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Botanical
Journal
ofthe Linnean
Socie&
(1988),
97:
357403. With 66 figures
A
taxonomic revision
of
the genus
Tordyliurn
L.
(
Apiaceae)
DAWUD AL-EISAWI F.L.S. AND STEPHEN L.
JURY
F.L.S.
Department
of
Botany, Plant Science Laboratories, University
of
Reading,
P.
0.
Box 221,
Whiteknights, Reading, Berkshire,
RG6
2AS
Received October,
1985,
accepted
for
publication February,
1987
AL-EISAWI, DAWUD
&JURY,
STEPHEN, L., 1988.
A
taxonomic
revision
of
the
genus
Tordylium
L.
(Apiaceae).
The
four
related genera:
Ainsworthia
Boiss.,
Tordylium
L.,
Synelcosciadium
Boiss. and
Mandenovia
Alava are revised. Data are presented from detailed gross morphology,
mericarp surface features and anatomy, and palynology.
The results show:
(1)
Ainsmorthia
and
Synelcosciadium
are congeneric with the genus
Tordylium,
and
that
Mandenovia
is a good monotypic genus;
(2)
Tordylium persicum
is synonymous with
7.
cappadocicum
and
7.
aegaeum
with
7.
pestalozzae;
(3) the genus
Tordylium
is best divided into
subgenus
Tordylium
(including
Synelcosciadium)
and subgenus
Ainsworthia
(Boiss.) Drude. The latter is
divided into section
Condylocarpus
(Hoffm.) DC, section
Hasselquistia
(L.)
Boiss. and section
Univittata
Drude.
A new species
of
Tordylium
is described, and two new combinations made.
A
taxonomic treatment
of
the genus
'Tordylium,
together with a key to the species, is given.
ADDITIONAL KEY
WORDS:-Aimwortha
-
anatomy
-
Mandenouia
-
mericarps
-
palynology
-
scanning electron microscopy
-
Synelcosciadium
-
systematics
-
Umbelliferae.
CONTENTS
Introduction
.
. .
.
. .
.
. .
.
.
.
.
.
.
. . . .
358
Historical review
.
.
.
. .
.
. .
. .
. . .
.
.
. . .
358
Morphological characters
.
. .
.
. .
. .
.
.
.
. . . . .
361
Habit and duration
.
.
.
. . .
. .
. . . . . . . .
361
Stem and indumentum
. .
.
.
.
. .
.
.
.
.
. .
.
.
361
Leaves
. .
.
. . . . .
.
. .
.
.
.
. .
.
.
.
361
Bracts
.
.
.
. .
.
.
.
.
. .
.
.
.
. .
.
. .
363
Bracteoles.
. . . .
.
. .
.
. . . . . . .
. .
.
366
Sepals
. . . .
.
.
.
.
.
. .
.
.
. .
. .
. .
368
Petals.
. .
. . . . . .
.
. . .
.
. .
.
. . .
368
Style and stylopodium.
. .
. .
.
. .
.
.
. .
.
.
.
.
370
Umbels
. .
. . . .
. .
.
. .
.
.
.
.
.
. . .
370
Fruits.
.
. . . . .
. .
. . .
.
. .
.
. . . .
371
Vittae
.
.
.
. . .
.
.
. .
. . .
.
.
. .
.
.
371
Scanning electron microscopy
of
mericarps
. . . . .
.
. . . .
.
.
37
1
Mericarp margin
. . . . . .
.
. . .
.
.
. .
.
. .
372
Hairs.
. . .
.
.
. .
. .
.
.
.
. .
. .
. . .
379
Style and stylopodium.
.
.
. .
.
.
. . .
. .
.
. . .
379
Calyx-teeth
.
.
. . . . .
.
.
. . . . .
.
. . .
379
Results
. . .
.
. . . .
.
.
. . . . . .
.
.
.
380
357
0024-4074/88/080357 +47 $03.00/0
0
1988 The Linnean Society of London
358
DAWUD AL-EISAWI AND
S.
L.
JURY
Comparative fruit anatomy
...............
380
Wings
...................
380
Vascular bundles
.................
381
Number and position
of
the vittae
.............
381
Shape and size of the vittae
..............
381
Endocarp.
..................
38
1
Mesocarp.
..................
381
Pollen morphology.
.................
381
Description
..................
382
Discussion and conclusions.
..............
386
Cytology
....................
386
Taxonomic treatment
.................
388
Tordylium
L.
..................
388
Synopsis of taxa
.................
388
Key
to the species.
................
389
Species excluded from
Tordyliurn
L.
............
402
Appendix: index to species
................
402
Acknowledgements
.................
402
References.
...................
403
INTRODUCTION
This study arose out of research on the Apiaceae of Jordan. The generic
delimitation of the genus
Tordylium
L. was found to be somewhat confused. To
revise this genus it was necessary to study the allied genera
Ainsworthia
Boiss.,
Synelcosciadium
Boiss. and
Mandenovia
Alava. The centre of distribution
of
all
these genera is Western Asia and the Mediterranean.
The earlier Flora writers covering parts of this region have treated these
genera very differently, as is outlined below. Previous classifications were
wholely based on morphological characters, such
as:
whether the mericarp
margin was smooth or corrugated, the presence or absence of calyx-teeth, the
number and shape of bracts, and whether the petals were hairy or glabrous. In
this study, additional information from scanning electron microscopy of fruits
and pollen grains, anatomy of mericarps and cytology has been obtained and
found to be important in delimiting the genera.
HISTORICAL REVIEW
Linnaeus
(1
753)
recognized seven species of
Tordylium
as follows:
7.
syriucum,
T.
oficinale,
T.
apulum,
T,
maximum,
T.
latiflium,
1.
anthriscus,
and
T.
nodosum.
The last three species are recognized today respectively as:
Turgenia lutzfolia
(L.)
Hoffm.,
Torilis japonica
(Houtt.) DC. and
T.
nodosa
(L.) Gaertner. Two years
later Linnaeus
(1
755),
described a new genus
Hasselquistia,
related to the genus
Tordylium,
but differing from it by its dimorphic fruits, with the sole species
H.
aegyptiaca.
Jacquin
( 1772),
described another species,
Hasselquistia cordala,
on the basis of
dimorphic fruits. This genus was not recognized
by
Lamarck (1792), who
transferred
Hasselquistia aegyfitiaca
to
Tordylium,
or by Poiret
(
1806),
who
transferred Jacquin’s
H. cordata
also to
Tordylium.
Other species were described, such as
Tordylium magnum
Brot.,
T.
grandijorum
Moench, and
T,
humile
Desf., all now reduced to synonymy.
Hoffmann
(
18
16), split
off
Tordylium apulum
as
Condylocarpus apulus,
a
monotypic
genus on the basis of the fruits having many vittae and the petals equal lobes.
This treatment was not accepted by De Candolle (1830), who again recognized
REVISION
OF
THE
GENUS
'TORDYLUM
359
the genus
Hasselquistia,
although he stated this separation was difficult, and
divided the genus
Tordylium
into two sections.
Labillardi2re
(
1812), described a new species of
Heracleum,
H.
carmeli
from
Syria, but this was later transferred by Boissier (1844) to a new genus
Synelcosciadium
Boiss. Boissier mentioned that
Synelcosciadium carmeli
(Labil.) Boiss.
has the general appearance of
a
Tordylium.
Boissier (1844) also split Jacquin's
Hasselquistia cordata
into a new genus
Ainsworthia,
differing from
Hasselquistia
and
Tordylium
by the absence of calyx-teeth and the non-rugose fruit margin.
He
also
described a new species of
Hasselquistia, H. lanata,
which he said was closely
related to
T.
aegyptiacum,
this he later transferred in his
Flora
Orientalis
(1872) to
Tordylium
as
T.
lanaturn.
He described four new species of
Tordylium:
1.
macropetalum,
T,
cappadocicum,
T.
pestalozzae
and
T.
brachytaenium,
as well as
three other species of
Ainsworthia: A. trachycarpa,
with uniform fruits, and
A.
carmeli,
with dimorphic fruits, and yet another species
A.
elegans
(Boissier,
1856). These last species have recently been treated by Alava (1971) in
preparation for his
Flora
of
Turkey
account (1972): the first as synonymous with
A.
cordata
and the last as
Tordylium elegans
(Boiss.) Alava.
Boissier in
Flora Orientalis
(
1872), recognized three genera
Ainsworthia,
Tordylium
and
Synelcosciadium,
considering the genera
Hasselquistia
and
Condylocarpus
congeneric with
Tordylium.
He divided
Tordylium
into three
sections,
Hasselquistia
(L.)
Boiss,
Eutordylium
and
Condylocarpus
(Hoffmann) DC.
He considered the genus
Ainsworthia
to have four species:
A. cordata
(Jacq.)
Boiss.,
A. trachycarpa
Boiss.,
A. carmeli
Boiss. and
A. elegans
Boiss.
&
Bal.
The genus
Synelcosciadium
was also still considered to be monotypic and
intermediate between
Heracleum
and
Tordylium,
being placed immediately after
Heracleum.
Boissier in the supplement
to
the
Flora Orientalis
(1888), described the new
species
Tordylium pustulosum
which was referred by Post
to
T.
hasselquistiae.
Boissier distinguished
it
from
T.
hasselquistiae
by the cauline leaves having
cuspidate leaflets and the mericarps being smaller in size.
Bentham
&
Hooker (1867), simply lumped all these genera
(Tordylium
Hasselquistia, Ainsworthia, Condylocarpus
and
Synelcosciadium)
,
recognizing only
Tordylium.
Candargy (1897) described a new species of
Tordylium,
T.
hirtocarpurn,
in
section
Eutordylium.
In his description he said that his species differed from
1.
persicum
by the form of the leaves, fruit hairs and the commissural face, and
from
T.
cappadocicum
by the fruit pedicels and shorter rays.
Drude in Engler
&
Prantl,
Die Naturlichen PfEanzenfamilien
(
1898) divided
Tordylium
into two subgenera, based on whether the fruit margin was corrugated
or not. He divided the subgenera into sections, according to the number of
vittae in each furrow as follows:
Subgenus
I
Tordyliastrum
Drude.
Section
I
Plurivittata
Drude
(T.
apulum
only).
Section
I1
Univittata
Drude.
Subgenus
I1
Ainsworthia
(Boiss.) Drude.
Halacsy, in his
Florae Graecae
(1901), gave three species in the genus
Tordylium,
but recognized a new variety
of
T.
apulum
var.
heterocarpum
Heldr. based on the
different sizes of the mericarps.
360
DAWUD AL-EISAWI
AND
S.
L.
JURY
Holmboe, in
Studies on the Vegetation
of
Cyprus
(1914), treated
Ainsworthia
as
synonymous with
Tordylium,
with
A.
cordata
as
T.
cordatum,
and
A.
trachycarpa
as
a subspecies,
T.
cordatum
subsp.
trachycarpum.
Thellung, in Hegi
Illustrierte Flora von Mitteleuropa
(
1926), used
De
Candolle’s
classification by recognizing two sections as follows:
1.
Section
Eutordylium
DC.
(=
Tordylium
Hoffmann
=
subgenus
Tordyliastrum
2.
Section
Condylocarpus
(
=
subgenus
Tordyliastrum
Drude Section
Plurivittata
Drude Section
Univittata
Drude)
.
Drude).
Aznavore, in
Flore
de Constantinople
(
1897), described
Ainsworthia byzantina
from
Istanbul as a new species. This species was later treated by Alava (1971) as
synonymous with
A.
trachycarpa.
Hayek, in
Prodromus Florae Peninsulae Balcanica
(
1927), considered
Ainsworthia
byeantina
as
a
species of
Tordylium,
T,
byzantinum
(Azn.) Hayek.
Zohary (1941) described a new species of
Tordylium,
T.
palaestinum.
He
said
that his new species differed from
T.
aegyptiacum
by the smaller size of its
mericarps and the shorter rays. Later, in
Flora Palaestina
(1972), he changed the
species to a variety,
T.
aegyptiacum
var.
palaestinum.
In the same Flora he treated
Ainsworthia
and
Tordylium
as separate genera.
Mandenova described the new species
Tordylium komarovii,
recently transferred
to a new monotypic genus by Alava (1970) to become
Mandenouia komarovii
(Mandenova) Alava.
Dinsmore, in Post
Flora
of
Syria, Palestine and Sinai
(
1932), treated
Ainsworthia
and
Tordylium
as two separate
genera.
Schischkin, in his account of
Tordylium
in the
Flora
of
U.S.S.R.
(1951), gave
two species:
T.
komarovii
and
T.
maximum;
the latter with a figure (Vol. 17: 267,
fig.
1)
showing two types of fruits, one with a smooth margin and the other with
a corrugated-strigose margin.
Runemark, in
Studies in the Aegean
Flora
(1968), recognized six species,
one
of
them,
7.
aegaeum
being a new species differing from
T.
pestalotzae
Boiss. by the
subulate bracteoles and the many unequal rays.
Townsend (1968) treated the genus
Ainsworthia
as
a
subgenus of
Tordylium.
Mouterde, in
Nouvelle Flore du Liban et de
la
Syrie
(1970), treated
Ainsworthia
as
The genus Ainsworthia was last revised by Alava (1971) who considered it a
(1)
The different number and shape of involucral bracts and bracteoles.
(2) Petals glabrous in
Ainsworthia
and not in
Tordylium.
(3)
Calyx-teeth obsolete in
Ainsworthia
and present in
Tor@ium.
(4) Mericarp margins in
Ainsworthia
are not moniliform.
In his revision he excluded the species
Ainsworthia elegans,
placing it in the
genus
Tor@ium,
without explanation; and considered
Ainsworthia carmeli
a
synonym
of
A.
cordata.
Alava in Davis’
Flora
of
Turkey
(1974), recognized 15 species of
Tordylium,
but
does not give any subgenera or sections.
a
separate genus from
Tordylium.
good genus, separable from
Tordylium
by the following reasons:
REVISION
OF
THE GENUS
TORDYLUM
361
MORPHOLOGICAL CHARACTERS
Habit and duration
All species of
Tordylium, Ainsworthia
and
Qnelcosciadium
are annual herbs,
except
T.
maximum
which is sometimes biennial.
Mandenovia
is a perennial herb
with tuberous roots and stem branching from the base. It differs from all the
species of the other genera mentioned; these have tap roots and are usually
single-stemmed, branching dichotomously above.
The height of the plants varies according to the ecological conditions, but
usually the drier the conditions the smaller the plants. This is particularly
evident in
A.
trachycarpa
which, when collected from dry places, never reaches
more than 300 mm, but in less extreme habitats often grows to
800
mm. In
general, for most species, the height of the plants ranges 150 to 500 mm, but in
A.
trachycarpa,
T.
maximum
and
S.
carmeli
they usually reach
800
mm or more.
The flowering period for most of the species is between March and early May,
except for
T.
maximum,
T.
macropetalum
and
S.
carmeli
which flower between May
and early July, and
Mandenovia komarovii
July-August.
Stem and indumentum
The stem in all species is erect, terete, finely striate to ridged, with few to
many branches.
The stem indumentum in most of the species is pubescent
to
scabrous,
sometimes hispid at the base, but in
T.
aegyptiacum
and
T.
lanatum
villous to
villous-scabrous; retrosely strigose to setose in
T.
maximum,
T.
macropetalum
and
Synelcosciadium carneli;
and glabrous to sparsely papillate in
M.
komarovii.
Leaves
(Figs 1-5)
The leaves are alternate, usually distinctly petiolate, the bases of the petioles
have small sheaths; the margins are membranous or membranous-ciliate.
The basal leaves are simple or pinnate, usually with long petioles. The simple
leaves are cordate to ovate with a crenate margin. The leaves are usually
1-
pinnate, with two to six pairs of leaflets; the terminal segments are always
larger, ovate to cordate, with crenate to serrate margins, but in
T.
aeryptiacum
and
T.
lanatum
(Fig.
1)
the leaves are
2-
to 3-pinnate with ovate-oblong
segments, the margins entire to dentate. The cauline leaves are sessile, usually
pinnate; segments are ovate
to
oblong, with crenate to serrate margins, the
leaflet tips are sometimes cuspidate, for example
T.
pustulosum
(Fig.
2)
and
T.
elegans
(Fig. 4). The upper leaves are sessile, pinnate or 3-lobed, rarely
simple, the terminal lobes lanceolate to linear, with margins entire or coarsely
serrate, for example
T.
Maximum
and
T.
macropetalum
(Fig. 5).
In the species
A. trachycarpa
(Fig.
4),
the size, shape and dissection of the
leaves and leaf-segments is very much affected by the environment: often one
finds plants all with simple leaves or with pinnate to deeply dissected leaves. The
leaflet size range is 15-40
x
15-60 mm.
362
DAWUD AL-EISAWI AND
S.
L.
JURY
T.
aegyptiacum
\"
T.
lanotum
-
cm
A
T:
syriarum
Figure
1.
Leaf silhouettes of five species
lettered up the plant.
Tordylium
aegypliacurn
I
T;
apulum
T;
hasselquistioe
of
Tordylium;
basal leaves labelled
A
and
sequc
and
7.
lanaturn
have
2-
to 3-pinnate leaves.
mtially
REVISION
OF
THE
GENUS
TORDYLUM
363
I
crn
-
T.
persicom
4
'H
G
+*
T.
cappadocicum
8%
T.
elego
ns
Figure
2.
Leaf silhouettes of three
species
of
Tordylium:
basal leaves labelled
A
and sequentially
lettered
up
the plant.
The leaves in
M.
komarovii
(Fig.
5)
are small, with thin, long petioles and a
pinnate lamina; segments
1-2
x
5-8
mm, oblong-lanceolate, with entire
margins, quite different from all leaf types (Figs
1-5).
Most species have leaves with soft, appressed hairs on both sides, some with
villous hairs, for example
T.
maximum,
T.
macropetalum
and
Synelcosciadium carmeli.
Bracts
(Figs
6-9)
The number and shape of the bracts varies from one species to another. In
general, the number ranges from
0
to
18
and the shape varies from filiform to
filiform-linear, linear-lanceolate, lanceolate or
oblanceolate-spathulate.
Bracts are always absent in
M.
komarovii
(Fig.
9)
and in the new species
described later Dudley D354
77
(Fig.
8);
one to five bracts are found in
T.
aegyptiacum,
T.
lanatum,
T.
apulum
and
T.
hirtocarpum;
always five in
T.
syriacum;
five to eight in
T.
pestaloaae,
T.
brachytaenium,
T.
persicum,
T.
cappadocicum,
T,
maximum,
7.
macropetalum,
and
S.
carmeli;
eight to
14
in
T.
pustulosum,
T.
hasselquistiae,
T.
elegans;
12-18
in
T.
oficinale,
A.
cordata
and
A.
trachycarpa.
Filiform bracts are found in
T.
pustulosum,
T.
hasselquistiae
(Fig.
6)
and
T.
elegans
(Fig.
6);
filiform-linear in
A.
trachycarpa,
A.
cordata
(Fig.
6)
and
364
DAWUD AL-EISAWI AND
S.
L.
JURY
T.
brachyreenium
BA
T.
officinale
I
cm
H
tE
%
tBbC
T.
hirtocerpum
Figure
3.
Leaf silhouettes
of
four
species of
Tordylium;
basal leaves labelled
A
and sequentially
lettered
up
the plant. The extremes
of
variation in
T.
ojicinale
are shown.
T.
oficinale
(Fig.
7);
linear
in
T.
aegyptiacum,
T.
maximum,
T.
lanalum
(Fig.
6)
and
T.
macropetalum,
T.
maximum
and
S.
carmeli
(Fig.
9);
linear-lanceolate in
T.
apulum
(Fig.
6)
and
T.
pestalotzae
(Fig.
7);
lanceolate in
T.
persicum;
T.
cappadocicum,
and
T.
bruchytuenium
(Fig.
8);
and oblanceolate-spathulate in
T.
syriacum
(Fig.
6).
REVISION
OF
THE
GENUS
TORDYLUM
365
1.
1
A.
trachycarpa
I
I
A.
cordafa
L
I
crn
H
A.
tracbycorpo
T:
pustulosum
Figure
4.
Leaf silhouettes
of
Ainsworthia cordata, A. trachycarpa
and
Tordylium
pustulosum;
basal leaves
labelled A and sequentially lettered up the plant.
366
DAWUD AL-EISAWI
IF
f
AND
S.
L.
JURY
K
T.
macropetolum
Mandenovia
komarovii
S.
carmeli
T.
maximum
Figure
5.
Leaf silhouettes of
Tordylium macropetalum,
T.
maximum, Synclcosciadium carmeli
and
Mandenovia komarovii;
basal leaves labelled A and sequentially lettered up the plant.
The bracts are pubescent in the filiform type, setose in the filiform-linear type,
ciliate in the lanceolate type, but strigose in
T.
maximum,
T.
macropetalum
and
S.
carmeli.
Bracteoles
The bracteoles are similar
to
the bracts, but usually unequal and five
to
eight
in number.
dpr
ma
I
dv
cdv
A
st
ma
cv
car
B
F
YE
A.
cordata
C
T.
etegans
P
I
T.
aegyptiacum
I T.
lanatum
I@
ii
B
@
A
T.
syriacum
I
C
I@
B
T.
pusiutosurn
C
I@
B
7:
hasselquisfiae
1
C
T.
apulum
Figure
6.
A.
The dorsal face of a mericarp (dorsally compressed) showing margin, stylopodium,
vittae and primary ribs. B. The commissural face of a mericarp (dorsally compressed) showing
margin and vittae;
C.
Bracts; F. Sernisphaerical mericarps. Abbreviations:
st
=
stylopodium,
dpr
=
dorsal primary rib, ma
=
margin, Idv
=
lateral dorsal vitta, cdv
=
central dorsal vitta,
cv
=
commissural vitta and car
=
carpophore rib. The differences between smooth margin (e.g.
Ainsworthia
trachycarpa),
corrugated margin (e.g.
Tordyfium
pustulosum)
and
rugose
margin (e.g.
7.
aegyptiacum)
are shown. The number, position and size of the vittae are also important
taxonomically.
368
DAWUD AL-EISAWI AND
S.
L.
JURY
C
E
T;
officinofe
#
T;
officinofe
A
YE
$D
0
A
T.
hirtocorpum
i
C
I
C
T:
oegoeum
F
D
A
T.
pestalozzae
Figure
7.
A. The dorsal face of the mericarp showing the corrugated margin, stylopodium, primary
ribs and two central vittae which are clearly visible in
Tor4lium
oficinale
and
7'.
hirtocarpum
but
reduced to two spots near the stylopodium in
7'.
aegaeum
and
7'.
pestalozzae.
The lateral dorsal vittae
are invisible.
8.
The cornrnissural face showing margin, stylopodium, carpophore rib and the two
divergent vittae.
C.
The bracts showing linear-filiform type in
7'.
oficicinale
and
7'.
aegaeum
and the
lanceolate type in
7'.
hirtocarpum
and
7'.
pestalouae.
D. Umbels at anthesis showing length of rays
which
are
equal in
7'.
hirtocarpurn
and
1.
pestalouac
and not equal in
'I,
oficinale
and
7.
aegaeum.
E.
Umbels in fruit showing
rays.
Sepals
The sepals are absent in most of the species, but in
T.
aegyfitiacum
and
T.
syriacum
they are present, although minute, and in
T.
maximum,
T.
macropetalum
and
S.
carmeli
well-developed and strigose.
Petals
The petals are obovate, 2-lobed, white, yellow or pink; the white colour at
anthesis often appears yellow in dried material.
REVISION
OF
THE GENUS
TORDTLUM
369
T.
brachyteeniurn
T.
ebracteaturn
u
T.
C8ppadocicU171
Figure
8.
A.
The dorsal face of the mericarp showing smooth margin in
Tordylium brachytaenium,
corrugated in
7.
sp.
Dudley D.35477,
T.
persicum
and
T.
cappadocicum,
stylopodium, primary ribs and
the vittae: two central vittae in
T.
persicum
and
T.
cappadocicum;
vittae absent in
1.
sp.
Dudley
0.35477
and absent or two broken central vittae in
T.
bractytacnium.
The lateral dorsal vittae are
invisible in all the mericarps of this group.
B.
The dorsal face of the mericarp showing margin,
stylopodium, carpophore ribs and vittae: two parallel vittae in
7.
persicum
and
T.
cappodocicum,
two
divergent in
T.
brachytaenium
and two incomplete in
1.
sp.
Dudley 0.35477.
C.
Bracts showing the
lanceolate type in all species except
T.
sp.
Dudley 0.35477
in which the bracts are absent.
T.
maximum
S.
carmelt
T.
moctvpetalum
M.
kom arovii
Figure
9.
A.
The dorsal face of the mericarp showing smooth margin in
Tordylium maximum,
Synelcosciadum carmeli
and
Mandenova komarovii,
corrugated margin in
7.
macropetalum,
stylopodium,
primary ribs and the
4
dorsal vittae.
B.
The commissural face of the mericarp showing margin,
carpophore ribs and the two parallel vittae.
C.
Bracts showing the linear, strigose type, and equal
rays without bracts in
Mandenouia komarovii.
370
DAWUD AL-EISAWI AND
S.
L.
JURY
The outer flowers in all species have 2-radiate, equally or unequally 2-lobed
petals;
T.
apulum
is an exception, the outer flowers having only one radiate,
equally 2-lobed petal.
The size of the petals varies from 2 to
10
mm. The smallest size (1.5-2.5 mm)
is found in
T.
hirtocarpum
and the largest (7-10 mm) in
7:
oficinale
and
T.
macropetalum.
The petals are glabrous, only rarely hairy at the base, except for
T.
maximum,
T.
macropetalum
and
S.
carmeli,
where all the adaxial surface is covered with
strigose hairs.
Style and stylopodium
The style in
all
species is 0.5 mm long and glabrous, except in
T.
maximum,
T.
macropetalum
and
S.
carmeli
where it is 1-2 mm long and strigose. The
stylopodium is conical-depressed, rarely with an undulate margin and glabrous,
except in the same three species where it is conical and strigose.
Umbels
The diameter of the umbels
at
athesis varies according
to
the number and size
of the rays and whether the rays are equal or not. In general, the diameter
ranges from 20-200mm, with two
to
40
rays 0-60mm long. For details see
Table
1.
TABLE
1.
Some morphological details
of
the
umbels
Species
Diameter
of
umbel
at Length Rays
anthesis Number
of
rays equal
(
+
)
(mm)
of
rays
(mm)
or not
(-)
Ainsworthia cordata
A. trachycarpa
Tordylium aegaeum
7.
aegyptiacum
T.
apulum
7.
brachytaenium
1.
cappadocicum
7.
elegans
T.
hasselquistiae
T.
hirtocarpum
1.
lanatum
1.
macropetalum
T.
maximum
T.
oficinale
7.
persicum
1.
pestalozcae
1.
pustulosum
1.
yiacum
7.
sp.
Dudley 0354
77
Synelcosciadium carmeli
Mandenovia komarovii
30-50
40-
I00
20-30
40-
100
30-70
30-40
30-70
30-60
40-80
10-20
40-70
30-40
30-50
30-50
20-50
20-30
30-50
30-60
10-20
40-
100
4-60
12-25
15-30
5-12
6-16
3-8
10-16
10-25
10-16
8-20
2-4
2-8
6-15
6-15
10-25
5-20
5-12
10-15
5-10
2-3
3-7
3-8
10-30
20-60
3-18
10-70
30-50
5-20
10-50
15-50
5-40
5-10
15-40
5-30
10-40
3-30
5-40
2-15
8-30
5-40
0-80
0-80
30-40
REVISION
OF
THE GENUS
TORDYLUM
37
1
Fruits‘
(Figs 6-9)
The fruits may be dimorphic or uniform. In species with dimorphic fruits the
mericarps are either orbicular and dorsally compressed or hemispherical. The
dorsally compressed mericarps are 6-10 mm in diameter, with thick, smooth to
rugose margins in
T.
aegyptiacum
and
T.
lanatum
(Fig. 6), and 5-8 mm in
diameter, with smooth, thin margins in
A.
cordata
and
7.
elegans
(Fig. 6). The
hemispherical mericarps are
4-5
mm in diameter in
T.
aegyptiacum
and
T.
lanatum
(Fig.
6),
and 2-3 mm in diameter in
A.
cordata
and
T.
elegans
(Fig. 6).
In species with uniform fruits the mericarps are 3-12 cm long, orbicular to
sub-orbicular or ovate
to
broadly elliptical, with thick smooth to corrugated
margins.
Tordylium syriacum
(Fig. 6) has mericarps 8-12 mm long, orbicular with
thick, rugose margins.
Tordylium pustulosum,
T.
brachytaenium
and
A.
trachycarpa
(Fig. 6) have mericarps 4-6mm long, orbicular to sub-orbicular with thick,
smooth margins.
Tordylium maximum
and
S.
carmeli
(Fig. 9) have mericarps
6-8 mm long, sub-orbicular to broadly elliptical again with smooth margins.
Tordylium ojicinale,
T.
hirtocarpum
and
T.
pestalozzae
have mericarps 3.0-5.5 mm
long, ovate, with thick, corrugated margins.
Tordylium macropetalum
(Fig.
7)
and
T.
hasselquistiae
(Fig. 6) have mericarps 6-8 mm long, broadly elliptical with
thick, corrugated margins. Positions of vittae, vascular bundles, etc. are seen in
Figure 10 showing mericarp anatomy.
Vittae
(Figs 6-9)
The number of vittae, their shape and position in the mericarp are important
taxonomic characters in the Apiaceae. In this group
of
genera the number
of
vittae has been used to split the genus
Tordylium
and to recognize subgenera and
sections. Hoffmann (1806) segregated
T.
apulum
as a monotypic genus,
Condylocarpus
on the basis
of
it having ten to 12 vittae on the dorsal face and ten
on the commissural; De Candolle (1832), Boissier (1872) and Drude (1896)
divided the genus on the basis of the number
of
vittae, but the position and
shape of these vittae have not been used by previous taxonomists, although we
have found these valuable characters.
SCANNING ELECTRON MICROSCOPY
OF
MERICARPS
Heywood
&
Dakshini (1971) demonstrated the importance of the detailed
study of the mericarp to understand better and discriminate between genera
of
the Apiaceae tribe Caucalideae. Their study was based on the structure
of
the primary and vallecular ridges and the hairs or spines which are found on
them.
Similarly, in this group, the mericarps provide valuable taxonomic
characters, and mericarps of 20 species were examined by scanning electron
microscopy. Dried mericarps were mounted on copper stubs by double-sided
‘Sellotape’ and coated with a thin layer of gold in a vacuum coating unit with a
revolving stage. The samples were then examined in a JSM-2 or JSM-35
372
DAWUD AL-EISAWI AND
S.
L.
JURY
rvb
cv
A
B
C
Figure
10.
Median transverse section in mericarps
of
Tordylium
showing the anatomical structure.
A.
Tordylium hitrocarpum.
B.
7.
maximum.
C.
T.
hasselquistiae.
Key to abbreviations: w
=
wing,
cvb
=
commissural vascular bundle,
h
=
hair, Idv
=
lateral dorsal vitta, Ivb
=
lateral vascular
bundle, end
=
endosperm, emb
=
embryo, dvb
=
dorsal vascular bundle, cdv
=
central dorsal
vitta, me
=
mesocarp, en
=
endocarp, wn
=
wing neck, cv
=
commissural vitta, rvb
=
carpophore
vascular bundle.
scanning electron microscope in the Plant Science Laboratories, University of
Reading, at accelerating voltages of 15-25 kV.
This group
of
plants has dorsally compressed mericarps with thickened
margins and inconspicuous primary and vallecular ridges. The features showing
taxonomic variation are:
(1)
(1)
the mericarp margin;
(2)
the hair types on the
mericarp surface;
(3)
the style and stylopodium; and
(4)
the presence or absence
of calyx-teeth and their indumentum. These features are illustrated in Figs
11-46.
Mericarp margin
The margins of the mericarps vary from thin and smooth in
T,
elegans
and
A.
cordata,
to thick and smooth in
T.
pustulosum,
7.
brachytaenium,
A.
trachycarpa
REVISION
OF
THE
GENUS
TORDrLUM
Figures
11-16.
Fig.
11.
Tordylium
apufum:
mericarp showing surface with vesicular hairs and
corrugated margins. Scale bar
=
2.2
mm. Fig.
12.
Tordylium
elegans:
mericarp showing surface with
papillate hairs and smooth surface; scale bar
=
2.2
mm. Fig.
13.
Ainsworthia
cordala:
mericarp
showing surface with sparse vesicular hairs and smooth margin; scale bar
=
1.5
mm. Fig.
14.
Ainsworthia trachycarpa:
mericarp showing large vesicular hairs and smooth margin; scale
bar
=
0.7
mm. Fig.
15.
Tordyium
grincum:
mericarp showing surface with small, large vesicular and
linear hairs and a rugose margin; scale bar
=
I,
1
mm. Fig.
16
Tordjdium hasselquistiae:
mericarp
showing surface with vesicular hairs and corrugate margins; scale bar
=
1.1
mm.
373
374
DAWUD AL-EISAWI AND
S.
L.
JURY
Figures
17-22,
Fig.
17.
Tordylium
sp.
Dudley
0.35477
mericarp showing surface with large and small
vesicular hairs and corrugated margin; scale bar
1.0
mm.
Fig.
18.
Tordyliurn
pestalouae:
mericarp
with large and small vesicular hairs and corrugated margin; scale bar
1.1
mm. Fig.
19.
Tordylium
aegaeum:
mericarp showing surface with small vesicular, long linear hairs and corrugated margin;
scale bar
=
1.2
rnm.
Fig.
20
Tordyliurn
o@cinale:
mericarp showing surface with small and mostly
long vesicular hairs and corrugated margin; scale bar
=
1.1
mm.
Fig.
21.
Tordylium
hirlocarpum:
mericarp surface with small vesicular and long linear hairs and corrugated margin; scale
bar
=
1.1
mm.
Fig.
22
Tordylium
pushlosum:
mericarp with
small
and large vesicular hairs and
smooth margin; scale bar
=
1.0
mm.
Figures
23-28.
Fig.
23.
lordylium
macropetalum:
mericarp with strigose surface and thick corrugated
margin; scale bar
=
I
.7
mm.
Fig.
24.
Tordylium macropetalum
with hairy style, stylopodium and calyx
teeth; scale bar
=
0.6
mm.
Fig.
25.
Tordylium maximum:
mericarp with strigose surface and smooth
margin; scale bar
=
1.7
mm.
Fig.
26.
lordylium maximum:
hairy style, stylopodium and calyx teeth;
scale bar
=
0.3
mm.
Fig.
27.
Synelcosciadzum
curmeli:
mericarp with strigose surface and smooth
margin; scale bar
=
I
.7
mm.
Fig.
28.
Synelcosczudzum
curmeli:
hairy style, stylopodium and calyx
teeth; scale bar
=
0.5
mm.
376
DAWUD AL-EISAWI AND
S.
L. JURY
Figures
29-33.
Fig.
29.
Tordylium elegans:
smooth margin of mericarp with papillate hairs; scale
bar
=
220
pm.
Fig.
30.
Tordylium
aegaeum:
corrugated margin
of
mericarp with reticulate surface
covered by reduced vesicular hairs; scale bar
=
25
pm. Fig.
31
Tordylium syrzacum:
rugose margins
of
mericarp with strigose and papillate hairs; scale bar
=
315
pm.
Fig.
32.
Tordylium
sp.
Dudley
D.35477
corrugated margin of mericarp with reticulate striate surface and reduced hairs; scale
bar
=
36
pm.
Fig.
33.
Tordylium acgyptiacum:
rugose margin
of
mericarp with papillate hairs; scale
bar
=
115
pm.
REVISION
OF
THE
GENUS
TORDYLUM
Figures 34-40. Fig. 34.
Ainsworthia trachycarpa:
large vesicular hairs
of
rnericarp; scale bar
=
35
prn.
Figs 35
&
36.
Tordylium
pustulosum:
small and large stalked hairs; scale bar
=
45 pm. Fig. 37.
Tordylium
sp.
Dudley
0.35477:
vesicular hairs
of
rnericarp; scale bar
=
30
krn.
Fig.
38.
Ainsworthia
cordatu:
peltate stalked hairs; scale bar
=
25
pm.
Fig. 39.
Tordylium uegaeum:
strap-like linear hair
with thickened margin; scale bar
=
70
km.
Fig.
40.
Tordyliiurn
elegans:
papillate and linear hairs;
scale bar
50
prn.
377
378
DAWUD AL-EISAWI AND
S.
L. JURY
Figures
41-46.
Tordyliurn
cappadocicum.
Fig.
41.
Mericarp showing the surface and the thick
corrugated margin; scale bar
=
I.
1
mm. Fig.
42.
Glabrous style and stylopodium; scale
bar
=
220
pm.
Fig.
43.
Mericarp surface with small vesicular
and
long tubular hairs; scale
bar
=
220
pm. Fig.
44.
Long hairs; scale bar
=
440
pm.
Fig.
45.
Striate mericarp surface showing
small vesicular hairs; scale bar
=
200
pm. Fig.
46.
Long hair base; scale bar
=
22
pm.
REVISION
OF
THE
GENUS TORDTLUM
379
and
T.
maximum,
to rugose in
T.
syriacum
and
T.
aegyptiacum,
to thick and
corrugated in
7.
macropetalum,
T.
ofkinale,
T.
hirtocarpum,
T.
aegaeum,
T.
pestalotzae,
7.
sp.
(Dudley D.35477),
T.
persicum,
T.
cappadocicum
and
T.
apulum.
Although some species have thick, corrugated mericarp margins, it is not
necessary to assume that they are related. The detailed structure obtained by
the electron microscope has shown that
T.
macropetalum,
which has a thick,
corrugated margin with strigose hairs on the surface, is different from
T.
hasselquistiae
which has a thick, corrugated margin and only rarely covered
with vesicular hairs. Both
T.
hasselquistiae
and
7.
macropetalum
are different from
T.
oBcinale,
T.
hirtocarpum,
T.
pestalozzae
and
T.
cappadocicum,
which have thick,
corrugated margins and papillae.
Likewise
T.
brachytaenium
which has a smooth margin with reticulate surface
and reduced papillae, differs from
T.
maximum
which has a smooth margin with
strigose hairs. Both of these were different from
T.
lanatum,
T.
pustulosum,
T.
elegans,
A.
cordata
and
A.
trachycarpa
which have
a
smooth margin with
vesicular hairs.
Hairs
Mericarps with strigose indumentum.
These are only present in
T.
maximum,
7.
macropetalum,
and
S.
carmeli.
All the parts of the mericarps including the
styles, stylopodiums, calyx-teeth and the margins have one type
of
sharp,
pointed, tuberculate, strigose hair.
Mericarps with vesicular hairs.
These are found in all species except the above.
These vesicular hairs can be: (a) small, which are found in all species; (b) large,
found in
A.
hachycarpa,
T.
apulum,
7.
pustulosum,
7.
hasselquistiae,
7.
lanahm,
T.
aegvptiacum;
(c) long, found in
T.
oJicinale,
T.
aegaeum,
T.
pestalozzae,
T.
ebracteatum
and
T.
brachytaenium;
(d)
long tubular, found in
T.
persicum,
T.
cappadocicum,
7.
hirtocarpum,
T.
syriacum,
T.
lanatum,
T.
aegaeum
and
T.
elegans,
or
(e)
peltate, found in
A.
cordata,
T.
syriacum
and
T.
hasselquistiae.
Usually more than
one
type of hair is found in one species.
We
believe that all
these five types mentioned, have a single origin because they have the same
granular surface and the same basal structure.
It
is possible that all these hair
types are derived from the small, vesicular hairs which are found in all
species.
Sole and stylopodium
The style and stylopodium are glabrous in all species except
T.
maximum,
1.
macropetalum
and
S.
carmeli
where they have strigose hairs similar to those
found on the mericarps.
Calyx-teeth
The calyx-teeth are absent, or minute, and glabrous in all species except
T.
maximum,
T.
macropelalum
and
S.
carmeli
which have well-developed, strigose
calyx-teeth.
380
DAWUD AL-EISAWI AND
S.
L.
JURY
Results
From this evidence it can be seen that:
(1)
Synelcoscadium carmeli
must be grouped with
T.
maximum
and
T.
macropetalum.
(2)
It
is very difficult to separate
Ainsworthia
from
Tordylium
on the basis of the
detailed structure of the mericarps.
(3)
All
Tordylium
species which have the four dorsal vittae visible, have
mericarp margins with vesicular hairs; and the species which have invisible
lateral dorsal vittae have papillate mericarp margins.
(4)
It
is
difficult to distinguish
T.
aegaeum
from
7.
pestalozzae
and
T.
persicum
from
T.
cappadocicum
on the basis
of
the detailed structure
of
the mericarp
surface. This supports the other evidence obtained from the morphology,
anatomy and the palynology.
COMPARATIVE FRUIT ANATOMY
The internal anatomy of the mericarp provides a number of extra valuable
taxonomic characters. Thellung
(
1926) illustrated the mericarp anatomy for
three species of
Tordylium-T. apulum,
T.
maximum,
and
T.
ojicinale-and
showed the number and position of vittae in the dorsal and commisural faces
and the hairs
on
the mericarp surfaces, as well as the general appearance
of
the
fruits. Theobald (1971) studied the comparative anatomy and development of a
few genera of the Umbelliferae including the genus
Tordylium.
He noted that
although the genus has often been subdivided into several genera, with
Ainsworthia
and
Hasselquistia
as possible segregates, in either circumscription the
taxa which make up the group have several distinctive anatomical characters in
common.
Although the species examined do have many anatomical characters in
common, they have other characters which can be used to differentiate between
them.
Wings
Some species have very long wings differentiated into a head and neck, for
example
1.
syriacum,
T.
aegyptiacum,
7.
apulum,
T.
hasselquistiae,
T.
elegans,
A.
cordata;
others have
a
short neck, for example
T.
pustulosum, A. trachycarpa,
T.
macropetalum,
T.
maximum
and
S.
carmeli;
a third group is without a neck, just
having a thickened margin, for example
T.
persicum,
T.
cappadocicum,
T.
pestalozzae,
T.
ojicinale,
T.
hirtocarpum,
T.
brachytaenium
and
T.
sp.
(Dudley
0.35477).
Although some species have mericarps with smooth margins,
it
is not
necessary to assume that they are related to each other, or that they differ from
those with corrugated margins, because in
T.
hasselquistiae,
T.
apulum,
T.
aegyptiacum
and
T.
macropetalum
with thick, rugose or corrugated margins, the
position
of
the lateral vittae and the wing differs from
T.
ojicinale,
T.
hirtocarpum,
T.
pestalozzae,
T.
persicum
and
T.
cappadocicum,
even though all of the latter have
mericarps with thick, corrugated margins. The same applies to
A.
trachycarpa
and
T.
pustulosum
with smooth margins which differ from
T.
brachytaenium
(also
REVISION
OF
THE GENUS
'TORDTLUM
38
1
with a smooth margin)
by
the position of the lateral vittae and the neck of the
wing.
Vascular bundles
The vascular bundles are rarely recognized because they are embedded in the
lignified tissue of the endocarp.
Number and position
of
the vittae
All species of
Tordylium, Ainsworthia
and
Synelcosciadium
have four dorsal and
two commissural vittae, except
T.
apulum
which has ten to 12 dorsal and
10
commissural;
T.
oJicinale,
T.
hirtocarpum,
T.
pestaloxae,
7.
aegaeum,
T.
brachytaenium,
T.
sp.
(Dudley
D.35477),
T.
persicum
and
T.
cappadocicum
have
four dorsal vittae but only the central ones are visible externally. This
is
because
the lateral dorsal vittae are situated under the thickened margin of the
mericarp.
Shape and size
of
the vittae
The shape of the vittae is elliptical
to
oblong. The size varies from 60-240 pm.
According to the size, the species can be grouped into four categories as follows:
(1)
60-80 pm in
T.
aegyptiacum
and
T.
lanatum.
(2) 100- 140
pm
in
T.
ojicinale,
T.
hirtocarpum,
T.
pestalozzae,
T.
cappadocicum,
(3)
160-200
pm
in
T.
pustulosum,
T.
hasselguistia,
A.
trachyarpa,
A.
cordata,
(4) 200-240 pm in
T.
apulum.
T.
persicum,
T.
sp.
(Dudley 0.35477),
T.
aegaeum
and
T.
syriacum.
T.
elegans,
T.
maximum,
T.
macropetalum
and
S.
carmeli.
Endocarp
The endocarp in all the species examined has one to six lignified layers which
surround the endosperm, and extend through the wings connecting with the
commissural primary vascular bundles. The species can be divided, according to
the endocarp layers, into three groups:
(1)
one or two layers in
T.
pustulosum,
T.
hasselquistiae,
1.
elegans,
A.
trachycarpa
and
A.
cordata;
(2) four to six layers in
T.
aegyptiacum,
1.
lanatum
and
T.
syriacum,
and (3) two to four layers all other
species.
Mesocarp
All the species have one or two layers of parenchyma except
T.
maximum
and
S.
carmeli
which have four to six layers of parenchyma.
As seen from the previous discussion, it is very difficult to separate
Ainsworthia
from
Tordylium
on the basis of the fruit anatomy, as Theobald
(197
1)
noted.
Likewise it is difficult to discriminate between
Synelcosciadium
and
Tordylium.
POLLEN
MORPHOLOGY
Of special relevance to this study
of
the genera
Ainsworthia, Tordylium,
Synelcosciadium
and
Mandenovia
are the works of Cerceau-Larrival
(
1962, 1963).
382
DAWUD AL-EISAWI AND
S.
L.
JURY
In
1962
she examined the pollen of
Tordylium
persicum,
T.
syriacurn,
T.
maximum
and
Synelcosciadium carmeli,
and described them all as belonging to the group
where the pollen is equatorilly constricted. In
2.
maximum
and
S.
carmeli
there is
a second order symmetry although some species (not specified) of
Tordylium
have
third order symmetry.
In
1963
she described
a
further species,
Tordylium apulum,
which has third order symmetry.
From this one can deduce that pollen-grains with second order symmetry are
in fact bi-aperturate whereas third order symmetry implies tri-aperturate pollen.
These early papers were based on light microscope studies. Her later work
involved the use of the scanning electron microscope, but does not figure any
further species of this small section of the family. She classified the pollen-grains
of the sub-tribe Tordylinae under the fifth type which she calls ‘The
equatorially-constricted
type’. She distinguished this type by having a size range
from
50-70
pm.; the exine assumes a proportion such that in the equatorial
zone, the pollen-grain becomes ‘winged’ on account of the carina
of
the exine,
the ectoapertures get smaller and are often scarcely visible.
All the work was carried out using the scanning electron microscope. This
only shows the surface topography of the pollen and reveals its symmetry in
strict botanical (not crystolographic) sense. The ectoapertures are revealed. This
method does not reveal the outline of the endexine,
so
important in Cerceau-
Larrival’s studies, nor does it show the form of the endoaperture.
Description
In this work, strict botanical terminology is used based on Cerceau-Larrival
&
Roland-Heydacker
(1976),
especially for the pollen tectal surface. The pollen
of
Ainsworthia, Tordylium, Synelcosciadium
and
Mandenovia
have been examined and
can be divided into the following groups:
I.
Tricolporate with radial symmetry. The outline may be elliptical and the
carinae continuous. In this case, the side of the ellipse can be either flat or
convex.
Or
the outline can be dumb-bell shaped due to a constriction in
the equatorial region caused
by
a
discontinuity or constriction of the
carinae.
A.
Outline elliptical, carinae continuous.
(i)
Sides of ellipse flattened.
1.
The pollen surface sculpturing is secondary-perforate, in
T.
pestalozzae
(Fig.
47)
and
T.
aegaeum
(Fig.
49).
These two
species have identical shape and sculpturing.
T.
oJicinale
(Fig.
51)
has similar shape and sculpturing to the previous
two
species,
but the sculpturing becomes more pronounced around the apertures.
T.
branchytaenium
(Fig.
53)
has similar pollen sculpturing but
the grains are less prolate.
2.
The pollen surface sculpturing
is
striate, rugulose on the
carinae becomes more pronounced around the apertures,
T.
apulum
(Fig.
48).
(ii) Sides of ellipse convex. The pollen surface sculpturing is cerebroid
to rugulose, e.g.
Mandenovia komarovii
(Fig.
59).
REVISION
OF
THE
GENUS
TORDYLUM
Figures 47-54.
Pollen
of
Tordylium
species.
Fig. 47.
T.
pestalorzae:
scale bar
=
14
pm.
Fig. 48
T.
apulum:
scale bar
=
I1
pm.
Fig. 49.
7.
aegaeum:
scale bar
=
14
prn.
Fig. 50.
1.
lanatum:
scale
bar
=
8.5
pm.
Fig. 51.
T.
ojicinule:
scale bar
=
14
pm.
Fig. 52.
T.
hasselquistiu:
scale bar
=
8.5
pm.
Fig. 53.
T.
brachytaenium:
scale bar
=
8.5
pm.
Fig. 54.
T.
pustulosum:
scale bar
=
8.5
prn.
383
384
DAWUD AL-EISAWI AND
S.
L.
JURY
Figures 55-60. Fig. 55.
Tordylium
cappadocicum:
scale bar
=
9.2
pm.
Fig. 56.
7.
maximum:
scale
bar
=
I1
pm.
Fig. 57.
T.
persicurn:
scale bar
=
7
pm.
Fig. 58.
7.
macropetalum:
scale bar
=
8.5
pm.
Fig.
59.
Mandanouia
komaravii:
scale bar
=
10
pm.
Fig.
60.
Synelcosciadium
carmeli:
scale bar
=
11
prn.
B.
Outline dumb-bell shaped, equtorially constricted, carinae
discontinuous. The pollen surface sculpturing is perforate, rugulate to
rugulose striate.
T,
hasselquistia
(Fig.
52),
T.
pustulosum
(Fig.
54),
1.
lanatum
(Fig.
50),
1.
syriacum
(Fig.
65),
T.
aegyptiacum
(Fig. 61),
A.
cordata
(Fig.
62),
A.
trachycarpa
(Fig.
63).
Owing to their close
similarity, it has not proved possible to separate these species further on
the basis of their pollen morphology.
11.
Dicolporate with second-order symmetry. This group has pollen grains
with two flattened sides. They are oblong to broadly elliptical and may be
subdivided on the basis of their outline into an elliptical group and a
dumb-bell shaped group.
REVISION
OF
THE GENUS
TORDYLUM
385
Figures
61-65. Fig. 61.
Tordylium
ocgvptiacum:
scale bar
=
8.5
pm.
Fig. 62.
Ainsworthiu
cordah:
scale
bar
=
8.5
pm.
Fig. 63.
Aimworthiu
trachycurpu:
scale bar
=
18.4
pm.
Fig. 64.
Tordyliurn
elguns:
scale
bar
=
7.5
pm.
Fig. 65
Tordyllium
syriucu:
scale bar
=
12.2
pm.
A.
Outline dumb-bell shaped, equatorially constricted. The pollen
surface sculpturing is secondary-perforate, and becomes more
pronounced around the colpi and
in
the equatorial regions.
T.
cappadocicum
(Fig.
55),
T.
persicum
(Fig.
57),
and
T.
macropetalum
(Fig.
58).
B.
Outline elliptical, pollen not equatorially constricted. The pollen
surface sculpturing is perforate-rugulate.
S.
camzeli
(Fig.
60)
and
T.
maximum
(Fig.
56)
386
DAWUD AL-EISAWI AND
S.
L.
JURY
Discussion and conclusions
From the results obtained from this study, the following points have been
noticed:
(1)
It is very difficult to separate
Ainsworthia
from
Tordylium
on the basis
of
the
pollen morphology because at least five other species have identical pollen
grains (group I(ii)) which cannot be separated from each other.
(2)
All species with the four dorsal vittae visible have pollen grains with a
tricolporate dumb-bell shaped and discontinuous carinae (group
I
(ii)
)
.
(3)
All species with the lateral dorsal vittae invisible (hidden under the wings)
have pollen grains which are tricolporate, elliptical in outline and continuous
carinae (group I(i)), except
T.
persicum
and
T.
cappadocicum
which have
dicolporate pollen grains.
(4)
T.
apulum
with 12 dorsal and ten commissural vittae, has a combination of
characters of shape and sculpturing of the pollen groups I(i) and I(ii).
(5)
Tordylium maximum,
T.
macropetalum
and
S.
camzeli
have dicolporate pollen
grains which correlates with other evidence from morphology and the detailed
structure of the mericarp surfaces.
(6)
Mandenovia komarovii
pollen grains have
a
shape and sculpturing which are
different from those of
Tordylium
and
Synelcosciadium
pollen, which supports
keeping this group separate.
CYTOLOGY
Chromosome information for the
Tordylium
and
Ainsworthia
group has been
summarized by Moore (1971). The most important previous work was that of
Runemark (1968), which dealt with six species occurring in The Aegean region.
In view of the much wider distribution of the group, and the lack of
chromosomal data from other regions, it is perhaps still premature to attempt a
final analysis of the cytological relationships within the group. However, further
cytological data
is
unlikely to be gathered easily because most of the species are
very difficult to germinate, some of them are endemic to very small regions from
which it is difficult to collect suitable material. Indeed, some of the species are
only known from the type specimens (for example
T.
brachytaeniurn).
All the
cytological records known are presented in Table
2.
As shown in Table
2,
the number
2n
=
8
in
T.
hirtocarpum
is equal to the
lowest number ever recorded in the family. The occurrence of
2n
=
20
in most of
the species made Runemark suggest that the basic number in this group is
x
=
10, but on this assumption it was difficult to explain the
2n
=
8
in
T.
hirtocarpum,
so
he said that this could be explained
by
having a species with a
chromosome number
2n
=
10
(based on
x
=
5)
then all the species would be
treated as tetraploid, except
T.
hirtocarpum,
with
2n
=
8, which may have been
reduced by reciprocal translocation.
If
one accepts Runemark’s assumption that the basic number in this group is
x
=
10, then
2n
=
10
and
2n
=
8
must be derived from the haploid number by
losing two chromosomes, and this usually will be sterile, but since the plant with
2n
=
8
is fertile and there is pairing between the chromosomes, as Runemark
illustrated, there is no reason to assume that the species with
2n
=
8
is derived
from
2n
=
20.
REVISION
OF
THE GENUS
TORDTLUM
387
TABLE
2.
Chromosome counts recorded
for
Toordylium, Ainsworthia, Synelcosciadum
and
Mandenovia
Species Chromosome
No.
Reference
____
~~ ~~
Tordylium maximum 2n
=
22
Tamaschjan
(1933)
2n
=
20
Runemark
(1968)
T.
apulurn 2n
=
20
Runemark
(1968)
7.
pestalozzae
(as
7.
aegaeurn) 2n
=
20
Runemark
(1968)
1.
oJ'icinale 2n
=
18
Runemark
(1968)
7.
hirtOCaTpUrn 2n
=
8
Runemark
(1968)
7.
yiacum 2n
=
20
Gardi:
&
Gardt
(1954)
T.
cordaturn
(as
A. cordata)
Gardt
&
Gardi:
(1954)
Constance
el
a!. (1976)
T.
aegyptiacurn
n=
10
Present paper
(Fig.
66A)
Ainsworthia trachycarpa
n=8
Present paper (Fig.
6B).
2n
=
20
n=
10
n=8
n=9
n
=
10
Constance
et al.
(1
97
1
)
Constance
et al.
(
1976)
Constance
et al.
(1
976)
Synelcosciadium carrneli
n=
10
Present paper (Fig.
66C)
Mandenovia komarovii 2n
=
22
Alava
(1973)
Since the number
2n
=
16 has been recorded, then it is a more likely
assumption that the basic number is
x
=
4
and that
2n
=
16
is
thus tetraploid
and
2n
=
20
is either pentaploid derived from crossing between tetraploid with
2n
=
16 and haxaploid
2n
=
24,
or
it
is derived from one or the other by either
losing or gaining four chromosomes.
However,
T.
uegyptiucum
(Fig.
66A)
with
2n
=
20
shows irregular meiosis;
some of the cells had one quadrivalent and eight bivalents or two quadrivalents
and six bivalents. This might suggest that the
2n
=
20
in
this species must be of
Figure
66.
A. Meiotic configurations in
pollen
mother
cells
at first metaphase.
Tordyliurn aegyptiacurn,
N
=
10
(Jordan Irbid,
1975 Al-Eisawi s.n.).
B.
Ainsworthia trachycarpa, n
=
8
(Jordan,
8
km
W
of
Karak, along the road
to
Chor,
1975, Al-Eisawi s.n.). C. Synelcosciadiurn carrneli, n
=
10
(Jordan,
Irbid, Al-Hemma,
1975, Al-Eisawi s.n.).
388
DAWUD AL-EISAWI AND
S.
L.
JURY
a hybrid origin, and this might support the second idea of
x
=
4, since the basic
numbers in the family range from
x
=
4 to
x
=
11.
Therefore, on the basis
of
the cytological information available, it is difficult
to separate the genus
Ainsworthia
with
A.
cordata,
2n
=
20
and
A.
trachycarpa
2n
=
16 (Fig. 66B) from the genus
Tordylium,
in which the number ranges from
2n
=
8 to
2n
=
20.
TAXONOMIC TREATMENT
Tor&ium
L.
Tordylium
L.
Genera Plantarum,
ed. 5:
1
1
1
(
1
754).
SYNONYMS:
Hasselquistia
L.,
Cent.
Pl.,
1:
9
(1755);
Amoen. Acad.,
4:
270 (1759).
Condylocarpus
Hoffm.,
Gen. Umb.,
ed. 2: 202 (1816).
Ainsworthia
Boiss.,
Ann. Sci.
Nut.,
ser.
3,
I:
343 (1844).
Synelcosciadium
Boiss.,
Ann.
Sci.
Nut.,
ser.
3,
1:
346 (1844).
Annual herbs, 150-800 mm tall, retrorsely setose, scabrous-pubescent to villous.
Stems erect, terete, striate to ridged, usually single. Basal leaves simple, or
1-
to
3-pinnate with ovate to cordate segments, with crenate to serrate margins;
upper leaves usually 3-lobed, rarely simple, with ovate-lanceolate segments.
Umbels
15-100
mm diameter. Rays (2-)5-20(-40),
0-80
mm, equal or
unequal. Bracts
0-
18,
filiform, linear, lanceolate, to
oblanceolate-spathulate,
deflexed, equal or unequal. Bracteoles 5-8, usually unequal. Flowers usually the
outer hermaphrodite, the central hermaphrodite and male. Sepals usually
absent,
or
present and well-developed. Petals white, pinkish or yellow, radiate,
the outer flowers have
1
or 2 large, equally or unequally 2-lobed) glabrous or
rarely strigose petals. Fruits uniform or dimorphic; semispherical mericarps;
dorsally compressed mericarps, orbicular to broadly elliptical or ovate; mericarp
margins smooth or corrugated, the mericarp surfaces covered with vesicular to
tubular or strigose hairs; primary ridges filiform; vittae 4 dorsal and
2
commissural, except in
T.
apulum,
which has 10-12 dorsal and 10 commissural
vi
t
tae.
TYPE:
T.
maximum
L.
Synopsis
of
Taxa
I.
Subgenus
Tordylium
A.
Section
Tordyliurn
1.
T.
maximum
L.
2.
T.
macropetalum
Boiss.
3.
T.
carmeli
(Labill.) Al-Eisawi &Jury
11.
Subgenus
Ainsworthia
Drude
4.
T.
apulum
L.
5.
T.
syriacum
L.
6.
T.
hasselquistiae
DC.
7.
T.
trachycarpum
(Boiss.) Al-Eisawi
&
Jury
8.
T.
pustulosum
Boiss.
B.
Section
Condylocarpus
(Hoffm.)
DC.
C. Section
Hasselquistia
(L.)
Boiss.
REVISION
OF
THE
GENUS
i0RDYLUM
9.
1.
lanatum
(Boiss.) Boiss.
10.
T.
aegptiacum
(L.)
Lam.
11.
T.
cordatum
(Jacq.) Poiret
12.
T.
elegans
(Boiss.
&
Bal.) Alava
&
Hub.-Mor.
13.
T.
cappadocicum
Boiss.
14.
T.
hitrocarpum
Cand.
15.
T.
oficinale
L.
16.
T.
bruchytanium
Boiss.
&
Heldr.
17.
T.
pestalozzae
Boiss.
18.
T.
ebracteatum
Al-Eisawi &Jury
389
Key
to the species
1
Plants retrorsely setose: sepals, petals, styles, stylopodium and
mericarps all covered with strigose hairs
Petals 8-12 mm, yellow; mericarps with corrugated margin
Petals
4-7
mm, usually white; mericarps with smooth margins
2
2
2.
macropetalum
3
3
Rays 10-16 not more than 30 mm long, not contracted in fruit;
Rays 3-7, up to
80
mm long, very much contracted in fruit; petals
1 Plants scabrous
to
villous; sepals, petals, styles and stylopodiums
glabrous, mericarps covered with vesicular hairs
Outer flowers with 1 large, equally 2-lobed petal; mericarps with
Outer flowers with 2 large, unequally to subequally 2-lobed petals;
petals 3-5 mm
.
.
Lmaximum
5-7mm
. .
3.
carmeli
4
4
10-12 dorsal and 10 commissural vittae
.
.
4.
apulum
mericarps with
4
dorsal and 2 commissural vittae
5 Mericarps with
4
dorsal vittae visible
6 Bracts and bracteoles oblanceolate-spathulate
.
.
5.
syriacum
6
Bracts and bracteoles filiform to narrow linear
7
Fruits uniform
8
8
Outer petals equally to subequally 2-lobed; mericarps 5-8 mm
Outer petals unequally 2-lobed; mericarps usually not more than
9
Leaf segments up to 30
x
30 mm or more, cordate; cauline
leaves acute; brackets
1-18
mm, filiform linear, mericarps
orbicular, cream.
.
7.
trachycarpum
9
Leaf segments not more than 15
x
15 mm, ovate; cauline leaves
cuspidate; bracts 6-
10
mm, filiform, mericarps orbicular to
broadly elliptical, reddish
. .
8.
pustulosum
diameter with thick corrugated margin
.
.
6.
hasselquistiae
5 mm diameter, with thick, smooth margin
7
Fruits dimorphic
10
Plants with dense, white, woolly hairs; leaves
2-
to 3-pinnate;
11 Entire plant with dense, white, woolly hairs; rays 2-8,
11 Plants with white, woolly hairs at the base, scabrous above;
bracts 0-5
subequal; mericarps blackish to greyish
.
.
9.
ianatum
rays 6-16, unequal; mericarps brownish
.
10.
aegyptiacum
390
DAWUD AL-EISAWI AND
S.
L.
JURY
10 Plants pubescent to scabrous; leaves 1-pinnate; bracts always
more than 5
Leaf segments up to 30x30 mm or more, cordate; outer
petals unequally 2-lobed; mericarps not more than 5 mm
diameter; central dorsal vittae continuous
Leaf segments not more than 15
x
15 mm, ovate; outer petals
subequally 2-lobed; mericarps up to
8
mm; diameter;
central dorsal vittae broken, not continuous
.
12.
elegans
12
.
11.
cordatum
12
5
Only the two central dorsal vittae visible
13 Outer petals subequally to equally 2-lobed; bracts broadly
13 Outer petals unequally 2-lobed; bracts narrowly lanceolate to
lanceolate; commissural vittae parallel
.
.
13.
cappadocicum
linear-filiform; commissural vittae divergent
14
Rays 2-4(-5), 5-12 mm long, equal
15 Bracts present
.
.
14.
hirtocarpum
15 Bracts absent
.
18.
ebracteatum
Bracts linear-filiform, more than 10; mericarps 2.5-3.5 mm
diameter
.
.
15.
oficinale
Bracts lanceolate-linear, less than 10; mericarps 4.5-5.5 mm
or more in diameter
1
7
Bracts lanceolate; mericarps orbicular to suborbicular with
17
Bracts linear-lanceolate; mericarps ovate, with thick,
14
Rays always more than 5, up to 30 mm long, unequal
16
16
thick, smooth margins
. .16.
brachytaenium
corrugated margins
. .
17.
pestaloxxae
I.
Subgenus
Tordylium
L.
Plants retrorsely setose; sepals well-developed, strigose; all the lower petals,
styles, stylopodia and mericarps covered with strigose hairs; pollen grains
dicolporate.
A.
Section
Tordylium
Petals radiate; vittae 6,
4
dorsal, 2 commissural, parallel.
1.
T. maximum,
L.,
Species
Plantarum:
240
(
1753).
TYPE:
Herb. Clifford,
90,
Tordylium
4 (BM, syntype).
Annual or biennial, 300-800 mm tall or more, retrorsely setose. Stems erect,
terete, ridged. Basal and cauline leaves 1-pinnate with 2-4 pairs
of
leaflets;
segments 10-18
x
20-35 mm, ovate, oblong, serrate, both sides covered with
setose hairs; upper leaves 3-lobed, with linear-lanceolate, coarsely serrate
terminal lobes. Umbels 30-50 mm in diameter. Bracts 3-8, 2-10
mm,
linear,
strigose. Rays 6-15, 10-40 mm, unequal. Bracteoles 5,
2-6 mm, unequal,
strigose. Sepals present, 0.2-1.2 mm, triangular, unequal, strigose. Petals white
to yellow, radiate, the outer 3-5 mm long, unequally 2-lobed, strigose at the
adaxial surface. Stamens 1.5 mm long. Styles
1
.O-1.3 mm; stylopodium conical,
both are strigose. Mericarps 6-8 mm in diameter, suborbicular to broadly
elliptical, with thick, smooth or rarely corrugate margin, and covered with
strigose hairs on the surface; primary ridges filiform; vittae
6,
4 dorsal and
2
REVISION
OF
THE
GENUS TORDTLUM
commissural, parallel. Flowering May-July.
2n
=
20,
22.
Most of Europe,
North Africa and the Middle East.
REPRESENTATIVE
SPECIMENS
EXAMINED.
ALBANIA:
Koci, 30 vi 1900,
Baldacci
22
(K);
Macedonia, near Karamudli, 26 vi 1917,
Turrill
442
(K); Gajtani,
18
vi
19 16,
Janchen s.n.
(WU)
.
ALGERIA:
terroius Pullives,
vi
19
10,
Faure s.n.
(E)
.
AUSTRIA:
Wein,
27
vi
1899,
Tgber
s.n.
(WU); Frankreich, Alpes Maritimes,
13
vi
1962,
Merxmuller
t3
Wiedmann
20251
(M); Grinzin near Vindobonam, vii
1881,
Halacy
2107
(M).
BRITISH
ISLES:
Essex,
S
of Essex, 30
vi
1942,
Lousley
840
(RNG);
S
of Essex,
2
ix 1944,
Lousley
s.n.
(RNG).
BULGARIA:
2
vi 1902,
Stribrny
s.n.
(E);
Bei Loutscha, 1895,
Hrumof
35
(WU).
CZECHOSLOVAKIA:
Moravia,
Mikulov, Klentnice,
3
viii 1947,
Frohlich
1349
(E,
K, M, WU).
FRANCE:
Fontan,
Les Buisarides et Moissons, 30 vi 1886,
Reuerchon s.n.
(E); Corailla, Vernet-Les-
Bains, 9 vi 1925,
Ellrnan
&'
Sandwith
142
(K);
Haute Garonne, Pyrenees,
Luchon,
8
vii 1975,
Souster
5
(RNG).
GERMANY:
Elbgehangeh,
0
viii 1902,
Hyne
s.n.
(M).
GREECE:
Thessalia, Kalanpak and Kaslreiki, 25 vi 1896,
Sintenis
996
(E);
Macedonia, Kyshakopega, 20 vi 1932,
Alston
&
Sandwith
420
(K).
HUNGARY:
vii 1877,
Tauscher s.n.
(E); Budapest, Harmashatarhegy, 14 viii 1944,
Papp 2620 (K); 18 vii 1917,
Xrzisch s.n.
(WU).
IRAN:
Azerbaijan, pass between
Astara and Ar'abil, 27 vii 1967,
Walton
69A
(E);
E of
Loweh,
Gorgan-Bijnwrd
road, 29
v
1959,
Lriiojenton
3975
(K).
ITALY:
Gallico, Hontmelas, 31 viii 1975,
Gandoger
&'
Lutson s.n.
(E); Eugane,
12
vii 1910,
Frirnmel s.n.
(WU).
ROMANIA:
dist. Adamclisi, Baneasa, 29 vii 1963,
Toma
s.n.
(M).
SPAIN:
Cerdange,
Angoustrine,
7
vii 1927,
Sennen
6084
(BM); Sierra de Segura,
8
vii 1850,
Bourgeau
679
(E,
K);
Prov. Jaen, Despenaperros,
28
v
1967,
Galiano et
al.
221
(RNG).
TURKEY:
Istanbul, near Biekilic,
11
vi 1968,
Buylop
13138
(E);
Prov. Bilecik,
Bilecik to Pazaryeri,
2
vii 1962,
Dauis
&
Coode 036456
(E); Suchak (Phyrgie),
25
vii 1857,
Balansa
1235
(K).
YUGOSLAVIA:
Macedonia, Tetovo, 470 m,
16 vii 1970,
Edrnondson
158
(BM, E, RNG); Sarjevosko, 25 vii 1920,
Mab
s.n.
(K);
Bosnieu, Tara-Gruppe,
1
km from Visegrad,
Thret s.n.
(M); Aydin,
Arslanli, E. of Naxilli, along the main highway, 2.5 km N of the highway on
Megan Cay river bank, 4 vi 1966,
Alma
&
Bocquet
5138
(E, W).
391
2.
T.
macropetalum
Boiss.,
Ann. Sci.
Nut.,
ser.
3,
1:
348 (1844).
TYPE:
Turkey, ad aggeres et sepes Cariae in valle Meandri prope Guzelhizar et
Nozlu, vi 1842,
Boissier s.n.
(BM, E, G,
K,
W).
Retrorsely setose, 300-700 mm tall. Stems erect, terete, striate to ridged. Basal
leaves oblong-ovate in outline, 1-pinnate with usually 2 pairs of leaflets;
segments 10-20
x
15-30 mm, ovate-oblong, coarsely dentate; upper leaves
sessile, 1-pinnate or 3-lobed)
the terminal segment 8-10
x
30-50(-80) mm,
lanceolate-linear, coarsely serrate, the leaves are covered on both sides with
setose hairs. Umbels 30-40 mm in diameter. Bracts
5-8,
4-6 mm, linear,
strigose. Rays 10-15, 5-30 mm, unequal. Bractoles
5,
2-5 mm,
linear-lanceolate, strigose, shorter than the fruiting pedicels. Sepals 0.5-
1
.O
mm,
triangular, strigose. Petals yellow, radiate, the outer 8-1 3 mm, unequally
2-
lobed, strigose at the abaxial surface. Stamens 2.5 mm long. Styles 1.0-1.3 mm
in fruit, strigose; Stylopodium conical, strigose. Mericarps 60-80 mm in
diameter, broadly elliptical, with thick, corrugated margins, the surface of the
mericarps reticulate when seen by the SEM, and covered with strigose hairs;
392
DAWUD
AL-EISAWI
AND
S.
L.
JURY
primary ribs filiform; vittae 6, 4 dorsal and 2 parallel, commissural. Flowering
May-July. Endemic to Turkey.
SPECIMENS
EXAMINED.
TURKEY: Caria, Vallis Meandri prope Guzelhizar, vi 1842,
Boissier s.n.
(BM, E, G, K, W); Odenisch-Vilayet d’oiden (osi Irdineure), 1933,
Lager s.n.
(BM);
B2
Manisa, Kula, 21 vi 1965,
Coode
€9
Jones
2778
(E).
3.
T.
carmeli
(Labill.) Al-Eisawi &Jury,
comb.
nov.
BASIONYM:
Heracleum carmeli
Labill.,
PI.
Syr.
Dec.,
5
3,
1.1
(1812).
Synelcosciadium carmeli
(Labill.) Boiss.,
Ann.
Sci.
Nut. Bot.
ser. 3,
1:
346 (1844).
TYPE: Palestine, Monte Carmelo,
Labillordibre
(G)
,
Retrorsely setose, 500-1000 mm tall. Stems erect, terete, ridged, dichotomously
branching above, hispid at the base, retrorsely setose above. Basal and cauline
leaves I-pinnate, with 2-3 pairs of leaflets; segments 10-20
x
30-40 mm, ovate,
lanceolate, crenate serrate; upper leaves,
1
-pinnate or 3-lobed, the terminal
segment lanceolate linear; both sides of the leaves covered with setose to
strigose hairs. Umbels 50-100 mm in diameter, open at anthesis, strongly
contracted in fruit. Bracts 3-5, 2-5 mm, linear, unequal, strigose. Rays 3-7,
0-70 mm, very unequal. Bracteoles 5-7, 3-5 mm, linear, strigose. Sepals
0.5- 1.5 mm, triangular, unequal, strigose. Petals white, radiate, the outer
6-8 mm, unequally 2-lobed, the lower surface strigose. Stamens 2-3 mm. Styles
1
mm; stylopodium conical, both strigose. Mericarps 6-8 mm diameter, broadly
elliptical to obovate, with thick, smooth margins and covered with strigose hairs;
primary ridges filiform; vittae
6,
4 dorsal and
2
parallel commissural. Flowering
late April-July.
n
=
10.
Endemic to the coastal areas from Syria south to
northern Palestine.
REPRESENTATIVE
SPECIMENS
EXAMINED.
LEBANON:
Said, 5 vi 1853,
Blanche
69
(K);
Nahr el Kalb, 5
v
1943,
Davis 6084A
(K);
Souk
el Ghurb, 27 vi 1959,
Maitland
519
(K); Hasbaya, 25 vi 1855,
Kotschy
162
(BM,
K);
Beirut, A.U.B. campus,
9 vi 1975,
Al-Eisawi s.n.
PALESTINE:
Carmel, 1846,
Boissier s.n.
(K);
Carmel,
1846,
Pinard s.n.,
(BM,
K).
Jerusalem, Kiriath-Anavim, 26 v 193
1,
Amdursky
153
(BM,
K); Carmel, Wadi Shomariya, 28 vi 1942,
Davis 4544
(K); Hsifa, Carmel,
6
v
1897,
Bornmuller
695, (K,
WU);
El-Huleh, 1863,
Lowne s.n.
(BM, K).
SYRIA:
Laftakia, 5 vi 1884,
Post s.n.
(BM);
Banias and Yarmuk river,
S.
of Saida, 191
1,
Myers
€9
Dinsmore 4949
(K)
.
II.
Subgenus
Ainsworthia
Drude
Plants scabrous-pubescent to villous; sepals usually absent; petals glabrous;
styles and stylopodium glabrous; mericarps covered with vesicular to tubular
hairs; pollen grains tricolporate (except in
T.
cappadocicum
which has
dicolporate pollen).
B.
Section
Condylocarpus
(Hoffm.) DC.
One petal only radiate; vittae 20-22, 10-12 dorsal, 10 commissural.
4.
T.
apulum
L.
Sp.
Pl.:
239
(1
753).
SYNONYMS:
T.
humile
Desf.,
Fl.
Atl.,
1:
235,
t.
38 (1800).
T.
grandgorum
Moench,
Meth.:
78
(1794).
Condylocarpus apulus
Hoffm.,
Gen. Umb.
ed.
2:
202 (1816).
REVISION
OF
THE GENUS
TORDYLUM
393
TYPE:
Herb. Clifford, 90,
Tordylium
3 (BM, syntype).
Stems erect 200-500 mm, terete, striate to ridged, hispid at the base and
scabrous above. Basal and cauline leaves 1-pinnate into 10-15
x
10-15 mm
ovate, crenate segments; upper leaves 3-lobed into dentate-lobate segments.
Umbels 30-70 mm in diameter. Bracts 2-5(-7), 4-10 mm, linear-lanceolate
with ciliate margins. Rays 3-8, 30-50 mm, equal, scabrous. Bracteoles 5,
3-6 mm, linear-lanceolate. Pedicels 4-6 mm, equal. Sepals absent, rarely
minute. Petals white,
only
one outer petal radiate, 4-7 mm, equally 2-lobed,
glabrous. Stamens
1
mm. Styles minute; stylopodium conical depressed.
Mericarps
7-
10 mm diameter, orbicular with thick, corrugated margin, covered
with vesicular hairs on the surface; ridges filiform; vittae 22,
12
dorsal and 10
commissural. Flowering March-May.
2n
=
20. Distributed throughout the
Mediterranean and east to Portugal and north
to
Austria.
REPRESENTATIVE
SPECIMENS
EXAMINED.
ALGERIA:
8
iv 1865,
Tribout
578,
(BM).
AUSTRIA:
Rovinj, 25 v 1953,
Meyer
s.n.
(M); Abruzzenreise, 8 v 1972,
Leute
2347
(W); Sudlich der Orbschaft Promontore,
1
vi 1907,
Janchen
s.n.
(WU).
CYPRUS:
near Orga,
12
iv 1956,
Poore
2636
(K); Hartcha, 9 iv 1950,
Chapman
270
(K).
GREECE:
Prov. Argolis, near Galata, 25
v
1964,
Phitos
1836
(M);
Prov. Attiki, E
of
Mt Penteli,
15
v
1967,
Maria
7999
(M); Corfu, 1896,
Baenitz s.n.
(E); Cos to
Asclepicion,
27
iii 1965,
Davis
40427
(BM, E, K,
W);
Crete, Knossos,
8
iv 1954,
Merxmuller
@
Wiedmann
23182
(M); Dist. of Sphakia, Kurssboden, 18 iv 1904,
DorJlr
528
(WU); Cyclades,
Is.
of
Armorgos,
11
iv 1940,
Davis
2407K
(E);
Dalmatia, Spalato, 1870,
Pichlet
555
(WU);
Dardanelli, 25 vi 1883,
Asherson
319
(BM); Delphi, along the road to Amphissa
just
outside the town, 27 v 1965,
Alma
4595
(W);
Elcusis, 1889,
Hausskenecht s.n.
(E); Ins. Euboea, Ozylithos,
3 v 1965,
Phitos
4039
(M);
Rodhos, Miramar Beach, near Rodhos, 21 iii 1965,
Davis
40281
(E,
K).
ITALY:
Florence,
iv
1965,
Polunin
8202
(E);
Etruria:
Florentia,
17
v 1914,
Fiori
2224
(BM, WU).
SARDINIA:
NE coast, Porto Rafad,
between Palau and Punta Sardegna, 4 iv 1973,
Humphries
&
Richardson
46
(BM,
RNG);
SICILY:
Kalampaka, 23 iv 1896,
Sintenis
265
(E); Toscana, 14 v 1951,
Roessler
462
(M).
LIBYA:
Cireaica, Jebel
El
Akhdar, Ain Belang,
6
v 1959,
Setni
334
(K);
El-Beda, El Madfa,
1
v 1934,
Pampanini
€5'
Pichi-Sermolli
5798
(K).
MALTA:
Mellicha,
c.
200 m,
11
iv 1970,
Davis
50630
(E,
RNG).
PORTUGAL:
Elvas,
Alentejo, village of Boim, 30 iv 1968,
Malato-Beliz
&
Abreu
6236
(M).
ROMANIA:
Via Laurenlina,
28
iv 1902,
White s.n.
(E).
SPAIN:
Badajoz; Almendral,
27
iv 1966,
<ubicarreta
(M); Madrid, Barcarrota, 26 iv 1966,
God9
et
al.
s.n.
(RNG).
TURKEY:
Prov. Aydin, Sultanhisar, between Aydin
&
Nazilli, 3 iv 1956,
Davis
@
Polunin
025586
(E,
K);
Mugla, Bodrum to Karaova,
12
iv
1965,
Davis
40972
(E); Istanbul, 50 km
W
of Istanbul,
1
v 1971,
Lamond
2620
(E); Lydia,
Soghan-Dere, 5 v 1906,
Bornmuller
9544
(WU).
YUGOSLAVIA:
Dalmatia,
Is.
Brazza, 15 iv 1895,
Ginrberger s.n.
(WU); Kroatia, Dubrovnik, 24 iv 1962,
Merxmuller
&?
Wiedmann
23180
(M)
.
C.
Section
Hasselquistia
(L.)
Boiss.
Outer
2
petals radiate; vittae 6, dorsal
4,
all visible, commissural
2,
parallel.
5.
T.
syriacum
L.,
S'.
PI:
239
(1
753).
TYPE:
Herb. Clifford, 90,
Tordylium
1
(BM, syntype); 337.1 (LINN, syntype).
394
DAWUD AL-EISAWI AND
S.
L.
JURY
Scabrous, 150-400 mm tall. Stems erect, terete, ridged, spreading. Leaves up to
150 mm, ovate-oblong in outline, 1-pinnate with 2-3 pairs of leaflets; segments
15-20
x
20-30 mm, ovate to obovate, crenate. Umbels 30-60 mm diameter.
Bracts 3-5, 10-35 mm,
spathulate-oblanceolate,
with scabrous margins,
unequal. Rays 5-10, 5-40 mm, very unequal. Bracteoles 5, 6-30 mm long,
similar to the bracts. Pedicels 0-2 mm. Sepals absent or minute. Petals white,
radiate, the outer 2-3 mm, equally 2-lobed, glabrous. Stamens 1.5 mm; anthers
brown. Styles
1
mm; stylopodium conical depressed. Mericarps 8-12 mm
diameter, orbicular, with thick, rugose margins and covered with vesicular and
tubular hairs; primary ridges filiform; vittae
6,
4
dorsal and
2
commissural.
Flowering March-May.
2n
=
20.
Endemic to the Middle East.
REPRESENTATIVE
SPECIMENS
EXAMINED.
CYPRUS:
Kythraea, 29 iii 1880,
Sintenis
325
(BM, K,
M,
M, WU); Kyrenia Range, above Kythea, 9 iii 1948,
Dube-
Wodley
28
(BM); Agios Philor, 19 ii 1941,
Davis
2222
(E).
LEBANON:
Shemlan,
21-26 iii 1948,
Maitland
210, 244
(E,
K);
Dahr el Litani,
11
iv 1933,
Samuelsson
3221
(K); Amaria, 16 iv 1889,
Bornmiiller
514
(BM);
N
of
Beirut,
27
iii
1933,
Meinertzhugen
s.n.
(BM); Jeb a1 Byblos, 1.5 km
N
of Jeb a1 Byblos, near village
of
Edde,
27
iv 1945, Roessler 5029 (M).
SYRIA:
Mt Nusairy, Bahama, 15 miles
aL’E de Ladikiea, iv 1909,
Haradjian
2805
(E,
K); Antioch,
22
v 1930,
Rogers
0617
(K);
Aleppo,
Rossell
248
(BM); Owasia, 16 iv 1889,
Bornmiiller
514
(WU).
TURKEY:
Prov. Mersin, dist. Tarsus, gorge of Tarsus River between Ulas and
Samlar, 5 iv 1957,
Davis
€9
Hedge
026458
(BM, E,
K);
Prov. Maras, valley
of
River Ceyhan, 15 km
W
of Maras, 3 v 1957,
Davis
€9
Hedge
027457
(BM, E,
K);
Solak Dagh, Zeytun,
11 v 1934,
Balls
€9
Gourlay
1063
(BM, E,
K);
N
of
Anatolia, 1890,
Mainssadjian
235
(M);
Boulouklii near Mersina (Cilicie),
23 iv 1855,
Balansa
578
(E,
K);
Hatay, along the road from Iskenderun to
Antakya, about 5 km E of Belen,
12
v
1967,
Alava
6606
(E,
W);
Cilicia, 1895,
Bemerkungen
30
(WU)
.
6.
T.
hasselquistiae
DC.,
Prodr.,
4:
198 (1830).
TYPE:
Syria, Aleppo, Michaux
(G).
Pubescent, scabrous, 300-500 mm tall. Stems erect, terete, striate to ridged,
pubescent at the base, scabrous above. Basal leaves 1-pinnate, with long
petioles; segments 13-25
x
15-30 mm, cordate, ovate, crenate; upper leaves
sessile, 1-pinnate or 3-lobed, with ovate to lanceolate segments, both faces
of
the
leaves with soft, appressed hairs. Umbels 40-60 mm diameter. Bracts 10-15,
8-15 mm long, filiform, pubescent, equal. Rays 8-20, 5-40 mm long, very much
equal. Bracteoles 5-8, 5- 15 mm long, unequal, longer than the fruiting pedicels.
Sepals absent. Petals white or pinkish, radiate, the outer 4-6 mm, equally
to
subequally 2-lobed, glabrous. Stamens
1.5
mm. Styles
1
mm in
fruit;
stylopodium conical depressed. Mericarps 6-8 mm, with thick, corrugated
margin and covered with vesicular hairs; primary ridges filiform; vittae
6,
4
dorsal and
2
commissural, the dorsal all visible, the commissural parallel.
Flowering March-May. Endemic to Syria and Turkey.
REPRESENTATIVE
SPECIMENS
EXAMINED.
SYRIA:
Aleppo, orient,
Michaux s.n.
(G)
,
19301,
Aucher-Eloy
3646
(BM, G); Biredschik,
18
v
1865,
Huussknecht s.n.
(G,
K);
Aintab,
v
1889,
Post
s.n.
(BM); Antiochia, Kasr el Banal, 39 v 1933,
Sumuelsson
REVISION
OF
THE
GENUS
TORDYLUM
5515
(K).
TURKEY:
Prov. Hatay, dist. Antakya, Iskunderun-Antakya W of Amik
Gol,
25 iv 1957,
Davis
@
Hedge
027131
(BM,
E);
Urfa, northern edge of the
town, 26 v 1967,
Alaua
6718
(E,
W); Adiyaman, along the road to Maras,
4 vi 1968,
Alava
6953
(E,
W);
Hatay, C6, Belen Pass, 14
iv
1966,
Albury
et
al.
748,
702
(BM, K); Hatay, Kirikham to Haman, 6 v 1965,
Coode
€3'
Jones
577
(E).
7.
T,
trachycarpum
(Boiss.) Al-Eisawi &Jury,
comb.
nov.
SYNONYMS:
Ainsworthia trachycarpa
Boiss.,
Biagn.
ser.
1
(10)
:
43
(
1849).
395
T.
cordatum
subsp.
trachycarpum
(Boiss.) Holmboe,
Stud. Veg.
Cyprus:
141 (1914).
T.
trachycarpum
Holmboe ex Prain in
Index Kewensis
Suppl.
5:
26
1
(
192
1
)
,
nomen
T.
byzantinum
(Azn.) Hayek,
Prodr. F1. Penins. Balcan.,
1:
1045 (1927).
Ainsworthia byzantina
Azn.,
Bull.
SOL.
Bot.
Fr.,
44:
170 (1897).
ivalidum
(Art, 34).
TYPE:
Palestine,
In
collibus Judeae prope Hierosolymam et St. Saba, iv-v 1846,
Stems erect, terete, ridged, densely scabrous, 200-800 mm. Basal and cauline
leaves simple or I-pinnate; segments 15-40
x
20-60 mm, cordate, crenate; upper
leaves I-pinnate or 3-lobed with ovate-cordate, crenate to dissected segments;
both faces of the leaves covered with soft, appressed hairs. Umbels 40-100 mm
diameter. Bracts 10-16(-20), 12-18 mm, filiform linear, setose-scabrous. Rays
15-30(-40), 20-60 mm long, unequal, scabrous. Bracteoles 5-8, 5-15 mm long,
unequal, scabrous. Pedicels 3-10 mm long. Sepals absent. Petals white, radiate,
the outer 5-8 mm long, unequally 2-lobed, glabrous. Stamens 1.5 mm long.
Styles
0.5
mm; stylopodium conical depressed. Mericarps 3-6 mm diameter,
orbicular with soft, thick margins and covered with vesicular hairs; primary
ridges filiform, vittae 6, 4 dorsal and
2
commissural, the commissural vittae
parallel. Flowering March-May.
n
=
7,
8, 9, 10. Endemic to the Middle East.
REPRESENTATIVE
SPECIMENS
EXAMINED.
CYPRUS:
Kythrae,
v
1880,
Sintenis
324
(K,
WU); Bella paise,
7
ii 1948,
Ingram
21
(BM);
Ayios, Amurosios, 30 iii 1974,
Meikle
4030
(K); Ayio Irini (Mophou) Grand of Kings Potrous,
12
iii 1941,
Davis
2556
(E); Near Ayia Irini, 26 iii 1962,
Meikle
2394
(W).
IRAQ:
Mod,
Aqra, 30 v 1946,
Rawi
11337
(K);
Shaqlawah, 13 v 1956,
Haines
764
(E,
K).
PALESTINE:
Haifa, Mt Carmel,
6
v 1897,
Bornmuller
687,
69Oa
(BM,
E,
K,
W,
WU); Wadi Kafrein,
20
v
191
1,
Myers
€3'
Dinsmore
8074
(K,
K); Jerusalem,
Talpioth, 15 v 1933,
Zohary
273
(BM,
E,
K,
W).
SYRIA:
Amaro prope Beilan,
1862,
Kotschy
261
(G, W); Tel Kalakh, 5 iv 1945,
Trench s.n.
(BM).
TURKEY:
Constantinople, v 1873,
De
Fontenay s.n.
(G);
Istanbul,
12
vi 1896,
Aiyzavour s.n.
(E, W); Istanbul, 20
vi
1897,
Nemetz s.n.
(WU); Prov. Diyarbakir, Cinar,
8
km
from Diyarbakir, 31
v
1957,
Davis
@
Hedge
028778,
(E,
K);
Plaine de Mersina
(Cilicie), 1855,
Balansa
570
(BM,
E,
K,
W).
Boissier
s.n.
(G)
.
8.
T.
pustulosurn
Boiss.,
Fl.
Or.
Suppl.:
268 (1888).
TYPE:
Turkey, ad rivulos prope Anamour Ciliciae Tracheae, 1872,
Pironin
(BM,
Pubescent to scabrous, 200-600 mm tall. Stems erect, terete, striate to ridged,
pubescent at the base, scabrous above. Basal leaves have long petioles, I-pinnate
G,
K).
396
DAWUD AL-EISAWI AND
S.
L.
JURY
with usually a pair of leaflets; segments 10-15
x
20-30 mm, cordate to ovate,
crenate; upper leaves sessile, with cordate-ovate, crenate, cuspidata segments.
Umbels 30-50 mm diameter. Bracts 7-12, 6-8 mm, filiform, pubescent. Rays
10-15, 8-25 mm, unequal. Bracteoles 3-5, 3-7 mm, unequal. Sepals absent.
Petals white or pinkish, radiate, the outer 3-4 mm, unequally 2-lobed, glabrous.
Stamens 1.3 mm. Styles 0.5 mm in fruit; stylopodium conical depressed.
Mericarps 4-6 mm, suborbicular to broadly elliptical, with thick, smooth
margins, and covered with vesicular hairs; primary ridges filiform; vittae 6, 4
dorsal and 2 parallel, commissural. Flowering March-May. Endemic to
Turkey.
REPRESENTATIVE
SPECIMENS
EXAMINED.
TURKEY:
Anamour, 1872,
Pironin 85
(BM,
G, K); Iter Orientale, 1859,
Xotschr 691
(G);
Anamur, Anamur-Gilindire,
14 v 1956,
Davis
€b’
Polunin 025999
(BM,
E,
K); Icel, Silifke,
SW
of Silifke, near
the old Roman ruins, 16 v 1967,
Alava 6648
(E,
G,
W);
Icel, 16 v 1967,
Alava
6650
(E,
W);
Cukurova, Kis Kalesi Beah, 90 km
W
of
Tarsus, 19 v 1967,
Deaver
7-15,?
(E)
.
9.
T.
lanatum
(Boiss.) Boiss.,
Fl.
Or.,
2:
1030 (1872).
SYNONYM:
Hasselguistia lanata
Boiss.,
Ann. Sci. Nut., ser.
3,
1:
347 (1844)
TYPE:
Turkey, Caria, 1943,
Pinard
(G,
K).
Stems densely villous, to 400 mm tall, erect, terete, spreading. Leaves up to
150 mm long, ovate-oblong in outline, l(-3) pinnate with ovate-oblong, lobate
to dentate segments. Umbels 40-70 mm diameter. Bracts 0-2, 1-3 mm, linear,
pubescent. Rays 2-8, 15-40 mm long, unequal to subequal, scabrous. Bracteoles
0-2, 1-2 mm, linear. Pedicels 4-6 mm, subequal. Sepals absent, rarely minute.
Petals white, radiate, the outer 20.0-3.5 mm, equally to subequally 2-lobed,
glabrous. Stamens 0.7 mm. Styles
0.7
mm. Styles 0.3 mm; stylopodium conical
depressed. Mericarps dimorphic; semispherical, 4-5 mm diameter and dorsally
compressed or 7-10 mm in diameter, grey-blue with thick, smooth to rarely
rugose margins and covered with vesicular hairs; primary ridges filiform; vittae
6, 4 dorsal and
2
commissural. Flowering March-May. Endemic to Turkey.
REPRESENTATIVE
SPECIMENS
EXAMINED.
TURKEY:
Caria, 1943,
Pinard s.n.
(G,
K);
Tcheltikchi, v 1845,
Heldreich s.n.
(G,
K); Lycia, Elmalu, 1860,
Bourgeau 120
(E,
G,
K); Pisidia, Tcheltikchi, v 1849,
Heldreich s.n.
(BM, G); Elmalu, Aliolagh,
29 vi 1883,
Pichler 299
(G,
WU); Liguriae,
11
v
1890,
Barla
@
Penig s.n.
(K,
M);
Anatalay, 5 km
S
of Kizilkaya on the way from Korkuteli to Burdur,
19
vi 1966,
Alava
€b’
Bocquel5313
(E, W); Anatalya, Sutlegen,
11
v 1964,
Jackson
5049
(E)
.
10.
T.
aegyptiacum
(L.) Lam.,
Tabl.
En&. Meth. Bot., l(2):
336
t.
193
(1 792).
SYNONYMS:
Hasselguistia aegyptiaca
L.,
Cent. P1. 1:
9 (1755),
“in Amoe. Acad. Bot., 4:
TYPE:
Hasselquist s.n.
(348.2, LINN, syntype).
Stems erect, 150-400 mm tall, terete, striate to ridged, spreading, villous at the
base and scabrous above. Leaves up
to
150 mm long, ovate-oblong in outline,
270
(1
788)”.
REVISION
OF
THE
GENUS
TORDYLCJM
397
l(-3) pinnate with oblong, lobate-dentate segments. Umbels (40-)SO-100 mm
diameter, often the central umbellule strate with purple to black head. Bracts
1-5(-7), 2-10 mm, linear, pubescent. Rays 6-16, 10-65 mm long, very
unequal, contracted in fruit. Bracteoles 3-5, 2-6 mm, filiform-linear. Pedicels
5.0-1.3 mm, filiform, unequal. Sepels absent or minute. Petals white, radiate;
the outer
5-7
mm, equally to subequally 2-lobed, glabrous or rarely villous at
the base. Stamens 1.5-2.0 mm, with greenish anthers. Styles
1
mm; stylopodium
conical depressed. Mericarps dimorphic; semispherical, 4-5 mm diameter and
dorsally compressed or 7-10 mm in diameter, brown with thick, rugose margins
and covered with vesicular hairs on the surface; primary ridges filiform; vittae 6,
4 dorsal, and 2 commissural. Flowering March-May.
n
=
10. Endemic to the
Middle East, Egypt and Libya.
REPRESENTATIVE
SPECIMENS
EXAMINED. CYPRUS:
Nicosia, along the line at Nicosia
Rly Station, iii 1930,
Norman
452
(BM); Gostria dist., Famagusta, 27 ii 1941,
Davis
2413
(E);
Kardemeros, 9 iv 1967,
Merton
ARI 107
(W).
EGYPT:
Alexandria,
27 vi 1910,
Muschler
s.n.
(K);
Mariut, 14
iii
1913,
Bolland
13591
(K).
IRAQ:
Irbil,
18 v 1958,
Haines
1408
(E, K); MOUSUI, Kursi, Jebel Sinjar, 23 v 1948,
Gillett
10867
(K);
MOSUI, Tioris Plain, 21 iii
1036,
Alou
109
(BM); Kirkuk,
Sulaimanieya, 23 v 1951,
Rawi
21488
(K).
LEBANON:
Saida, 6 iv 1853,
Blanche
18
(E,
K);
Amioun, 23 v 1959,
Polunin
5370
(E,
K); Jounieh,
N
of Beirut,
27 iii 1933,
Meinertzhagen s.n.
(BM); Beirut, 15 iv 1910,
Permy
s.n.
(WU).
LIBYA:
Tripolis, 24 iv 1877,
Blanche s.n.
(WU).
PALESTINE:
Jaffa, 24 iii 1897,
Bornmuller
692
(K, WU); Ramleh, Kobab, 13 v 1932,
Dinsmore
10550
(K). Palestine, 1846,
Pinard
s.n.
(BM,
K);
1846,
Boiss. s.n.
(K);
Lake Hula,
8
iv 1945,
Cappen s.n.
(BM);
Samaria, between Bat Shelomo and Eliakim, 17 iv 1957,
Lurch
&
Crizi
674
(BM,
M); Latrone, 240 m, 26 iv 1905,
Dinsmore
25.50
(E); Northern Negen, 10 km N of
Beer Sheba,
18
iv
1967,
Alava
6153
(W).
SYRIA:
Aboudouhour, near Hammah,
6-12
v 1968,
Haradjian
1975
(E,
K); Aleppo, v 1841,
Kelschy s.n.
(BM,
K);
Bludan,
8
vi 1855,
Xotschy
86
(BM); Beka, near Rajak, 16
v
1910,
Bornmiiller
11815
(BM); Zebdani, 6 vi 1943,
Davis
6135A
(K); Jebel Druze, v 1956,
Werkmeisher s.n.
(M).
TURKEY:
Prov. Urfa, Akcalkale, 10 km from
Urfa,l7 v 1957,
Davis
@
Hedge
D28149
(BM,
E,
K); Prov. Mardin, Mardin-
Nusaybin, 5-10 km from Nusaybin, 22 v 1957,
Davis
@
Hedge
028400
(BM,
K);
Urfa, northern edge of the town, 24 v 1967,
Alava
6717
(E).
11.
T.
cordatum
(Jacq.) Poir. in Lam.,
En&. Meth.
Bot.,
7:
712 (1806).
SYNONYM:
Hasselquistia cordata
Jacq.,
Hort. Vindob.,
2:
91,
t.
193 (1772).
Ainsworthia cordata
(Jacq.) Boiss.,
Ann. Sci. Nat., ser.
3,
1:
343 (1844).
Ainsworthia carmeli
Boiss.,
Diagn. ser.
1 (10)
:
44
(
1849).
TYPE:
Palestine,
Jacq.
47,
(LINN: 348.3).
Stems erect, 200-500 mm tall, terete, striate to ridged, hispid to scabrous. Basal
leaves simple, cordate crenate; cauline leaves simple or 3-lobed with
ovate-cordate, crenate to serrate segments; upper leaves sessile, simple, cordate
to lanceolate, the leaves with
soft,
appressed hairs
on
both sides. Umbels
30-50 mm diameter. Bracts
10-18,
10-15 mm, filiform-linear, covered with
long, soft hairs. Bracteoles 5-7, 4-12 mm, unequal, covered with long, soft hairs.
Pedicels
3-7
mm, unequal. Sepals absent. Petals white, radiate, the outer
398
DAWUD AL-EISAWI AND
S.
L.
JURY
4-6 mm, unequally 2-lobed. Stamens 1.5 mm. Styles
0.5
mm; stylopodium
depressed. Mericarps dimorphic; semispherical, 2-3 mm in diameter; dorsally
compressed or 4-6 mm in diameter with thin, smooth margins and sparse
vesicular hairs
on
the surface; primary ridges filiform; vittae 6, 4 dorsal and
2
commissural. Flowering March-May.
n
=
10. Endemic to South Lebanon and
Palestine.
REPRESENTATIVE
SPECIMENS
EXAMINED.
LEBANON: Wadi Hammana,
6
vi 19 10,
Bornmiiller
II813B
(BM,
E).
PALESTINE: Mt Carmel, In a wadi on the north
facing slope of Mt Carmel, 21 iv 1967,
Alavu
5364
(W);
Mt Carmel,
1
v 1909,
Myers
€5'
Dinsmore
I387
(E, M); Hafa, Mt Carmel, 6 v 1897,
Bornmuller
689
(E,
K,
WU); Haifa, Mt Carmel, 15 iv 1897,
Bornmuller
690
(BM,
E,
K,
WU); Mt
Carmel, 1846,
Pinard
s.n.
(BM,
K);
Mt Carmel,
1846
Boissier
s.n.
(E,
G,
K).
12.
T.
elegans
(Boiss.
&
Bal.) Alava
&
Hub.-Mor.,
Notes
R.
B.
G.
Edin.,
31:
117
SYNONYM:
Ainsworthia elegans
Boiss.
&
Bal.,
Diagn.
ser.
S
(5):
100 (1856).
TYPE: Turkey, in regione montana Tauri Cilicici prope Gulek Boghas,
Stems erect, 200-400 mm tall, terete, striate to angular, many branched,
pubescent to scabrous. Basal and cauline leaves 1-pinnate with 2-3 pairs of
leaflets; segments 10-15
x
10-15 mm, ovate, lobate crenate; upper leaves 3-
lobed into lanceolate, entire, mucronate to cuspidate segments. Umbels
30-60 mm in diameter. Bracts 8-14, 5-10 mm, filiform, pubescent. Rays 10-16,
15-50 mm, unequal,
'
scabrous. Bracteoles
1-5,
4-6 mm, filiform. Pedicels
3-10 mm, unequal. Sepals absent. Petals white, radiate, the outer 2-3 mm,
equally or subequally 2-lobed. Stamens
1
.O-1.3 mm; anthers brown. Styles
0.7 mm; stylopodium conical depressed. Mericarps dimorphic; semispherical,
3-4 mm in diameter; dorsally flattened, 6-8 mm in diameter, orbicular, with
thin, smooth margins, glabrous or covered with vesicular to tubular hairs on the
surface; primary ridges filiform; vittae 6, 4 dorsal (the 2 central are broken) and
2 commissural. Flowering March-May. Endemic to Syria and Turkey.
REPRESENTATIVE
SPECIMENS
EXAMINED.
SYRIA:
Mounts Amanus, region de
Duldul, vi 1908,
Hurudjiun
2364
(E,
K).
TURKEY: Ciliciennes, Taurus, prts du
defile des Portes Ciliciennes, 25 vi 1855,
Balansa
569
(BM,
G,
K,
W);
Adana,
Osmaniye, Toprakkale,
20
iv 1957,
Davis
€5'
Hedge
026908
(BM,
K);
Adana,
dist. Bahee Haruniye, 17 vi 1957,
Davis
€5'
Hedge
026754
(E);
Adana,
Toprakkale, 19v 1967,
Alava
66Y4
(E,
W);
Cilicia, 29 iv 1859,
Kotscb
69
(G,
W).
Adana, 0.5 km below Aurdagi Pass, 14 v 1966,
Albury
tY
Watson
993
(K).
(1971).
25 vi 1855,
Balansa
569,
(BM,
G,
K,
W).
D.
Section
Univittata
Drude.
Outer
2
petals radiate; vittae 6, dorsal 4, only the
2
central visible or not,
commissural 2, divergent (except in
I.
cappadiocicum).
13.
T.
cappadocicum
Boiss.,
Ann. Sci.
Nut.,
ser.
3
(I):
349 (1844).
SYNONYM:
1.
persicum
Boiss.
&
Hausskn,
FZ.
Or.,
2:
1032 (1872).
TYPE:
Turkey, Cappadocia ad Euphrateum,
Aucher-Eloy
3632-2642
(G,
K)
REVISION
OF
THE
GENUS
TORDYLUM
Stems erect, 150-500 mm tall, terete, striate to ridged, dicotomously branched,
rubescent to scabrous. Basal and cauline leaves 1-pinnate with 2-4 pairs of thick
leaflets; segments 15-20
x
30-40 mm, ovate-cordate, crenate, serate; upper
leaves 1-pinnate, or 3-lobed with ovate-cordate segments, both sides of the
leaves with soft, appressed hairs. Umbels 30-70 mm in diameter. Bracts 5-8,
5-10 mm, lanceolate with membranous, ciliate margins. Rays 10-25,
0.5-5.0 mm, unequal, the central ray usually sterile, short, thick, densely hairy.
Bracteoles similar to the bracts. Pedicels 2-6 mm, unequal. Sepals absent or
minute, membranous. Petals yellow, radiate, the outer 5-8 mm, equally or
subequally 2-lobed, glabrous or with soft hairs at the base. Stamens 2 mm.
Styles 0.7 mm; stylopodium conical depressed. Mericarps 4.5-6.0 mm broadly
elliptical with thick, corrugated margins and covered with tubular hairs;
primary ridges filiform; vittae 6, 4 dorsal and 2 commissural, the lateral dorsal
vittae invisible, the commissural vittae parallel. Flowering March-May.
Endemic to Turkey, Iran and Kuwait.
REPRESENTATIVE
SPECIMENS
EXAMINED.
IRAN:
In arenosis, Buschir, 1868,
Huussknecht
s.n.
(G); Buschir,
17
iv 1885,
Starfs.n.
(K); Daleki, 25 iv 1885,
Stupf
s.n.
(K);
Borashdkum, 27 iv 1885,
Stupf
s.n.
(K); Kalafabad, Ahwas
to
Bebehen, 500 ft,
26 iii 1962,
Furse
1211
(K).
KUWAIT:
Adaliyah,
8
iv 1935,
Dickson
245
(K).
TURKEY:
Cappadocia, Euphratem,
Aucher-Eloy
3642
(G,
K); Orfa, near Velsch
Kuru,
22
v
1888,
Sintenis
833
(K,
WU).
399
14.
T.
hirtocarpum
Candargy,
Bull.
Soc.
Bot.
Fr.,
44:
159 (1897).
TYPE:
Greece, Olympus, Palaeocastron, Petrovuni.
Stems erect, 100-200 mm tall, terete, finely striate, pubescent. Basal and cauline
leaves simple or 1-pinnate; segments ovate, crenate; upper leaves 3-lobed with
lanceolate, crenate to entire segments. Umbels 10-15 mm in diameter. Bracts
2-5, 2-6 mm, unequal, lanceolate with ciliate margins. Rays 2-4, 5-10 mm,
equal, Bracteoles 5-6, 2.5 mm, lanceolate with ciliate margins. Pedicels
1-2 mm. Sepals absent. Petals white, radiate, the outer 2-3 mm, unequally 2-
lobed, glabrous. Stamens 1.5-2.0 mm. Styles 0.7-1
.O
mm long; stylopodium
conical depressed. Mericarps 4.5-6.0 mm, ovate with thick, corrugated margins
and covered with tubular hairs; primary ridges filiform; vittae 6, 4 dorsal and 2
commissural, the lateral dorsal vittae invisible, the commissural divergent.
Flowering March-May. Endemic to Greece and Turkey.
REPRESENTATIVE
SPECIMENS
EXAMINED.
GREECE:
Insula Phurni, 25-26 iv
Rechinger
jil.
4694
(BM,
K);
Kos,
Oros Dikairo, 13 v 197
1,
Brenun
11142
TURKEY:
Ismir: Cesme, 26 iv 1965,
Duuis
41835
(K);
Loryma, Pisidien,
Luschun
s.n.
(WU).
934,
(K).
881,
15.
T.
oflcinale
L.,
Sp.
Pl.:
239 (1753).
TYPE:
337.2
(LINN,
syntype).
Stems erect, 200-600 mm tall, terete, striate to ridged, dichotomously branched,
pubescent to scabrous. Basal leaves simple or 1-pinnate, with 2-3 pairs of
400
DAWUD AL-EISAWI AND
S.
L.
JURY
leaflets; segments ovate crenate; upper leaves usually 3-lobed, the terminal
segment lanceolate to linear, coarsely serrate; all leaves with soft, appressed
hairs. Umbels 30-50 mm in diameter. Bracts 10-16,
1
.O-1.7
mm, filiform-linear,
with scabrous to setose margins. Rays 10-20, 3-20 mm, very unequal.
Bracteoles 5-7, 3-13 mm, unequal, much longer than the fruiting pedicels.
Sepals absent. Petals yellow (rarely reddish), radiate, the outer 5-10 mm,
unequally 2-lobed, glabrous. Stamens
2
mm. Styles 0.5 mm; stylopodium
conical depressed. Mericarps 2.5-3.5 mm, ovate, with thick, corrugated margins
and covered with vesicular hairs on the surface; primary ridges filiform; vittae 6,
4 dorsal and 2 commissural, the lateral dorsal vittae invisible, the commissural
vittae divergent. Flowering March-May. 2n
=
18.
Endemic to Italy and the
Balkan Peninsula although given in the
Flora
of
Turkey
as possibly occurring on
the basis
of
a reference in the literature. We have not seen any specimens
of
this
species from Turkey.
REPRESENTATIVE
SPECIMENS
EXAMINED.
ALBANIA:
Dist. Gjinokastre, 7 vi 1933,
Alston
B
Sandwith
1445
(BM,
K);
Skoplje (Uskub), 19 v 1905,
Adamovics.n.
(WU).
GREECE:
Thessaliae, Karditza, 1889,
Haussknecht
s.n.
(E)
;
Thessaliae, near
Pharsalum, 13-
18
vi 1885
Heldreich
s.n.
(WU); Crete, Kissamos, Lieux arides,
12 v 1884,
Reverchon
254
(E, M, WU); Dalmatia, near Spalato, v 1870,
Pichler
s.n.
(WU); Prov. Attiki, dist. Klissura, inter Mesolongin ad Agriniou, 22 v 1964,
Phitos
1579
(M); Peloponnese; Megalopolis, near the theatre, 15 vi 1966,
Whitefoord
I51
(BM).
ITALY:
Topygia, 29
v
1875,
Porta
&
Riga
105
(WU);
Apulia-Gargano, Vieste, Coppetelle,
22
v 1913,
Fiori
2125
(BM, K, WU);
Peschici, Gargano Peninsula, 5 iv 1960,
Brummitt
985
(K).
YUGOSLAVIA:
Dalmatia, 15 v 1905,
Crawford
41
((E); Dalmatia Mariana, 27 v 1906,
Janchen
s.n.
Dalmatian Coast, northern outskirts of Sibenik, 12
v
1969,
Brumitt
81
(K);
Dalmatia, near Makarsha, v 1935,
Gilliat
B
Smith
3476
(K).
16.
T.
brachytaenium
Boiss.
&
Heldr.,
Diagn., ser.
1(10):
45 (1849).
TYPE:
Turkey, in Pamphyliae collibus ad Kourmalu prop Adalia, iv 1845,
Stems erect, 300-500 mm tall, terete, striate, dichotomously branched,
pubescent to minutely scabrous above. Basal leaves
1
-pinnate with long petioles;
segments 13-20
x
20-54 mm, ovate, crenate; upper leaves sessile with ovate
to
lanceolate, dentate segments; all leaves with soft, appressed hairs. Umbels
30-40 mm in diameter. Bracts 558 mm, lanceolate with ciliate margins,
unequal. Rays 10-16, 5-20 mm, unequal, scabrous. Bracteoles 5-7, 2-7 mm,
lanceolate, with ciliate margins, unequal. Sepals absent. Petals radiate, white to
yellow, the outer 6-8 mm, unequally 2-lobed, glabrous, rarely with soft hairs at
the base. Stamens 1.5 mm. Styles 0.5 mm; stylopodium conical depressed with
undulate margin. Mericarps 30-40
x
40-55 mm, semiorbicular with thick,
smooth margins, (the immature mericarps has a corrugated margin near the
stylopodium, which disappears at maturity), and covered with vesicular hairs on
the surface; primary ridges filiform; vittae 6, 4 dorsal and 2 commissural, the
lateral dorsal and 2 commissural, the lateral dorsal vittae are invisible, hidden
under the wings. Flowering April. Endemic
to
Turkey.
Heldreich
(BM,
E,
G,
K,
M,
WAG).
REVISION
OF
THE
GENUS
TORDTLUM
40
1
REPRESENTATIVE
SPECIMENS
EXAMINED.
This species is known only from the type
collection.
17.
T.
pestaloxxae
Boiss.,
Diagn., ser.
1 (10):
45 (1849).
SYNONYM:
T.
aegaeum
Runem.,
Bot.
Not.,
121
(2):
253 (1968).
TYPE:
Turkey, prope Elmal Lyciae ubi,
Pestalozza
(G).
Stems erect, 100-300 mm tall, terete, striate to ridged, pubescent at the base,
scabrous above. Lower leaves simple or 1-pinnate with
1-2
pairs of leaflets;
segments ovate, crenate to dentate; upper leaves I-pinnate or 3-lobed with
ovate-lanceolate, terminal segments, both leaf surfaces covered with soft,
appressed hairs. Umbels 20-30 mm in diameter. Bracts 5-8, 4-9 mm, linear to
rarely lanceolate. Rays 5-12, 3-15 mm, unequal. Bracteoles 5-8, 4-8 mm,
linear, equal or longer than the fruiting pedicels. Sepals absent. Petals white
to
yellow, radiating, the outer 5-7 mm, unequally 2-lobed, glabrous. Stamens
1.5
mm. Styles
1
mm in fruit; stylopodium conical depressed. Mericarps
5-6 mm, ovate with thick, corrugated margins and covered with vesicular hairs;
primary ridges filiform; vittae 6, 4 dorsal and
2
commissural, the dorsal vittae
usually all invisible, except the central, which are sometimes shown near the
stylopodial end,
the commissural vittae divergent. Flowering March-May.
2n
=
20. Endemic
to
Greece and Turkey.
REPRESENTATIVE
SPECIMENS
EXAMINED,
GREECE:
Aegean Islands, Astipalea,
between Maltesana and Vriseu Pund,
11
v 1960,
Runemark
8
Nordenstam
13542
(LUND); Rhodes, on Mountain pophet Elias near Salakos, 11 v 1935,
Rechinger
Jl.
7116
(BM); Rhodes Batida, 21 v 1870,
Bourgeau
59
(BM, E, G,
K);
K rionero,
'
1
vi 1893,
Grimburg
493
(WU);
Chois, 4 km
S
of Pyrgi,
18
v 1966,
Ludtke
440
(M); Insida, Paros near Naussa, 6
v
1898,
Leonis
76,
(WU);
E of
Kos,
11
iv 1974,
Butler
18161
(M).
TURKEY:
Elmalu, 1846,
Pestalozza s.n.
(G);
Smyrne,
18-30 v 1854,
Balansa
50
(BM,
K,
G, WAG); Adalia (Pamphylia), 14
iv
1860,
Bourgeau
I19
(G, E,
K);
Ismir, Kemalpasa, Karabel, 25 v 1966,
Alava
4758
(E,
W);
Mugla, Marmanis, Sogul to Bozburun, 15 v 1965,
Davis
41198
(E,
K);
Ismir, Kenalpasa, 6 vi 1966,
Alava
5046
(E, W); Lydia, Sipgli, 19 v 1906,
Bornmiiller
9514
(K); Anatalya, Dudenbas,
9
km
N
of Antalya, 1944,
Hennipman
et
al.
407
(WAG).
18.
T.
ebracteatum
Al-Eisawi &Jury,
sp.
nov.
Ab
Tor&io hirtocarpo
Cand. caule elatiore bractearum absentia, petalis minimis,
staminibus et stylis parvis differt.
Stem erect to ascending, 300
mm
tall, terete, finely striate, many branched,
pubescent
to
scabrous. All leaves simple, cordate, crenate to dentate, both sides
covered with soft, appressed hairs. Umbels 10-20 mm in diameter. Bracts
absent. Rays 2-3, 1.3-1.8 mm long, equal, scabrous. Bracteoles
5,
2-5 mm,
unequal, lanceolate. Pedicels 0.5-3.0 mm, unequal. Sepals absent. Petals white,
minute, the
outer
1
mm, 2-lobed. Stamens
0.7
mm. Styles 0.5 mm in fruit;
stylopodium conical depressed. Mericarps 5-6 mm, ovate with thick, corrugated
margins, covered with vesicular hairs; primary ridges filiform; vittae 6, 4 dorsal
402
DAWUD AGEISAWI AND
S.
L. JURY
and
2
commissural, the dorsal vittae often all invisible, the commissural
incomplete, divergent. Flowering and fruiting June. Endemic to Turkey.
TYPE:
Turkey Prov. Mugla, limestone sea-cliffs just before entrance of Bozburn
The species
is
only known from its type collection.
Harbour,
50-100
m,
8
vi 1962,
Dudley
035477
(E).
Species excluded from
Tordylium
L.
Tordylium absinthifolium
Pers.
=
Xosima absinthifolia
(Vent.) Link.
T.
anthriscus
L.
=
Torilisjaponica
(Houtt.) DC
T.
artedia
Crantz
=
Artedia squamata
L.
T.
aucheri
Jaub.
&
Spach.
=
Ormosciadium aucheri Boiss.
T,
jiodosum
L.
=
Torilis nodosa
(L.)
Gaertner
T.
peregrinum
L.
=
Capnophyllum peregrinum
(L.) Lag.
7.
latifolium
L.
=
Turgenia
latiflia
(L.) Hoffm.
T.
komarovii
Mandenova
=
Mandenouia komarovii
(Manden.) Alava
The genus
Mandenouia
was described by Alava in (1975), monotypic, based on
Tordylium komarovii
Mandenova.
It
differs from
Tordylium
by being perennial
with fusiform roots, having ternate leaves, lacking bracts and bracteoles, the
rays being equal in length and the mericarps having vittae which do not run the
entire length
of
the mericarp.
APPENDIX: INDEX TO SPECIES
Ainsworthia byrantina
Azn.
=
7
. . .
A. carmeli
Boiss.
=
11
.
,
.
.
A. cordata
(Jacq.) Boiss.
=
11
.
. .
A.
elegans
Boiss.
&
Bal.
=
12
,
. .
A.
trachycarpa
Boiss.
=
7
. . .
.
Condylocarpus apulus
Hoffm.
=
4.
. .
Hasselquistia aegyptiaca
L.
=
10
. . .
H. cordata
Jacq.
=
11
. . .
.
H. lanata
Boiss.
=
9
.
. .
.
.
Heracleum carmcli
Labill.
=
3
. .
.
Synelcosciadium carmeli
(Labill.) Boiss.
=
3
.
Tordylium aegacum
Runem.
=
17
.
.
7.
ae&ptiacum
(L.) Lam.
10 .
.
.
T.
apulum
L. 4.
.
. .
. .
T.
brachytaenium
Boiss.
&
Heldr. 16
.
.
7.
bycantinurn
(Azn.) Hayek
=
7
. .
7.
cappadocicum
Boiss. 13
. .
.
.
T.
carmeli
(Labill.) Al-Eisawi &Jury 3
.
T.
cordatum
(Jacq.) Poir. in Lam.
11
.
.
395
7.
cordatum
(Jacq.) Poir. in Lam.
subsp.
.
397
trachycarpum
(Boiss.)
Holmboe
=
7
.
.
395
.
397
'I:
ebracteatum
Al-Eisawi
&Jury
18
. . .
401
.
398
7.
elegans
(Boiss.
&
Bal.) Alava
&
Hub.-
.
395
Mor.
12
.
.
.
. .
. .
398
.
392
T.grandtj7orum
Moench
=
4
.
. . .
392
.
396
T.
hasselguistiae
DC. 6
. .
.
.
.
394
.
397
7.
hirtocarpum
Cand. 14
. . .
.
.
399
.
396
T.
humile
Desf
=
4
. . .
.
.
.
392
.
392
T.
lanatum
(Boiss.) Boiss. 9
.
. .
.
396
.
392
7.
macropetalum
Boiss.
=
2.
. .
.
.
391
.
401
1.
maximum
L.
=
1
.
,
.
.
,
.
390
.
396
T.
oJicinale
L. 15
.
.
.
.
.
.
399
.
392
1.
persicum
Boiss.
&
Hausskn.
=
13
. .
.
398
.
400
7.
pestalouae
Boiss.
17
. .
.
.
.
401
.
395
7.
pustulosum
Boiss. 8
.
.
.
. .
395
.
398
7.
syriacum
L. 5
.
. . .
.
.
393
.
392
T.
trachycarpum
(Boiss.) Al-Eisawi &Jury
7
.
395
.
397
T.
trachycarpum
Holmboe ex Prain
=
7
.
.
395
ACKNOWLEDGEMENTS
We wish
to
thank the University of Reading for facilities and Professor
V.
H
Heywood for help and advice. We especially thank Dr C.
E.
Jarvis for help with
the typification of the Linnean species.
REVISION OF THE GENUS
TORDI‘LUM
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