximate
species
num-
bers
in
some
regions
is
shown
in
Fig.
1.
While
the
majority
of
species
are
found
in
southern
and
tropical
Africa,
in
Madagascar
and
on
the
Indian
Ocean
islands,
a
few
species
occur
in
India,
Thailand,
the
Phillipines
and
New
Guinea.
This
is
a
distribution
similar
to
Satyrium
(Orchidoideae
-
Satyriinae).
Unlike
the
other
Coryciinae
which
are
mostly
grassland
species,
most
Disperis
species
thrive
in
humid
montane
cloud
forests.
The
enormous
and
frequently
bizarre
lip
appendage
links
Disperis
with
the
other
Coryciinae.
However,
the
genus
is
very
distinct
in
the
subtribe,
and
appears
to
be
rather
isolated
(BoLus,
1888;
VOGEL,
1959).
In
contrast
to
the
other
Coryciinae,
in
Disperis
the
thecae
of
the
anther
are
not
separated
by
an
enlarged
connective,
and
the
anther
is
entirely
covered
by
an
enormous
flat
rostellum.
I
F
In
the
present
paper the
floral
structure
of
various
Disperis
species
is
described.
It
has
been
attempted
to
compare
a
reasonably
large
number
of
species
(41
out
of
approx.
90).
Special
emphasis
is
paid
to
the
little
known
tropical
species,
which
are
structurally
rather
different
from
the
better
known
South
African
species.
Line
drawings,
macro-photos
and
SEM
i
micrographs
are
used
to
illustrate
the
highly
complicated
gynostemium
and
lip
structure.
Brief
comments
on
the
phylogeny
of
Disperis
are
given
in
the
end
of
the
paper.
The
information
presented
in
the
paper
will
form
the
mor-
phological
basis
for
a
study
of
the
detailed
taxonomy
and
phylogeny
of
the
genus
at
a
later
stage.
d
19
2
14
28
Fu;
1.
Distribution
area
of
the
genus
Disperis
highlighting
the
areas
of
highest
species
concentrations.
The
numbers
represent
approximate
species
numbers
(mainly
after
STEWART
et
al.,
1982,
GEERINCK,
1984,
VERDCOURT,
1968
a,
SUMMERHAYES,
1936,
SCHELPE,
1976).
Material
and
methods
Material
was
obtained
from
the
spirit
collections
held
in
the
Kew
Herbarium,
Eng-
land,
and
in
the
Bolus
Herbarium,
University
of
Cape
Town.
Additional
material
was
collected
in
various
parts
of
South
Africa.
The
vouchers
are
listed
in
the
appendix.
Out
of
the
41
species
studied,
7
species
are
described
in
detail
while
the
rest
is
briefly
treated
in
order
to
provide
a
broader
comparison
base
(Table
1).
Macrophotographs
of
living
flowers
were
taken
under
a
WILD
dissecting
micro-
scope.
All
other
material
was
preserved
in
Kew
Cocktail.
For
SEM
purposes,
samples
were
dehydrated
and
critical-point-dried
according
to
the
technique
described
by
GERSTBERGER
&
LEINS
(1978).
Subsequently
the
samples
were
sputter-coated
with
Au/
Pd
and
viewed
and
photographed
in
a
CAMBRIDGE
STEREOSCAN
S200
scanning
29
Beitr.
Biol.
Pflanzen
66/3
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
435
436
H.
Kurzweil
and
H.
P.
Linder
electron
microscope
at
5
kV.
High
resolution
settings
had
to
be
used
throughi
order
to
minimize
the
effect
of
charging.
Abbreviations
AN
anther
AU
lateral
gynostemium
appendages
LA
lip
appendage
LB
lip
blade
LC
lateral
carpel
apices
LCL
lip
claw
0
ovary
P1
-
2
petals
RO
rostellum
ROA
rostellum
arms
S1
-
3
sepals
STG
stigma
Observations
Disperis
katangensis
Summerh.
The
flowers
(Fig.
2
a
-
b)
are
comparatively
large
in
this
species
from
cen-
tral
Africa.
The
ovary
is
elongate
like
in
many
other
Disperis
species,
and
its
six
ovary
ribs
are
not
very
prominent
and
therefore
not
clearly
visible.
Like
in
all
other
Coryciinae
the
median
sepal
is
adnate
to
the
petals,
and
together
with
them
forms
a
hood
enclosing
the
gynostemium
and
the
lip.
The
hood
is
more
or
less
hemi-spherical,
and
almost
as
deep
as
wide.
It
is
mostly
made
up
by
the
enlarged
petals,
while
the
median
sepal
is
rather
narrow.
The
pet-
als
are
unlobed
and
have
slightly
incurved
undulate
front
margins.
Like
in
most
Disperis
species
the
petals
are
extremely
asymmetrical,
with
their
anterior
portion
enormously
developed
while
the
posterior
portion
is
reduced
to
a
narrow
strap.
The
flat
lateral
sepals
are
lanceolate
and
pendent,
and
are
unfused
in
the
specimens
studied
here
(Fig.
2
a).
However,
flowers
with
basally
fused
lateral
sepals
apparently
occur
in
most
other
specimens
(VERDCOURT,
1968
a,
p.
219;
SUMMERHAYES,
1935).
The
lateral
sepals
exhibit
short
saccate
spurs
in
the
middle
near
their
anterior
margin.
As
is
typical
of
Disperis
species
the
lip
is
long
and
narrow,
but
is
relatively
insignificant
as
it
is
projecting
into
the
hood.
It
is
divided
into
four
parts
(Fig.
3
a).
(1).
The
narrow
basal
portion
is
erect
and
fused
to
the
gynos-
temium.
Just
above
this
portion,
the
lip
is
reflexed
forming
a
knee-like
bend
and
is
closely
adpressed
to
the
flat
rostellum.
It
becomes
gradually
widened
in
this
part.
The
two
portions
together
may
be
referred
to
as
the
'lower
claW
(2).
At
the
apex
of
this
claw,
an
enormous
papillose
two-lobed
structure
is
situated
(LA?
in
Fig.
3
a).
It
is
a
massive
body
clearly
wider
than
the
lip
claw.
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
437
S
I
.741
1
4iii
a
2.
Disperis
katangensis
Summerh.;
flower
in
ventral
and
lateral
view
(Milne-
Redhead
3699).
Bar:
5
mm.
Usually
it
is
referred
to
as
the
lip
appendage,
but
it
has
been
suggested
that
it
is
not
homologous
to
the
lip
appendages
found
in
other
Disperis
species
(see
discussion).
(3).
A
further
flat
claw,
which
is
tapering
and
adpressed
to
the
'lower'
claw,
is
developed
between
the
papillose
bilobed
structure
and
the
actual
lip
blade.
The
term
'secondary
claw'
may
be
used
in
easy
reference
to
this
case.
(4).
The
actual
lip
blade
(LB
in
Fig.
3
a)
is
flat
but
hairy
and
papillose
and
has
a
prominent
projecting
crest.
While
the
flat
part
is
hairy
except
along
its
margins
(Fig.
4
a),
the
crest
is
papillose
at
its
apex
and
on
the
anterior
face.
The
gynostemium
(Fig.
3
a
-
d,
4
c
-
d)
is
dominated
by
the
flat
rostellum
covering
both
the
top
and
the
sides
of
the
horizontally
reflexed
anther.
Only
the
apical
part
of
the
anther
is
not
covered
by
the
rostellum
and
is
therefore
visible
when
seen
from
above
(Fig.
3
b).
The
anther
canals
(the
narrow
basal
extensions
of
the
thecae)
are
short
and
not
well
differentiated
from
the
wide
anther
sacs
(Fig.
4
c).
The
thecae
are
not
separated
by
a
prominent
connec-
tive,
and
are
widely
open
after
the
removal
of
the
pollinia.
This
is
typical
of
29*
ROA
RO
438
H.
Kurzweil
and
H.
P.
Linder
AN
1
;1
'0
LA?
A
x
,
LB
RON
1,,
RO
AU
AU
LC
ti
V
a
ROA
—AU
AU
I
C
d
Fig.
3.
Disperis
katangensis
Summerh.;
gynostemium
(Milne-Redhead
3699).
a:
three-quarter
view,
lip
attached.
b:
top
view.
c:
front
view.
d:
lateral
view.
Bar:
1
mill
the
genus
Disperis
(VOGEL,
1959),
(Fig.
4c).
Apically,
the
anther
has
a
short
rounded
connective
process
(Fig.
3
b).
Lateral
gynostemium
appendages
are
clearly
visible
in
this
species
(AU
in
Fig.
3
b
-
d,
4
c).
Two
distinct
portions
can
be
distinguished.
While
the
small
lower
part
is
horizontal
and
heavily
sculptured,
the
elongate
upper
part
is
erect
and
smooth.
It
is
fused
to
the
lip
base
and
is
visible
from
the
front
side
440
H.
Kurzweil
and
H.
P.
Linder
beyond
the
lip
insertion
(Fig.
3
d).
Their
terminal
portions
are
widened
and
bent
upwards.
The
receptive
surface
is
borne
on
two
separate
stigmas
which
are
situated
in
the
anterior
and
lateral
parts
of
the
central
rostellum
lobe.
They
face
upwards
and
sideways,
are
flat
and
thus
not
raised
above
the
rostellum
sur-
face,
and
are
papillose
throughout
(STG
in
Fig.
4d).
Disperis
thomensis
Summerh.
This
species,
distributed
in
West
and
East
Africa,
can
be
readily
distin-
guished
from
other
species
by
the
flowers
with
deeply
saccate
hoods
point-
ing
backwards
and
lobed
petals
(Fig.
5b).
The
median
sepal
is
a
narrow
strap
connecting
the
two
petals,
although
it
is
naviculate
throughout
and
bent
around
the
sac.
The
petals
are
the
dominating
structures
of
the
galea,
and
extend
far
into
the
sac.
They
have
prominent
basal
anterior
lobes
and
somewhat
reflexed
triangular
anterior
apical
lobes.
Like
in
the
preceeding
species
the
petals
are
strongly
asymmetrical
with
their
posterior
half
con-
LC
L
S
i
ROA
i%
1
a
Fig.
5.
Disperis
thomensis
Summerh.;
flower
in
ventral
and
lateral
view
(Milne-Retl
'
head
4227).
Bar:
1
mm.
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
441
fined
to
a
narrow
strap
only.
The
lateral
sepals
are
pendent
and
free
at
their
base.
They
are
apically
acute,
and
have
comparatively
long,
cone-like
and
broadly
rounded
spurs.
Like
in
D.
katangensis
these
lateral
sepal
spurs
are
situated
half-way
up
the
sepal
and
close
to
the
anterior
margin.
The
narrow
lip
claw
is
suberect
in
its
basal
portion
which
is
fused
with
the
gynostemium
(Fig.
6
a).
Higher
up,
it
is
knee-like
bent
and
strongly
reflexed.
Further
above
(behind)
the
lip
claw
takes
a
"U"-shaped
bend,
and
the
termi-
nal
part
which
is
then
projecting
forwards
is
a
complex
structure
made
up
by
two
deeply
bilobed
processes.
This
complex,
consisting
of
altogether
four
arms
obviously
represents
the
lip
appendage
(LA
in
Fig.
6
a).
The
lip
blade
is
represented
by
the
narrow
lorate
lobe
projecting
into
the
"U"
of
the
lip
claw
(LB
in
Fig.
6
a).
The
upper
surface
of
this
lip
blade
is
covered
with
multicel-
lular
trichomes
which
each
carry
numerous
globular
glands
on
short
stalks
(Fig.
6
d
-
e).
Interestingly,
the
glands
near
the
apex
of
the
lip
appendage
(Fig.
6
f)
are
very
similar
in
shape,
but
are
solitarily
attached
to
the
lip
appendage.
The
gynostemium
consists
mainly
of
the
horizontally
reflexed
anther
which
is
covered
by
the
flat
median
rostellum
lobe.
The
rostellum
arms
are
simple
spathulate
structures
pointing
upwards.
Their
terminal
viscidia
bearing
portions
are
somewhat
widened
and
angled
upwards
(ROA
in
Fig.
6
a
-
b).
As
in
D.
katangensis,
the
anterior
part
of
the
median
rostellum
lobe
forms
a
prominent
raised
ridge.
It
is
confluent
with
the
vestiges
of
the
lateral
carpel
apices
(LC)
and
with
the
lateral
rostellum
lobes
(ROA
in
Fig.
6
b).
The
stigmas
(STG
in
Fig.
6
c)
are
situated
in
the
anterior
and
lateral
parts
of
the
rostellum.
The
rostellum
and
consequently
also
the
stigmas
are
strongly
bent
in
this
portion,
resulting
in
the
stigmas
facing
upward
and
sideways.
Lateral
gynostemium
appendages
are
clearly
developed,
and
are
rather
small
structures.
Disperis
zeylanica
Trimen
D.
zeylanica
(Sri
Lanka,
India)
is
the
only
Asian
Disperis
species
studied
here.
The
hood
is
very
shallowly
concave
although
strongly
curved
(which
may
be
referred
to
as
`vexillum-like'),
and
is
mainly
made
up
of
the
large
petals
(Fig.
7
a
-
b).
The
latter
are
unlobed
and
have
undulate
front
margins.
Like
in
the
previous
species
they
are
extremely
asymmetrical,
and
mainly
Consist
of
the
enlarged
anterior
portions.
The
pendent
lateral
sepals
are
broadly
lanceolate
and
are
basally
slightly
connate.
They
exhibit
short
cone-
like
and
obtuse
spurs
in
their
lower
third
near
their
front
margin.
The
lip
is
basically
similar
in
shape
to
that
described
in
D.
katangensis
(Fig.
7
d).
It
is
differentiated
into
claw,
large
bilobed
structure
(LA?
in
Fig.
7
d;
possibly
homologous
with
the
lip
appendage),
secondary
claw
and
ha.
442
H.
Kurzweil
and
H.
P.
Linder
LA
LCL
ROA
LB
R
0
-
ROA
ST
G
LA
'A-Vt*.0
,
L8
LCL
orN
t
0
NAP5wIlimpr
Fig.
6.
Disperis
thomensis
Summerh.;
details
of
the
gynostemium-lip-compl
ex
(Milne-Redhead
4227).
a:
gynostemium
in
three-quarter
view,
lip
attached.
b:
anterior
part
of
gynostemium
in
top
view.
c:
stigma.
d
-
e:
glands
around
the
lip
blade.
f:
glands
at
the
apex
of
the
lip
appendage.
SEM
micrographs;
bars:
0.5
mm
in
a
-
d.
0.2
mm
e
-
f.
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
443
pia
,/
‘\`‘‘
L
A?
LB
a
AN
RO
LA?
-
LB
AU
TG
LC
C
g.
7.
Disperis
zeylanica
Trimen
(Barnes
s.
n.);
a
-
b:
flower
in
ventral
and
lateral
ew.
c:
gynostemium
in
top
view.
d:
gynostemium-lip-complex
in
three-quarter
view.
bars:
1
mm.
RO
AU
444
H.
Kurzweil
and
H.
P.
Linder
lip
blade.
The
basal
part
of
the
claw
is
again
extensively
fused
to
the
gynos-
temium.
The
'lip
appendage'
is
bilobed
with
two
narrow
horn-like
diverging
lobes,
which
are
papillose
on
their
upper
surface
(Fig.
8
b).
The
orientation
of
the
'secondary
claw'
could
not
be
determined
in
the
pickled
material
seen.
i
LB
of
1.
41.
VIt
.
:15$
NWAS'
1
/
4
.
Li
b
Fig.
8.
Disperis
zeylanica
Trimen;
details
of
gynostemium-lip-complex
(Barnes
s.n.).
a:
gynostemium
in
front
view
showing
the
lateral
appendages.
b:
papillae
of
the
'lip
appendage'.
SEM
micrographs;
bars:
0.5
mm
in
a,
0.2
mm
in
b.
The
anther
is
more
or
less
horizontal
and
is
largely
covered
by
the
flat
and
apically
emarginate
central
rostellum
lobe
which
is
infolded
in
the
middle
(RO
in
Fig.
7c).
The
apex
of
the
anther
is
emarginate
as
well
and
is
visible
from
above
(AN
in
Fig.
7
c).
The
lateral
gynostemium
appendages
are
dif-
ferentiated
into
small
sculptured
horizontal
portions
and
elongate
smooth
vertical
portions
which
are
fused
to
the
lip
base
but
terminate
in
two
free
tips
(AU
in
Fig.
7
d,
8
a).
In
contrast
to
D.
katangensis,
the
erect
smooth
por-
tions
fused
to
the
lip
base
are
very
narrow
here.
The
central
rostellum
lobe
bears
the
two
stigmas
near
the
bases
of
the
rostellum
arms.
The
latter
are
simple
flat
structures
projecting
forwards
and
upwards,
and
have
enlarged
apices.
Like
in
the
previous
species
they
are
confluent
with
a
fleshy
ridge
at
the
anterior
margin
of
the
rostellum.
The
two
vestiges
of
the
lateral
carpel
apices
are
comparatively
small
in
this
species
(LC
in
Fig.
7
c).
Disperis
wealei
Rchb.
f.
This
species
is
included
as
an
example
of
the
South
African
Disperis
species.
It
has
a
wide
distribution
area,
ranging
from
the
Eastern
Cape
to
Natal
and
Transvaal.
The
deeply
saccate
hood
(Fig.
9
a
-
b)
encloses
the
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
445
S
i
?.•
A
ROA
a
Fig.
9.
Disperis
wealei
Rchb.
f.;
flower
in
ventral
and
lateral
view
(Linder
2088).
Bar:
2
mm.
r[1,
gynostemium
except
the
lateral
rostellum
lobes
which
project
forward.
The
hood
is
mostly
made
up
by
an
enormous
median
sepal,
while
the
petals
are
rather
narrow.
The
petals
are
unlobed
and
more
or
less
symmetrical
with
two
almost
equally
wide
portions
on
both
sides
of
the
midnerve.
On
the
outer
side
the
petal
midnerve
is
extraordinarily
strongly
developed.
The
lanceo-
late
lateral
sepals
are
pendent,
strongly
diverging,
and
not
fused.
The
lateral
sepal
spurs
are
cone-like
and
rather
wide
at
their
point
of
insertion.
The
lip
has
a
narrow
claw
which
is
extensively
fused
to
the
gynostemium.
The
enormous
boat-shaped
complex
with
its
entire
terminal
process
(LA)
evidently
constitutes
the
lip
appendage,
while
the
lip
blade
is
represented
by
the
lanceolate
anterior
tip
of
the
'boat'
(LB
in
Fig.
10
a).
The
terminal
process
on
the
lip
appendage
is
superficially
seen
smooth,
but
exhibits
fine
papillae
on
closer
examination
(STEINER,
1989).
After
removal
of
the
lip
the
structure
of
the
gynostemium
can
be
seen.
The
anther
is
reflexed
at
an
angle
of
approximately
160°
(AN
in
Fig.
10
c).
Only
its
rounded
apex
is
visible
from
above
(Fig.
10
b).
The
widely
open
thecae
and
the
occurrence
of
comparatively
prominent
anther
canals
are
shown
in
Fig.
10
d.
Lateral
gynostemium
appendages
are
clearly
developed
(AU
in
Fig.
10
c).
They
are
horizontal
in
their
posterior
part
and
vertical
and
adnate
to
the
lip
in
their
anterior
part.
In
contrast
to
the
species
described
446
H.
Kurzweil
and
H.
P.
Linder
LA
AN
%
RO
LB
RO
STG
LC
ROA
a
b
STG
AN
fFl
ROA
AU
d
Fig.
10.
Disperis
wealei
Rchb.
f.;
gynostemium
(Linder
2088);
a:
three-quarter
view,
lip
attached.
b:
top
view.
c:
lateral
view.
d:
top
view,
rostellum
removed.
Bar:
1
Mrn•
AN
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
447
above
(in
which
the
erect
portion
is
smooth),
they
are
sculptured
through-
out.
The
central
rostellum
lobe
is
flat
and
covers
the
top
and
the
sides
of
the
anther
(RO
in
Fig.
10
b).
It
is
constricted
in
the
middle,
and
coarsely
dentate
at
the
margins.
Unlike
in
the
species
described
above,
the
two
stigmas
are
situated
in
the
median
anterior
part
of
the
rostellum,
and
touch
each
other
in
the
median
line
of
the
gynostemium
(STG
in
Fig.
10
b).
They
are
remark-
ably
large,
occupying
almost
one
third
of
the
entire
length
of
the
central
ros-
tellum
lobe.
The
lateral
rostellum
arms
are
very
complicated
structures
(ROA
in
Fig.
10
b
-
c).
They
are
canaliculate
and
knee-like
bent
in
their
lower
and
descending
portion.
Their
terminal
viscidia-bearing
parts
are
non-canaliculate
and
angled
upwards
and
to
the
sides.
The
tissue
of
the
lat-
eral
rostellum
arms
extends
to
just
in
front
of
the
stigmas.
It
is
not
confluent
with
the
vestiges
of
the
lateral
carpel
apices
which
take
the
shape
of
short
triangular
teeth
in
this
species
(LC
in
Fig.
10
b).
The
occurrence
of
a
small
structure
between
these
teeth
is
probably
an
abnormality
(see
discussion).
Disperis
virginalis
Schltr.
The
hood
of
this
southern-central
African
species
is
deeply
saccate
(Fig.
11
a
-
b).
The
median
sepal
is
comparatively
broad,
and
is
adnate
to
the
similarly
well
developed
petals.
The
latter
have
obscure
anterior
lobes
(the
degree
of
their
formation
is
variable
in
the
species)
and
are
apically
dilated
with
large
recurved
lobes.
The
broadly
lanceolate
lateral
sepals
are
descend-
ing
and
are
fused
in
their
lower
third.
They
exhibit
cone-like
obtuse
spurs
with
very
wide
entrances.
The
lip
has
an
elongate
narrow
claw
which
has
the
shape
of
a
horizontal
"S"
when
seen
from
the
side
(LCL
in
Fig.
11
c).
While
the
claw
is
fused
to
the
gynostemium
at
the
base,
it
is
reflexed
and
adpressed
to
the
rostellum
above
this
fused
portion,
recurved
behind
the
gynostemium
and
curved
upwards
in
its
distal
part.
The
terminal
portion
is
deeply
split
into
two
bilobed
struc-
tures
which
are
finely
papillose.
This
complex
made
up
of
altogether
four
lobes
is
basically
similar
to
that
of
D.
thomensis,
and
obviously
represents
the
lip
appendage
(LA).
The
lip
blade
itself
is
a
minute
lorate
process
and
is
densely
hairy
at
its
base
(LB
in
Fig.
11
c,
12
a).
Unlike
in
the
Disperis
species
described
above,
the
gynostemium
is
shortly
stalked
and
is
situated
well
in
front
of
the
hood
(Fig.
11
b
-
d).
The
horizontal
anther
is
almost
as
long
as
the
central
rostellum
lobe
which
covers
its
top
and
sides.
This
central
rostellum
lobe
bears
the
two
separate
stigmas
close
to
the
base
of
the
rostellum
arms.
The
narrow
rostellum
arms
are
straight
and
vertical,
and
have
spoon-like
enlarged
terminal
parts
bearing
the
viscidia
(ROA
in
Fig.
11
d,
12
b).
Like
in
Disperis
katangensis
the
rostellum
arms
are
confluent
with
a
fleshy
ridge
on
the
anterior
margin
of
the
central
rostellum
448
H.
Kurzweil
and
H.
P.
Linder
RO
A
S3
.1.
I
a
b
ROA
LA
LB
R
0
SIG
AN
c
d
n.
AU
LCL
Fig.
11.
Disperis
virginalis
Schltr.
(S.
G.
H.
114,
239);
a
-
b:
flower
in
ventral
and
lat-
eral
view.
c:
gynostemium
and
lip
in
three-quarter
view.
d:
gynostemium
in
lateral
view.
Bars:
1
mm.
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
449
LA
R
0
LC
ROA
LB
12.
Disperis
virginalis
Schltr.
(S.
G.
H.
114,
239);
a:
apex
of
lip.
b:
anterior
part
of
gynostemium
in
top
view.
SEM
micrographs,
bars:
1
mm.
be
and
with
the
rounded
teeth
of
the
lateral
carpel
apices
(LC
in
Fig.
12
b).
e
lateral
gynostemium
appendages
are
comparatively
small
structures
U
in
Fig.
11
d).
Disperis
anthoceros
Rchb.
f.
This
species
has
a
vast
distribution
area
in
Africa,
ranging
from
Nigeria
in
e
west
to
Kenya
in
the
east
and
Natal
(South
Africa)
in
the
south.
The
hood
of
the
flower
is
developed
into
a
slender
elongate
spur
which
entirely
is
made
up
by
the
median
sepal
(Fig.
13
a
-
b).
The
lip
and
its
appendage
are
hidden
in
this
spur,
although
they
reach
only
half-way
up
(Fig.
13
c).
Only
the
very
base
of
the
lip
is
fused
to
the
gynostemium.
The
petals
are
entire
and
are
adnate
to
the
median
sepal.
The
strongly
developed
rib
on
the
outer
petal
surfaces
adjacent
to
the
margin
of
the
median
sepal
clearly
indi-
cates
the
position
of
the
midnerve,
revealing
that
the
largest
petal
portion
is
derived
from
the
anterior
part
while
the
posterior
part
is
only
the
narrow
strap
adnate
to
the
median
sepal.
The
lateral
sepals
are
spreading
and
only
slightly
descending.
They
are
united
in
their
lower
third,
and
exhibit
short
conical,
rounded
spurs
in
their
middle.
The
narrow
lip
claw
is
erect
in
its
basal
part,
is
then
reflexed
and
adpres-
sed
to
the
rostellum,
and
is
erect
in
its
elongate
main
portion
(LCL
in
Fig.
13
c).
This
main
portion
is
of
rather
late
ontogenetic
origin,
as
it
is
still
lacking
in
an
earlier
ontogenetic
stage
(Fig.
14
b).
The
lip
appendage
on
the
apex
of
this
claw
is
developed
as
a
bunch
of
two
minute
fimbriate
lobules
which
are
smooth
(LA
in
Fig.
14
a).
The
lip
blade
is
a
minute
process,
and
is
completely
hidden
among
these
fimbrae.
It
is,
however,
clearly
visible
on
an
immature
gynostemium
(LB
in
Fig.
14
b).
450
H.
Kurzweil
and
H.
P.
Linder
LA
LC
L
r\\
7/.
./
C
a
\ ,
b
MO,
RO
STG
R
OA
d
e
Fig.
13.
Disperis
anthoceros
Rchb.
f.
(Eggeling
5949);
a
-
b:
flower
in
ventral
and
lat-
eral
view.
c:
gynostemium
and
lip.
d
-
e:
gynostemium
in
top
and
in
three-quarte
r
view.
Bars:
1
mm.
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
451
LA
LB
RO
R
OA
Fig.
14.
Disperis
anthoceros
Rchb.
f.
(Eggeling
5949);
a:
lip
appendage.
b:
gynos-
temium
and
lip
in
a
young
ontogenetic
stage.
SEM
micrographs,
bars:
0.5
mm.
The
gynostemium
mostly
consists
of
the
horizontally
reflexed
anther
and
the
slightly
shorter
and
infolded
central
rostellum
lobe
which
covers
it
on
top
and
on
the
sides
(RO
in
Fig.
13
e).
The
stigmas
are
very
narrowly
lanceo-
late
structures
at
the
base
of
the
short
untwisted
rostellum
arms.
The
lateral
gynostemium
appendages
of
this
species
are
very
insignificant
structures
in
the
mature
flower.
Disperis
kilimanjarica
Rendle
This
species,
known
from
Central
and
East
Africa,
is
very
distinct
among
the
tropical
Disperis
species
because
of
its
unusual
lip.
The
ovary
has
a
remarkably
narrow
neck.
The
hood
encloses
the
gynostemium,
and
mainly
consists
of
the
median
sepal
with
its
horizontal
sac-like
spur
(S
1
in
Fig.
15
a
-
b).
The
entire
petals
are
somewhat
clawed
and
are
slightly
longer
than
the
median
sepal.
The
pendent
lateral
sepals
are
acute-lanceolate,
and
are
fused
at
the
very
base
only.
They
exhibit
short
but
rather
wide
sac-like
spurs
with
very
broad
entrances.
The
lip
(Fig.
15
c)
has
a
narrow
claw.
It
is
adpressed
to
the
basally
erect
and
then
recurved
stigma,
and
is
fused
to
the
gynostemium
at
the
very
base
only.
At
the
upper
end
of
the
stigma
the
claw
forms
a
knee-like
bend
to
the
back.
The
lip
blade
is
an
elongate-triangular
process
and
is
pointing
for-
wards
(LB
in
Fig.
15
c).
The
lip
appendage
(LA)
is
solid
and
entire,
and
is
projecting
horizontally
into
the
spur.
The
gynostemium
is
unstalked
and
exhibits
a
weakly
reflexed
anther
(Fig.
15
c
-
d).
The
rostellum,
which
covers
the
front,
top
and
the
sides
of
the
30
Beitr.
Biol.
Pflanzen
66/3
0
s,
452
H.
Kurzweil
and
H.
P.
Linder
\
p
1
/2,
a
LA
LB
RO
AN
STG
RO
ROA
Fig.
15.
Disperis
kilimanjarica
Rendle
(Mabberley
&
McCall
13);
a
-
b:
flower
in
v
tral
and
lateral
view.
c:
gynostemium
and
lip.
d:
gynostemium
in
top
view.
BF
1
mm.
anther
is
also
only
weakly
reflexed
to
approximately
120°.
The
stigma
entire
and
apically
deeply
emarginate
(STG
in
Fig.
15d).
It
is
very
large
occupies
most
of
the
central
rostellum
lobe.
The
stigma
is
rather
narrow
its
basal
portion
where
it
is
restricted
in
width
by
the
rostellum
arms
(RC
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
453
in
Fig.
15
d).
The
latter
are
very
short
but
twisted
and
coiled
up,
so
that
the
viscidia-bearing
parts
are
facing
inwards.
Discussion
A)
Morphology
(see
also
table
1)
Perianth
The
hood,
generally
made
up
by
the
median
sepal
adnate
to
the
two
petals,
is
a
characteristic
feature
of
Disperis.
However,
a
hood
composed
of
the
median
sepal
and
the
two
petals
occurs
also
in
other
Coryciinae,
in
Brownleea
(usually
placed
in
Disinae)
and
in
some
Orchideae
(particularly
in
Cynorkis),
which
is
probably
due
to
parallel
evolution.
The
hood
may
take
the
form
of
a
variously
shaped
sac,
of
a
slender
spur
or
of
an
only
weakly
concave
but
strongly
bent
lobe
(vexillum-like).
The
spur
or
sac
may
be
derived
from
the
median
sepal
alone
(D.
anthoceros,
Fig.
13
a
-
b;
D.
kilimanjarica,
Fig.
15
a
-
b;
also
in
D.
woodii,
D.
stenoplectron,
D.
disaeformis
and
D.
cucculata).
This
condition
was
referred
as
the
`D.
anthoceros-type'
by
VOGEL
(1959,
p.
193).
In
other
species
the
median
sepal
is
a
narrow
organ
only,
and
posterior
petal
lobes
form
the
side
walls
of
the
spur.
This
is
here
shown
in
D.
thomensis
(Fig.
5
b)
and
D.
kerstenii
(Fig.
16),
but
is
also
found
in
D.
aphylla,
D.
bifida
and
D.
uzungwae.
The
term
`D.
kamerunensis-type'
was
used
for
this
situa-
tion
(VoGEL,
1959,
p.
193).
VOGEL
also
recognized
a
third
type,
the 'D.
lindle-
yana-type',
which
was
described
as
an
intermediate
condition.
The
lateral
sepals
are
basally
congenitally
more
or
less
united
or
are
com-
pletely
free.
The
fused
portion
is
usually
less
than
one
third
of
the
sepal
length,
only
D.
egregia
has
almost
entirely
fused
lateral
sepals
(VERDCOURT,
1968
a).
The
lateral
sepals
are
usually
descending,
pendent
or
rarely
slightly
reflexed
(D.
fanniniae).
They
are
either
almost
parallel
(D.
katangensis,
Fig.
2
a;
D.
zeylanica,
Fig.
7
a;
D.
kilimanjarica,
Fig.
15
a)
or
more
or
less
spreading
(D.
wealei,
Fig.
9
a).
In
some
species
the
lateral
sepals
diverge
at
an
angle
of
almost
180°
(most
notably
in
D.
circumflexa,
Fig.
18d;
D.
cue-
culata,
Fig.
19
a;
D.
villosa).
The
lateral
sepals
are
usually
more
or
less
acute,
but
caudate
tips
occur
in
D.
capensis
(Fig.
18
a).
They
are
probably
vestiges
of
apiculi
which
are
well
developed
in
Disa
(LINDER,
1981
a
-
b,
LINDER
&
KURZWEIL,
1990).
While
the
lateral
sepals
are
more
or
less
naviculate
in
most
species,
they
are
flat
in
some
species
and
thus
form
a
'platform'
dominating
the
whole
flower
appearance
(e.g.
D.
johnstonii,
D.
togoensis).
Spurs
on
the
lateral
sepals
are
found
in
all
Disperis
species
and
are
unique
the
Orchidoideae.
They
provide
the
best
diagnostic
character
of
the
genus,
d
generally
have
been
used
in
orchid
classifications
and
floristic
accounts.
30
Table 1.
4=.
cn
Distribution
of
features
of
particular
importance
in
the
species
studied.
Disperis
oppositifolia
has
been
omitted
from
the
table
as
the
mate-
rial
4=•
was
too
fragmentary
for
study.
Abbreviations:
distr
(geographical
distribution):
1
...
tropical
Africa,
2
...
tropical
Asia,
3
...
South
Africa,
4
...
Indian
Ocean
Islands
(including
Madagascar).
hood:
sp
spur-like,
sac
...
saccate,
vex
...
vexillum-like
(=
very
shallowly
con-
cave).
pet
(petals):
sym
...
symmetrical,
as
...
strongly
asymmetrical,
int
...
intermediate.
s.
fus
(lateral
sepal
fusion):
1/3
...
one
third,
±
base
only,
0
unfused.
lip
(lip
type):
virg
Virginalis
type,
kat
...
Katangensis
type,
weal
...
Wealei
type,
card
...
Cardiophora
type.
1.
fus
(fusion
of
the
lip
to
the
gynostemium):
ext
...
extensively,
±
...
weakly,
0
unfused.
roa
(rostellum
arms):
tw
...
twisted,
untw
untwisted,
±
...
slightly
twisted.
stg
(stigma):
bas
...
on
base
of
gynostemium
front
side,
top
...
on
top
of
the
rostellum,
lat
...
close
to
the
lateral
rostellum
arms.
lc
(vestiges
of
lateral
carpel
apices):
+
...
present,
...
absent.
au
(lateral
gynostemium
appendages):
+
...
clearly
bipartite
with
prominent
smooth
upper
portion
fused
to
the
lip
base,
...
not
clearly
bipartite
or
upper
portion
inconspicuous.
?
no
data
available
(material
collapsed).
NA
not
applicable
(see
text).
Jop
lin
'd
'H
P
u
s
'I
ma
m);
'H
sp
sp
sp
sac
sac
vex
sac
sp
sac
sp
vex
vex
sp
vex
vex
sac
vex
distr
`TROPICAL
GROUP'
D.
anthoceros
1,3
D.
aphylla
1
D.
bifida
1
D.
dicerochila
1
D.
elaphoceras
1
D.
johnstonii
1,3
D.
katangensis
1
D.
kerstenii
1
D.
leuconeura
1
D.
nemorosa
1
D.
pusilla
1
D.
reichenbachiana
1
D.
thomensis
1
D.
togoensis
1
D.
tripetaloidea
4
D.
virginalis
1,3
D.
zeglanica
2
pet
s.
fus
lip
1.
fus
roa
stg
lc
au
as
1/3
virg
±
untw
lat
as
±
virg
ext
untw
lat
+
as
1/3
virg
ext
untw
lat
+
+
as
0
virg
ext
untw
lat
+
+
as
0
kat?
ext
untw
lat
+
+
as
±
kat
ext
untw
lat
+
+
as
0?
kat
ext
untw
lat
+
+
as
0
virg
ext
untw
lat
+
+
?
0
kat
?
?
? ?
?
±
virg
? ?
lat
?
as
0
kat?
ext
untw
lat
+
as
±
kat
ext
untw
top-lat
+
+
as
0
virg
ext
untw
lat
+
as
1/3
kat
ext
untw
top-lat
+
+
as
0
kat
ext
untw
lat
+
+
as
±
--
1/3
virg
ext
untw
lat
+
+
as
±
kat
ext
untw
lat
+
+
hood
01
distr
hood
pet
s.
fus
lip
1.
fus
roa
stg
lc
au
`SOUTH
AFRICAN
GROUP'
D.
bodkinii
3
sp
int
0
card
±
tw
bas
D.
bolusiana
3
sac
int
0
weal
0
±
tw
bas
D.
breviloba
1
sp
as
0
card
0
?
?
?
?
D.
capensis
3
sac
as
0
card
ext
tw
NA
+
D.
cardiophora
3
sac
as
0
card
ext
tw
top
+
D.
circumflexa
3
sac
as
0
weal
ext
tw
top
+
D.
cooperi
3
sac
sym
0
weal
ext
tw
top
+
+
D.
cucculata
3
sac
int
0
weal
0
±
bas
D.
decipiens
1
sac
as
0
NA
±
tw
? ? ?
D.
fanniniae
3
sac
int
0
weal
ext
tw
top
+
D.
kilimanjarica
1
sp
as
±
card
±
tw
bas
D.
lindleyana
3
sac
int
0
card
ext
tw
top
+
+
D.
macowanii
1,3
sac
int
0
card
0
tw
bas
NA
D.
micrantha
3
sac
as
0
card
±
tw
bas
D.
oxyglossa
3
sac
as
0
weal?
ext
tw
top
+ +
D.
paludosa
3
vex
as
0
weal?
ext
±
top
+
D.
purpurata
3
sac
int
0
weal
0
tw
bas
+
D.
thorncroftii
3
sac
as
0
card
ext
tw
top
+
?
D.
tysonii
3
sac
int
0
?
ext
tw
top
D.
uzungwae
1
sp
?
1/3
card?
ext
±
lat
+
D.
villosa
3
sac
as
0
weal
0
tw
bas
D.
wealei
3
sac
sym
0
weal
ext
tw
top
+
+
D.
wood
ii
3
sp
as
0
card
±
tw
?
?
A
com
par
ati
ve
st
ud
y
of
th
e
fl
or
al
m
or
ph
ol
o
g
y
i
n
th
e
genus
Di
s
peri
s
456
H.
Kurzweil
and
H.
P.
Linder
S,
LA
RO
n1
1
STG
S3
Fig.
16.
Disperis
kerstenii
Rchb.
f.;
flower
in
three-quarter
view
(Eggeling
6334).
Bar:
1
mm.
They
are
always
short,
and
are
mostly
developed
as
cylindrical
or
conical
structures
of
various
shapes.
In
some
species
they
are
strongly
reduced
and
very
shallow,
but
species
with
obsolete
lateral
sepal
spurs
were
not
observed.
The
spurs
are
usually
inserted
half-way
up
the
sepals
near
their
anterior
margin.
While
most
tropical
and
South
African
species
are
rather
uniform
as
regards
their
lateral
sepal
spurs,
a
few
species
from
the
Western
Cape
of
South
Africa
differ
considerably
in
their
sepal
spur
shape
or
inser-
tion:
in
D.
purpurata
and
D.
bolusiana
the
spurs
are
not
clearly
developed
but
the
whole
of
the
lateral
sepal
has
the
shape
of
cone-like
sacs;
in
D.
vil-
losa
the
saccate
spurs
are
borne
in
subterminal
position
and
face
to
the
sides;
in
D.
cucculata
the
spurs
are
inserted
in
the
middle
of
strongly
curved
lateral
sepals,
which
have
their
apices
projecting
upwards
in
their
terminal
part
(Fig.
19
a).
The
petals
are
rather
diverse
in
shape
(Fig.
20
a
-
f).
They
are
usually
entire
and
prominent
basal
anterior
lobes
occur
only
rarely
in
the
genus
(D.
thomensis,
Fig.
5
a
-
b;
D.
kerstenii,
Fig.
16;
D.
dicerochila,
Fig.
20a;
D.
vir-
ginalis,
Fig.
20
b;
D.
nemorosa).
They
are
reminiscent
of
similar
structures
found
in
Disa
and
Herschelianthe
where
they
have
been
used
as
taxonomi
-
cally
important
characters
(LINDER,
1981
a
-
c).
LA
LB
ROA
ti
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
457
a
ST
G
R
0
LC
Fig.
17.
Disperis
kerstenii
Rchb.
f.;
details
of
lip
and
gynostemium
(Eggeling
6334).
a:
lip
appendage.
b:
lip
blade.
c:
gynostemium
in
top
view.
d:
lateral
rostellum
arm.
SEM
micrographs;
bars:
1
mm
in
a
-
c,
0.1
mm
in
d.
The
petals
are
generally
free
from
the
gynostemium
except
in
a
few
Dis-
peris
species
from
the
southwestern
Cape
of
South
Africa
(especially
in
D.
villosa;
to
a
much
lesser
extent
also
in
D.
bolusiana
and
D.
cucculata).
Such
a
fusion
petals
/
gynostemium
is
similar
to
what
is
found
in
the
Disinae
(in-
cluding
Brownleea),
and
in
Brachycorythis,
Neobolusia
and
Schwartzkopf-
fia.
However,
it
remains
to
be
shown
whether
the
sporadic
appearance
of
this
phenomenon
is
due
to
parallel
evolution.
The
petals
are
±
symmetrical
in
some
South
African
species
(here
shown
in
D.
fanniniae,
Fig.
20
f;
D.
cooperi,
Fig.
20
c).
In
the
majority
of
species
they
are
extremely
asymmetrical
with
their
dorsal
portion
confined
to
the
narrow
strap
of
tissue
which
is
adnate
to
the
median
sepal,
as
can
be
recognized
by
the
position
of
the
midnerve.
While
symmetrical
petals
are
probably
primi-
458
H.
Kurzweil
and
H.
P.
Linder
S,
P
2
LC
R
0
ROA
LA
STG
STG
O
S
1
Fig.
18.
a
-
c:
Disperis
capensis
Sw
.;
a:
flower
(Kurzweil
901).
b:
gynostemium
and
lip
in
lateral
view
(Kurzweil
901).
c:
gynostemium
in
ventral
view
(Hall
701).
d:
Disperis
circumflexa
(L.)
Dur.
&
Schinz,
flower
(Bol
p
s.n.).
c:
SEM
micrograph;
bars:
2
mm.
tive
in
the
orchids
in
general,
strongly
asymmetrical
petals
are
the
basic
condition
in
the
Coryciinae.
This
suggests
that
the
symmetrical
petals
of
some
Disperis
species
may
well
be
a
secondary
simplification.
The
petal
midnerve
is
frequently
extraordinarily
strongly
developed
on
the
outer
petal
surface
over
most
of
the
petal
length,
and
is
visible
on
the
closed
bud
between
the
sepals.
Similar
extraordinarily
strongly
developed
petal
nerves
are
also
found
in
other
Coryciinae,
in
Vanilla
(Epidendroideae)
and
in
Apostasia
(Apostasioideae).
Among
other
Orchidoideae,
strongly
AU
STG
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
459
ROA
S
i
RO
LCL
S
2
A
,R0
L
ROAD'
Fig.
19.
a
-
b:
Disperis
cucculata
Sw.;
a:
flower
(Bol
e
s.
n.).
b:
gynostemium
(Linder
1137).
c
-
d:
Disperis
villosa
(L.
f.)
Sw.
(Kurzweil
1206);
c:
gynostemium
and
lip.
d:
base
of
gynostemium
in
lateral
view.
b:
SEM
micrograph;
b
-
d:
SEM
micrographs;
bars:
5
mm
in
a,
0.5
mm
in
b,
1
mm
in
c
-
d.
developed
petal
midnerves
are
also
found
in
Brachycorythis
(Orchideae)
and
Brownleea
(Diseae)
where
they
are
extremely
short.
The
shape
of
the
lip
shows
the
greatest
diversity
of
all
flower
structures
found
in
the
genus
and
has
been
used
as
a
key
character
in
most
floristic
accounts.
The
lip
has
a
narrow
claw
in
all
species,
and
a
frequently
bizarre
appendage
on
top
of
this
claw.
The
appendage
and
the
lip
blade
(limb)
fre-
quently
form
together
a
complex
entity
making
it
difficult
to
clearly
distin-
guish
between
the
two
parts.
The
highest
degree
of
complexity
is
found
in
species
with
boat-shaped
and
`pseudopeltate'
lips
(sensu
VOGEL,
1959),
where
the
margins
of
the
lip
blade
are
united
with
the
appendage.
Disperis
decipiens
(Fig.
23),
a
species
described
from
Tanzania
(VERDCOURT,
1975),
has
an
inappendiculate
lip.
However,
the
lack
of
the
appendage
in
this
species
is
probably
due
to
secondary
loss.
4611
H.
Kurzweil
and
H.
P.
Linder
Fig.
20.
Petals
of
various
Disperis
species.
a:
D.
dicerochila
Summerh.
(Bridson
419).
b:
D.
virginalis
Schltr.
(Wild
46240).
c:
D.
cooperi
Harv.
(Schelpe
7252).
d:
D.
togoen-
sis
Schltr.
(Scholes
20).
e:
D.
kerstenii
Rchb.
f.
(Eggeling
6634).
f:
D.
fanniniae
Harv.
(Kurzweil
1086).
Bar:
5
mm
in
a,
b,
d,
e;
4
mm
in
c;
7
mm
in
f.
The
lip
is
usually
extensively
fused
to
the
gynostemium,
which
is
often
quoted
as
a
diagnostic
character
of
the
whole
subtribe
Coryciinae
(=
Dis-
peridinae)
(SENGHAS,
1973
-
1974,
DRESSLER,
1981).
However,
in
some
species
the
lip
is
almost
or
completely
free
from
the
gynostemium
(see
also
VERDCOURT,
1975,
1978,
1988),
for
example
in
D.
kilimanjarica
(Fig.
15a,
c),
D.
villosa
(Fig.
19d)
and
D.
cucculata
(Fig.
19
a).
This
appears
to
be
a
derived
state
within
the
genus,
as
a
lip
being
fused
to
the
gynostemium
is
the
more
general
state
in
the
Coryciinae.
In
spite
of
the
extreme
diversity,
a
few
basic
lip
shapes
can
be
distin-
guished.
(1)
Virginalis
type:
In
many
tropical
species
the
lip
appendage
consists
of
two
apically
bilobed
processes
on
top
of
an
elongate
and
usually
bent
claw.
The
lip
blade
itself
is
a
frequently
short
lobe
just
below
the
inser-
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
461
LA
I
LA
LA
LB
I
L
B
;
.'
I
B
O
LA
,
I
O
LA
-
LB
LB
Fig.
21.
Lips
of
various
Disperis
species,
seen
from
the
front
(b,
c),
from
the
side
(a,
d,
e)
and
from
the
top
(f
-
g).
Lip
claw
not
shown
in
b.
a:
D.
cardiophora
Harv.
(Kurzweil
1250).
b:
D.
villosa
(L.
f.)
Sw.
(Kurzweil
1206).
c:
D.
micrantha
Lindl.
(Kurzweil
1111).
d:
D.
paludosa
Harv.
(Kurzweil
1027).
e:
D.
kerstenii
Rchb.
f.
(Eggeling
6634).
f:
D.
zeylanica
Trimen
(Barnes
s.n.).
g:
D.
katangensis
Summerh.
(Milne-Redhead
3699).
SEM
micrographs,
bars:
0.5
mm.
LB
LB
LA
462
H.
Kurzweil
and
H.
P.
Linder
tion
of
the
lip
appendage
and
is
mostly
hairy
at
its
base.
This
form
is
found
in
D.
thomensis
(Fig.
6
a,
d),
D.
virginalis
(Fig.
11
c),
D.
kerstenii
(Fig.
16,
17
a
-
b,
21
e),
D.
dicerochila,
D.
nemorosa
and
D.
aphylla.
In
D.
anthoceros
and
D.
bifida
the
lip
appendage
consists
of
two
pendent
fimbriate
lobes.
(2)
Katangensis
type:
Several
other
tropical
species
exhibit
a
basically
anchor-shaped
structure
consisting
of
two
diverging
lobes
on
top
of
a
short
reflexed
claw
(`LA?'
in
the
above
descriptions).
The
anchor-shaped
structure
is
not
borne
at
the
point
of
insertion
of
the
lip
blade
on
the
claw
but
well
below
it.
This
structure
may
be
homologous
with
the
lip
appendage,
and
the
flat
claw
below
the
lip
blade
(here
termed
secondary
claw)
would
then
be
an
additional
claw,
similar
to
that
found
in
many
other
Coryciinae
(KuRzwEii,
et
al.,
1991).
SUM1VIERHAYES
(1935),
however,
suggested
that
the
two-lobed
structure
may
well
be
due
to
the
broaden-
ing
and
splitting
of
the
lip
claw.
The
'secondary
claw'
would
then
be
a
continuation
of
the
basal
lip
claw.
In
this
interpretation
the
true
lip
appendage
is
represented
by
the
papillose crest
of
the
lip
blade
(VERD-
COURT,
1968a).
In
D.
katangensis
(Fig.
21
g)
and
D.
tripetaloidea
the
lobes
of
the
two-lobed
structure
are
rather
stout,
while
in
D.
zeylanica
(Fig.
21
f),
D.
reichenbachiana,
D.
johnstonii,
D.
togoensis
and
D.
leuconeura
they
are
narrow
and
horn-like.
D.
pusilla
may
also
be
refer-
red
to
this
morphological
type,
but
its
lip
blade
is
unstalked
and
nar-
rowly
triangular
throughout.
(3)
Wealei
type:
In
many
South
African
species
the
lip
blade
and
the
entire
lip
appendage
are
marginally
fused
resulting
in
a
cup-
or
boat-shaped
structure.
Two
forms
can
be
distinguished.
Disperis
wealei
(Fig.
10
a),
D.
circumflexa,
D.
stenoplectron,
D.
concinna,
D.
cooperi
and
D.
fanniniae
have
elongate
boat-shaped
lips
with
inflexed
or
reflexed
papillose
lip
appendage
and
upcurved
lip
blade.
In
some
species
the
lip
blade
forms
a
typical
mentum
or
sac
with
the
rest
of
the
lip
(particularly
in
D.
fanniniae).
Generally,
this
kind
of
lip
is
borne
on
a
very
short
claw.
In
some
Disperis
species
from
the
southwestern
Cape
of
South
Africa
(D.
villosa,
Fig.
21
b;
D.
cucculata;
D.
bolusiana;
D.
purpurata)
the
lip
is
very
distinct:
the
slender,
elongate
claw
is
more
or
less
free
from
the
gynostemium
and
bears
a
cup-
or
funnel-shaped
structure
with
a
papil
-
lose
dorsal
process.
This
'cup'
was
referred
to
as
`pseudopeltate'
(VoGEL,
1959).
(4)
Cardiophora
type:
This
is
perhaps
the
most
heterogenous
morphological
type
of
all.
The
lip
appendage
is
solid
and
entire
or
apically
bifid,
and
the
blade
is
a
short
to
elongate
triangular
process
pointing
forwards
or
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
463
downwards.
The
lip
claw
is
variable
in
shape,
but
is
mostly
massive,
straight
and
only
moderately
long
(except
in
D.
capensis
and
D.
uzungwae,
where
it
is
very
elongate
and
bent).
This
lip
form
is
found
in
D.
kilimanjarica
(Fig.
15c),
D.
capensis
(Fig.
18b),
D.
cardiophora
(Fig.
21
a),
D.
lindleyana
(Fig.
22
a),
D.
parviflora,
D.
uzungwae,
D.
bre-
viloba,
D.
woodii,
D.
stenoglossa,
D.
renibractea
and
D.
thorncroftii.
The
whole
of
the
lip
may
be
suberect
to
somewhat
reflexed
(here
shown
in
D.
lindleyana,
Fig.
22
a)
or
more
or
less
horizontally
reflexed
(D.
kilimanjarica,
Fig.
15
c;
D.
capensis,
Fig.
18b).
In
D.
macowanii
and
D.
bodkinii
the
lip
blade
is
clearly
concave.
D.
micrantha
(Fig.
21c)
and
D.
disaeformis,
with
the
claw
abruptly
widened
below
the
blade
and
with
an
apically
strongly
papillose
lip
appendage,
also
seem
to
belong
to
this
type.
LA
ROA
LB
AU
ROA
LCL
AN
RO
LA
LC
Fig.
22.
Disperis
lindleyana
Rchb.
f.
(Kurzweil
1235);
a:
gynostemium
and
lip.
b:
gynostemium
in
front
view.
c
-
e:
initiation
of
the
lip
appendage.
SEM
micrographs,
bars:
1
mm.
464
H.
Kurzweil
and
H.
P.
Linder
The
lip
of
D.
tysonii
has
a
short
claw
bearing
a
reflexed
elongate
and
straight
lip
appendage
which
is
not
fused
marginally
to
the
broadly
ovate
lip
blade.
This
is
either
referable
to
the
Wea/ei-type'
with
reduced
side-walls
of
the
'boat',
or
is
a
further
development
within
the
'Cardiophora-type'
with
a
strongly
developed
broad
blade.
D.
paludosa
has
a
very
unusual
lip
(Fig.
21
d).
It
is
basically
also
a
'boat',
but
this
'boat'
is,
in
contrast
to
that
found
in
the
Wea/ei-type',
apparently
exclusively
derived
from
the
lip
blade.
S
LB
Fig.
23.
Disperis
decipiens
Verd.,
flower
(Cribb
&
Grey-Wilson
10748A).
Bar:
1
mm.
Ontogenetic
data
on
the
lip
of
Disperis
species
are
very
difficult
to
obtain.
The
observations
made
in
a
single
species
(D.
lindleyana,
Fig.
22
c
-
e;
illustrating
the
two-lobed
lip
appendage
primordium)
are
certainly
not
rep-
resentative
of
the
whole
genus.
However,
the
result
is
interesting,
and
might
suggest
that
the
lip
appendage
is
basically
two-lobed
even
in
this
Disperis
species
with
entire
lip
appendage.
This
observation
together
with
several
previous
reports
that
the
lip
appendage
of
some
other
Coryciinae
is
dis
-
tinctly
bilobed
early
in
ontogeny
(VoGEL,
1959,
KuRzwEn.,
1991)
suggests
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
465
that
a
bilobed
lip
appendage
is
the
basic
condition
in
the
Coryciinae
(KURZWEIL
et
al.,
1991).
This
is
supported
by
the
fact
that
the
adult
lip
appendage
of
many
tropical
Disperis
species
is
deeply
two-lobed.
This
would
imply
that
the
deeply
bibbed
lip
appendage
of
these
tropical
species
is
an
ancestral
feature.
The
lip
appendage
(or
in
case
of
the
Katangensis-type
the
two-lobed
structure
possibly
homologous
with
the
split
lip
claw)
is
papillose
in
most
species.
In
the
South
African
species
the
glandular
trichomes
are
cylindrical
or
clavate
and
produce
oil
(STEINER,
1989)
which
may
well
be
a
synapomor-
phy
of
the
'South
African'
Glade
(see
below).
Such
cylindrical
or
clavate
glandular
trichomes
were
not
observed
in
any
tropical
species
in
the
present
study.
STEINER
(pers.
comm.)
suggests
that
the
tropical
Disperis
species
do
not
produce
oil.
In
D.
katangensis
(Fig.
4
b)
and
D.
zeylanica
(Fig.
8
b)
the
glands
on
the
large
two-lobed
'lip
appendage'
are
ovoid-globular
and
appar-
ently
sessile,
and
occur
together
with
minute
filiform
structures
(probably
not
artifacts!).
The
glands
on
top
of
the
lip
crest
of
D.
katangensis
(Fig.
4a)
are
apparently
very
similar
in
shape
to
those
on
the
lip
appendage'.
In
D.
thomensis
(Fig.
6
f)
the
glands
are
globular,
and
are
born
solitarily
on
stalks.
They
are
born
on
the
front
side
of
the
upper
arms
of
the
lip
appendage
(Fig.
6
a).
Interestingly,
similar
globular
glands
are
also
found
on
the
lip
blade
of
this
species.
However,
these
are
attached
to
trichomes
in
large
num-
bers
(Fig.
6
d
-
e).
In
D.
virginalis
(Fig.
12
a),
D.
anthoceros
(Fig.
14
a),
D.
kilimanjarica
(Fig.
15c)
and
D.
kerstenii
(Fig.
17
a,
21
e)
the
lip
appendage
appears
to
be
smooth.
While
the
only
existing
sectional
classification
of
Disperis
(SCHLECHTER,
1898)
relies
on
vegetative
characters
only,
the
lip
shape
appears
to
provide
better
characters
to
divide
the
genus
into
subgroups.
However,
it
has
to
be
emphasized
that
the
above
four
types
are
merely
morphological
types.
They
might
on
closer
examination
prove
to
be
accumulations
of
different
lip
shapes
and
therefore
do
not
necessarily
imply
phylogenetic
relationship.
More
detailed
and
comparative
studies
of
lips
of
all
Disperis
species
are
needed
in
order
to
assess
the
diversity
in
the
genus,
to
examine
the
system-
atic
relevance
and,
eventually,
to
contribute
to
a
sound
phylogeny
of
the
genus.
A
diagram
of
some
lip
types
showing
the
possible
evolution
of
lip
types
in
Disperis
is
given
in
Fig.
27.
Gynostemium
The
general
gynostemium
structure
is
rather
uniform.
The
gynostemium
lacks
a
prominent
stalk,
although
short
stalks
occur
occasionally
(D.
wealei,
Fig.
10
c;
D.
virginalis,
Fig.
11
c
-
d).
Usually
the
anther
is
horizontally
reflexed,
and
is
rarely
only
weakly
reflexed
(D.
kilimanjarica,
Fig.
15
c;
D.
466
H.
Kurzweil
and
H.
P.
Linder
cucculata,
Fig.
19
b;
D.
bolusiana)
or
suberect
(D.
macowanii;
see
Fig.
1
K
in
VERDCOURT,
1975).
The
anther
has
parallel
thecae
with
short
anther
canals.
In
contrast
to
all
other
Coryciinae,
the
connective
is
very
narrow
and
the
thecae
are
therefore
close
to
one
another.
The
thecae
remain
widely
open
after
the
removal
of
the
pollinia
(here
shown
in
D.
wealei,
Fig.
10d)
(see
also
VOGEL,
1959).
This
feature
is
very
unusual
in
Orchidoideae
and
may
serve
as
another
synapomorphy
for
Disperis.
The
anther
apex
is
mostly
more
or
less
rounded
or
slightly
emarginate
like
in
the
other
Diseae.
If
the
view
of
an
acute
anther
apex
being
the
ancestral
condition
of
the
Orchidoideae
(DRESS-
LER,
1990)
is
accepted
the
non-acute
anther
apex
of
most
Diseae
has
to
be
interpreted
as
a
derived
condition.
An
enormous
flat
central
rostellum
lobe
curves
over
and
around
the
anther.
The
lateral
gynostemium
appendages
of
Disperis
were
previously
referred
to
as
auricles,
and
have
been
commonly
interpreted
as
lateral
staminodes.
They
were
usually
described
as
simple,
sculptured
appendages
on
both
sides
of
the
gynostemium.
However,
on
closer
examination
they
clearly
consist
of
two
portions
in
most
species:
a
small
horizontal
portion,
and
an
elongate
erect
portion
which
is
fused
to
the
lip
base
(unless
the
lip
is
free).
The
width
of
the
erect
portion
varies
considerably,
and
therefore
this
portion
has
fre-
quently
not
been
detected
by
previous
workers.
The
point
of
confluence
of
the
two
portions
may
be
developed
as
a
±
sharp
angle
or
as
a
gentle
curve
when
seen
from
the
side.
While
the•
horizontal
portion
is
generally
sculptured,
the
erect
portion
may
be
smooth
(especially
in
the
tropical
species).
The
latter
was
described
and
illustrated
above
for
D.
katangensis
(Fig.
4
c),
D.
zeylanica
(Fig.
7d,
8a),
D.
lindleyana
(Fig.
22
b)
and
D.
johnstonii
(Fig.
24
b),
where
the
erect
portion
is
rather
prominent.
In
a
few
species
the
lateral
appendages
are
not clearly
differentiated
into
two
por-
tions
(particularly
in
D.
paludosa,
D.
cucculata),
and
are
throughout
sculptured
lobes.
This
frequent
bi-partite
appearance
of
the
lateral
gynostemium
append-
ages
is
suggestive
of
their
organophyletic
origin
from
both
auricles
and
basal
bulges
(as
defined
in
KURZWEIL,
1987),
which
would
correspond
to
the
situation
found
in
numerous
other
Orchidoideae.
However,
in
the
absence
of
detailed
ontogenetic
studies
this
derivation
of
the
appendages
from
two
dif-
ferent
primordia
is
hypothetical.
In
D.
fanniniae
the
lateral
gynostemium
appendages
originate
from
significant
primordia
(obviously
homologous
to
staminodes)
initiated
early
in
ontogeny
(KuRzwrIL,
1991).
But
it
is
not
clear
whether
the
appendages
of
the
mature
gynostemium
do
not
incorporate
auricular
tissue
as
well,
as
is
suggested
by
their
sculptured
surface.
The
erect
portions
have
occasionally
been
noted
before
in
some
species
(e.g.,
MANNING,
pers.
comm.),
but
no
information
on
their
systematic
dis-
tribution
in
the
genus
Disperis
is
available to
date.
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
467
ROA
AN
fr*
ROA
STG
AN
AU
0
ROA
AU
I
Fig.
24.
Disperis
johnstonii
Rolfe,
gynostemium
(Cole
s.n.);
a:
top
view.
b:
front
view.
c:
lateral
rostellum
arm.
d:
lateral
view.
SEM
micrograph,
bars:
0.3
mm.
Two
Disperis
species
have
very
unusual
lateral
gynostemium
appendages.
In
the
specimen
of
D.
macowanii
studied
here
(see
also
VERDCOURT,
1975)
the
lateral
gynostemium
appendages
are
horizontal
horn-like
processes
project-
ing
forward,
and
are
strongly
thickened
at
the
base.
Very
similar
but
shorter
horn-like,
undifferentiated
and
projecting
processes
are
also
found
in
D.
bodkinii.
The
rostellum
is
deeply
three-lobed
into
a
flat
central
rostellum
lobe
cov-
ering
the
anther
and
the
two
rostellum
arms.
The
flat
central
rostellum
lobe
is
the
most
unusual
gynostemium
character
of
the
genus,
and
serves
as
another
synapomorphy.
It
may
either
be
as
long
as
the
anther,
or
may
be
slightly
shorter
leaving
the
anther
apex
visible
from
above
(D.
katangensis,
Fig.
3
b;
D.
zeylanica,
Fig.
7c;
D.
togoensis).
In
D.
fanniniae
and
D.
bodkinii
the
apex
of
the
central
rostellum
lobe
is
slightly
recurved
behind
the
gynos-
31
Beitr.
Biol.
Pflanzen
66/3
468
H.
Kurzweil
and
H.
P.
Linder
temium,
and
the
margins
are
coarsely
serrate
in
these
two
species.
The
cen-
tral
rostellum
lobe
is
almost
always
closely
adpressed
to
the
gynostemium,
although
it
was
observed
to
be
elevated
well
above
the
latter
in
the
specimen
of
D.
macowanii
studied
here
(see
also
VERDCOURT,
1975,
Fig.
1K).
The
central
rostellum
lobe
generally
bears
the
stigmas,
which
are
ontogenetically
derived
from
the
median
carpel
(KuRzwEIL,
1991).
These
are
generally
situated
in
the
anterior
rostellum
portion
above
the
lip
insertion.
The
exact
position
of
the
stigmas
is
rather
diverse
in
the
genus:
(1)
In
those
species,
in
which
the
lip
is
not
or
only
weakly
fused
to
the
gynostemium,
the
stigma
is
a
±
entire
(but
sometimes
apically
emarginate),
round
and
slightly
convex
cushion
situated
on
the
front
base
of
the
gynos-
temium
(D.
kilimanjarica,
Fig.
15
a,
c
-
d;
D.
cucculata,
Fig.
19
b;
D.
villosa,
Fig.
19
d;
D.
bolusiana;
D.
bodkinii;
D.
macowanii).
(2)
In
most
tropical
Disperis
species
the
flat
stigmatic
area
is
situated
on
two
separate
lanceolate
stigmas
near
or
at
the
base
of
the
rostellum
arms
(here
shown
in
D.
katangensis,
Fig.
3
a
-
d;
D.
thomensis,
Fig.
6
a
-
c;
D.
zeylanica,
Fig.
7
c
-
d;
D.
virginalis,
Fig.
11
c
-
d;
D.
anthoceros,
Fig.
13
c
-
e;
D.
kerstenii,
Fig.
16,
17
c;
D.
johnstonii,
Fig.
24
a,
c
-
d).
(3)
In
many
southern
African
Disperis
species
two
stigmas
are
developed
as
large
usually
convex
cushions
on
top
of
the
central
rostellum
lobe,
and
are
frequently
united
in
the
middle.
In
D.
fanniniae
the
two
flat,
separate
stig-
mas
are
positioned
on
bulges
in
the
anterior
rostellum
portion
but
well
above
its
anterior
margin
(KuRzwEIL,
1991).
(4)
A
very
unusual
stigmatic
condition
is
found
in
D.
capensis
where
the
stigmas
are
situated
in
the
anterior
portion
of
the
central
rostellum
lobe
which
is
curled
in
front
of
the
gynostemium,
and
is
therefore
situated
in
front
of
the
lip
insertion
(Fig.
18
c;
see
also
Abb.
137
in
VOGEL,
1959).
In
many
tropical
species
the
anterior
margin
of
the
central
rostellum
lobe
is
developed
as
a
fleshy
ridge-like
tissue
(shown
in
D.
katangensis,
Fig.
3
b;
D.
virginalis,
Fig.
12
b;
D.
kerstenii,
Fig.
17
c;
D.
johnstonii,
Fig.
24
a),
which
is
confluent
with
the
two
median
teeth
on
the
central
rostellum
lobe
and
the
lateral
rostellum
arms.
The
two
median
teeth
adpressed
to
the
basal
part
of
the
rostellum
occur
in
most
species
and
appear
to
be
the
basic
condition
in
the
genus
Disperis.
They
were
previously
regarded
as
lip
excrescences
(VoGEL,
1959),
but
ontogenetic
evidence
suggests
that
they
are
vestiges
of
the
lateral
carpel
apices
(KuRzwEIL,
1991).
In
D.
fanniniae
these
teeth
are
two-lobed.
The
occurrence
of
three
such
teeth
in
a
few
specimens
(observed
in
D.
wealei,
Fig.
lob,
and
D.
fanniniae,
KURZWEIL,
1991)
may
well
be
an
abnormality
and
does
not
necessarily
question
the
interpretation
of
these
structures
as
lateral
carpel
apices,
as
three
lateral
carpel
apices
were
occasionally
also
observed
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
469
P
z
LA
LB
RO
STG
AU
RO
ROA
O
LC
Fig.
25.
Disperis
paludosa
Harv.
(Drewe
s.n.);
a:
flower
in
ventral
view.
b:
gynos-
temium
and
lip
in
lateral
view.
c:
gynostemium
in
top
view.
d:
lateral
rostellum
arm.
c
-
d:
SEM
micrographs;
bars:
5
mm
in
a
-
b,
1
mm
in
c
-
d.
in
young
stages
of
other
Orchidoideae
(KURZWEIL,
1985).
While
these
ves-
tiges
of
the
lateral
carpel
apices
are
rather
insignificant
in
most
species,
they
are
present
as
conspicuous
triangular
teeth
in
D.
paludosa
(Fig.
25
c
-
d)
and
D.
aphylla.
The
rostellum
arms
are
narrow
and
straight
in
most
tropical
species,
e.
g.
in
D.
katangensis
(Fig.
3
a
-
d),
D.
thomensis
(Fig.
6a
-
b),
D.
virginalis
(Fig.
11
c
-
d,
12
b),
D.
anthoceros
(Fig.
13
d
-
e),
D.
kerstenii
(Fig.
16,
17
c
-
d)
and
D.
johnstonii
(Fig.
24d).
In
almost
all
South
African
and
a
few
tropical
species
the
rostellum
arms
are
very
complicated
as
they
are
variously
twisted
and
contorted
(e.g.
D.
wealei,
Fig.
10
a
-
d;
D.
kilimanjarica,
Fig.
15
d;
D.
villosa,
Fig.
19
c;
D.
lindleyana,
Fig.
22
a
-
b;
and
D.
cooperi,
Fig.
26).
In
D.
cucculata
(Fig.
19
b),
D.
paludosa
(Fig.
25c
-
d)
and
D.
uzungwae
the
rostellum
arms
are
intermediate
between
these
two
condi-
tions,
i.e.
they
are
semi-complicated.
31'
470
H.
Kurzweil
and
H.
P.
Linder
B)
Phylogenetic
considerations
The
genus
Disperis
is
usually
placed
as
a
distinct
genus
in
the
Coryciinae,
and
was
regarded
as
a
sister
group
of
the
`Coryciinae
s.
s.'
(=
Ceratandra,
Evotella,
Pterygodium,
Corycium)
by
KURZWEIL
et
al.
(1991).
The
unique
flower
and
gynostemium
structure
(lateral
sepal
spurs,
linear
lip
with
claw,
flat
rostellum
covering
the
anther,
elongate
and
projecting
rostellum
arms)
leave
no
doubt
regarding
the
monophyly
of
the
genus
Disperis.
The
infrageneric
classification
of
the
genus
Disperis
was
little
disputed
in
the
past,
as
only
one
classification
is
available
(SCHLECHTER,
1898).
SCHLECH-
TER
distinguished
two
sections
on
the
basis
of
the
arrangement
of
foliar
leaves
but
this
classification
is
largely
artificial
(SCHLECHTER,
1898).
The
recognition
of
two
subgroups
purely
on
the
basis
of
vegetative
characters
appears
not
to
be
justified
as
both
sections
are
very
heterogenous
in
their
floral
morphology.
Furthermore,
ScHmEcHTER's
classification
lists
only
the
40
species
then
known
(out
of
approx.
90
known
at
present).
Two
distinct
groups,
on
the
basis
of
the
floral
morphology
of
the
41
species
studied,
can
be
recognized
(see
also
table
1).
This
grouping
corresponds
well
to
what
was
postulated
by
MANNING
&
LINDER
(1992).
(1)
The
first
group
comprises
the
majority
of
species
in
the
genus,
and
was
referred
to
as
the
'tropical
group'
(MANNING
&
LINDER,
1992)
since
most
species
occur
in
the
tropics
although
three
species
spread
into
the
subtropi-
cal
parts
of
South
Africa
(D.
virginalis,
D.
anthoceros,
D.
johnstonii).
The
pollen
surface
was
given
as
a
synapomorphy
for
the
group
(MANNING
&
LIN-
DER,
1992),
but
the
species
in
this
group
also
share
the
simple
rostellum
Fig.
26.
Disperis
cooperi
Harv.
(Schelpe
7252);
lateral
rostellum
arm.
SEM
graph,
bar:
1
mm.
mic
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis 471
m
d
Fig.
27.
Hypothetical
diagram
of
lip
types
in
Disperis.
a
-
c:
Wealei
type.
a:
D.
wealei.
b:
D.
bolusiana.
c:
D.
villosa.
d
-
h:
Cardiophora
type.
d:
D.
cardiophora.
e:
D.
woodii.
f:
D.
micrantha.
g:
D.
lindleyana.
h:
D.
macowanii.
—i
-
j:
Virginalis
type.
i:
D.
virginalis.
j:
D.
anthoceros.
k
-
m:
Katangensis
type.
k:
D.
katangensis.
1:
D.
zeylanica.
m:
D.
pusilla.
472
H.
Kurzweil
and
H.
P.
Linder
arms
and
the
lanceolate
stigmas
near
or
at
the
base
of
the
rostellum
arms.
However,
simple
rostellum
arms
are
primitive
in
the
Orchidoideae,
and
two
separate
stigmas
are
obviously
the
basic
condition
in
the
entire
subtribe
Coryciinae.
Therefore,
both
characters
may
be
plesiomorphic
in
the
genus.
The
'lip
appendage'
is
deeply
bilobed,
but
is
possibly
not
homologous
within
the
group
(i.e.
compare
the
`Katangensis-type'
and
the
Virg/net/is-type).
On
the
basis
of
the
studied
species
two
subgroups
can
be
recognized.
a)
The
lip
structure
of
the
Virginalis-type'
might
be
regarded
as
a
synapomor-
phy
of
the
Disperis
virginalis
alliance.
However,
the
character
state
may
also
be
plesiomorphic,
as
a
two-lobed
lip
appendage
is
considered
to
be
the
basic
condition
of
all
Coryciinae.
This
may
be
a
morphological
homogenous
paraphyletic
group
close
to
the
ancestral
condition
of
Disperis.
b)
If
the
interpretation
of
the
lip
of
the
Katangensis-type'
made
by
SUMIVIERHAYES
(1935)
and
VERDCOURT
(1968
a)
is
accepted,
the
true
lip
appendage
of
this
type
would
be
represented
by
the
papillose
but
unlobed
crest
on
the
lip
blade,
while
the
enormous
bibbed
structure
is
a
split
and
enlarged
part
of
the
claw.
This
lip
crest
together
with
the
split
claw
may
serve
as
synapomor-
phies
of
the
group
referrable
as
Katangensis
Glade'.
(2)
The
'South
African
group'
comprises
almost
all
of
the
South
African
species
as
well
as
a
few
tropical
species.
All
species
belonging
to
the
group
share
the
entire
lip
appendage.
This
is
obviously
a
derived
state
as
outgroup
comparison
with
the
Coryciinae
s. s.
suggests
that
the
two-lobed
lip
appen-
dage
is
primitive,
and
consequently
this
group
can
be
regarded
as
monophyletic.
The
strongly
twisted
rostellum
arms
may
also
be
a
synapomorphy
for
the
group,
as
untwisted
rostellum
arms
are
certainly
the
primitive
condition
in
the
Orchidoideae.
The
few
cases
of
only
weakly
twisted
rostellum
arms
found
in
the
'South
African
group'
(D.
cucculata,
D.
paludosa,
D.
uzungwe)
may
well
be
due
to
secondary
loss
of
twisting.
The
South
African
group
is
extraordinary
heterogenous,
and
apparently
has
diversified
much
more
than
the
'tropical
group'.
A
phylogeny
of
the
Glade
was
presented
by
MANNING
&
LINDER
(1992).
It
is
interesting
that
the
species
of
the
'South
African
group'
have
retained
a
few
possibly
primitive
features
in
their
floral
morphology.
Alternatively,
these
may
also
be
due
to
secondary
developments.
(1)
Unlike
in
the
'tropical
group'
and
unlike
in
all
other
Coryciinae,
the
petals
of
the
'South
African
group'
are
often
rather
simple
and
almost
symmetrical
structures.
(2)
The
fusion
of
the
lateral
sepals
is
missing
in
most
species.
(3)
The
species
are
also
linked
by
the
incomplete
or
lacking
separation
of
the
two
stigmas.
This
is
probably
an
ancestral
feature,
as
one
entire
stigma
is
certainly
primitive
in
orchids
with
a
stigma
on
the
median
carpel.
However,
two
separate
stigmas
have
evolved
in
the
'tropical
group'
of
Disperis
species,
and
also
in
the
rest
of
the
Coryciinae
(KuRzwEIL
et
al.,
1991).
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
473
Conclusions
(a)
Detailed
studies
show
a
wide
range
of
characters
than
can
be
used
for
phylogeny.
(b)
Comparative
data
for
more
species
are
needed
for
a
plausible
phylogenetic
analysis.
(c)
SCHLECHTER'S
infrageneric
classification
of
the
genus
is
not
based
on
natural
groupings,
and
deeply
contrasts
a
phylogeny
based
on
details
of
floral
structures
(present
paper)
as
well
as
on
pollen
data
(MANNING
&
LIN-
DER,
1992).
Acknowledgements
The
study
was
funded
by
FRD
and
a
Smuts
Memorial
Fellowship.
We
express
our
sincere
thanks
to
the
staff
of
the
Electron
Microscope
Unit
of
the
University
of
Cape
Town
for
supplying
SEM-
and
darkroom-time
and
for
technical
assistance,
and
to
the
staff
of
the
Herbarium,
Kew
/
England,
and
the
Bolus
Herbarium,
University
of
Cape
Town,
for
providing
alcohol
material
for
study.
Thanks
are
also
due
to
the
Cape
Town
City
Council,
the
Cape
Department
of
Nature
and
Environment
Conservation
and
the
Natal
Parks
Board
for
issuing
the
necessary
collecting
permits.
Collecting
was
helped
by
Mrs.
P.
DREWE,
Prof.
P.
JACKSON,
Dr.
JACOT-GUILLARMOD
and
Dr.
J.
MANNING.
We
are
also
grateful
to
Ms.
B.
LOURENS
for
making
the
line
drawings.
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603
-
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of
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from
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9
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51
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57
(1988).
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S.:
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der
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Appendix:
Specimens
examined
The
specimens
studied
are
listed
below
in
alphabetical
order.
The
source
of
the
'
specimens
is
given
in
brackets
(K
=
The
Herbarium,
Royal
Botanic
Gardens,
Kew;
BOL
=
Bolus
Herbarium,
University
of
Cape
Town).
Material
collected
by
us
was
identified
according
to
STEWART
et
al.
(1982).
In
all
other
specimens
we
have
trusted
the
identifications
of
the
staff
of
Kew
and
the
Bolus
Herbarium,
respectively.
The
nomenclature
follows
VERDCOURT
(1988)
and
STEWART
et
al.
(1982)
unless
otherwise
stated.
Bol,
indicates
the
number
in
the
pickled
collection
of
the
Bolus
Herbarium.
Disperis
anthoceros
Rchb.
f.,
Kenya:
Tweedie
2852
(K);
Eggeling
5949
(K);
Zambia:
G.
Williamson
285
(K).
D.
aphylla
Krzl.,
Zambia:
Schaijes
2839
A
(K).
D.
bifida
Cribbl,
Malawi:
Dowsett-Lemaire
348
(K).
D.
bodkinii
Bol.,
South
Africa:
Kurzweil
1205
(BOL);
South
Africa:
Kurzweil
960
(BOL).
D.
bolusiana
Schltr.,
without
loc.
and
collector
(BOL).
D.
breviloba
Verdc.
2
,
Zambia:
Williamson,
Ball
&
Simon
362
(K).
D.
capensis
Sw.,
South
Africa:
Kurzweil
901
(BOL);
South
Africa:
Hall
701
(BOL);
South
Africa:
Kurzweil
909
(BOL).
476
H.
Kurzweil
and
H.
P.
Linder
D.
cardiophora
Harv.,
South
Africa:
Kurzweil
1250
(BOL);
South
Africa:
Kurzweil
1431
(BOL).
D.
circumflexa
(L.)
Dur.
&
Schinz,
South
Africa:
Kurzweil
1207
(BOL);
Bol
t
,
1323
(BOL);
Bol
t
,s.n.
(BOL).
D.
circumflexa
(L.)
Dur.
&
Schinz
var
aemula
Schltr.,
Oliver
s.n.,
Bol
e
1278
(BOL).
D.
cooperi
Harv.,
South
Africa:
Schelpe
7252
(BOL);
South
Africa:
Schelpe
6329
(K).
D.
cucculata
Sw.,
South
Africa:
Linder
1137
(BOL);
Bol
e
s.n.
(BOL).
D.
decipiens
Verdc.,
Tanzania:
Cribb
&
Grey-Wilson
10748A
(K);
Tanzania:
Leedal
11250
(K).
D.
dicerochila
Summerh.,
East
Africa:
Tweedie
2691
(K);
Ruanda:
Bridson
419
(K);
Tanzania:
Polhill
&
S.
Paulo
15308
(K).
D.
elaphoceras
Verde.,
Tanzania:
Thomas
3682a
(K).
D.
fanniniae
Harv.,
South
Africa:
Kurzweil
1086
(BOL);
South
Africa:
Kurzweil
1422
(BOL).
D.
johnstonii
Rolfe,
Nigeria:
Cole
s.n.
(K);
Tanzania:
Eggeling
6629
(K);
Bol
t
,
144
(BOL).
D.
katangensis
Summerh.,
Zimbabwe:
Milne-Redhead
3699
(K);
Zimbabwe:
Richards
10841
(K).
D.
kerstenii
Rchb.
f.,
Mabberley
1206
(K);
Tanzania:
Eggeling
6634 (K).
D.
kilimanjarica
Rendle,
Kenya:
Mabberley
&
McCall
13
(K);
Malawi:
Dowsett-
Lemaire
352
(K).
D.
leuconeura
Schltr.,
McLaughin
106
(K).
D.
lindleyana
Rchb.
f.,
South
Africa:
Hall
609
(BOL);
Transkei:
Kurzweil
1235
(BOL);
South
Africa:
Linder
790
(BOL).
D.
macowanii
Bol.,
Tanzania:
Cribb
&
Grey
Wilson
10748
(K).
D.
micrantha
Lind1.,
South
Africa:
Kurzweil
1111
(BOL).
D.
nemorosa
Rendle,
Malawi:
Dowsett-Lemaire
351
(K).
D.
oppositifolia
Sw.
3
,
Mauritius:
Braclay
375
(K).
D.
oxyglossa
Bol.,
South
Africa:
Trauseld
1052
(BOL).
D.
paludosa
Harv.,
South
Africa:
Kurzweil
1027
(BOL);
South
Africa:
Drewe
s.n.
(BOL).
D.
purpurata
Rchb.
f.,
Bol
t
,
135
(BOL).
D.
pusilla
Verc.
4
,
Zimbabwe:
Richards
17074
(K).
D.
reichenbachiana
Rchb.
f.,
Milne-Redhead
4295
(K);
Tanzania:
Polhill
&
Paulo
1467
(K);
Tanzania:
Cribb
&
Grey-Wilson
10673
(K).
D.
thomensis
Summerh.,
Angola:
Milne-Redhead
4227
(K);
Sierra
Leone:
Jaeger
1289
(K).
D.
thorncroftii
Schltr.,
South
Africa:
Trauseld
1028
(BOL).
D.
tog
oensis
Schltr.,
Togo:
Scholes
20
(K).
D.
tripetaloidea
(Thou.)
Ld1.
3
,
Seychelles:
Jeffrey
407
(K).
D.
tysonii
Bol.,
South
Africa:
Kurzweil
1098
(BOL);
South
Africa:
Linder
4665
(BOL)•
A
comparative
study
of
the
floral
morphology
in
the
genus
Disperis
477
D.
uzungwae
Verde.,
Tanzania:
Lovett
270
(K).
D.
villosa
(L.
f.)
Sw.;
South
Africa:
Kurzweil
1206
(BOL);
South
Africa:
Bol
s.n.
(BOL).
D.
virginalis
Schltr.,
Zimbabwe:
Wild
2810
(K);
Zimbabwe:
S.
G.
H.
114,
239
(K);
Zimbabwe:
Wild
46240
(K).
D.
wealei
Rchb.
f.,
South
Africa:
Linder
2088
(BOL).
D.
woodii
Bol.,
South
Africa:
Batten
s.n.
(BOL).
D.
zeylanica
Trimen
5
,
Sri
Lanka:
Barnes
s.n.
(K).
nomenclature
according
to
CRIBB
&
STEWART
(1985),
2
(VERDCOURT
1978),
3
(SCHLECHTER
1925),
4
(VERDCOURT
1968b),
5
(JAYAWEERA
1981).
Authors'
address:
Bolus
Herbarium
University
of
Cape
Town
7700
Rondebosch
South
Africa
H.
KURZWEIL*
and
H.
P.
LINDER
*
current
address:
Compton
Herbarium,
National
Botanical
Institute
Private
Bag
X7
Claremont
7735
South
Africa