Camelina neglecta (Brassicaceae, Camelineae), a new diploid species from Europe
Author
Brock, Jordan R.
Author
Mandakova, Terezie
Author
Lysak, Martin A.
Author
Al-Shehbaz, Ihsan A.
text
PhytoKeys
2019
115
51
57
http://dx.doi.org/10.3897/phytokeys.115.31704
journal article
http://dx.doi.org/10.3897/phytokeys.115.31704
1314-2003-115-51
5349FFA4CF57FFA2FFCDF042FFF5FFFD
2549002
Camelina neglecta J.Brock,
Mandakova
, Lysak & Al-Shehbaz
sp. nov.
Figs 1
, 2
, 3
, 4
Type.
France, Lozere, Causse
Mejean
, corn field, September 1996,
44°16'N
,
2°33'E
,
Henri Besancon s.n.
(holotype: MO-6869197; isotype: MO-6869196).
Description.
Annual herbs. Stems 50-60 cm tall, simple at base, branched about middle or above, densely pilose above base with exclusively simple, crisped trichomes 1-3 mm long, glabrous at middle and above. Basal leaves withered by anthesis; cauline leaves oblong-lanceolate, middle ones 4-5.5
x
0.5-1 cm, gradually reduced in size upwards and becoming narrowly lanceolate, sparsely hirsute with simple trichomes, ciliate with antrorse subsetose trichomes 0.1-1 mm long, base sagittate, margin entire, apex acute. Racemes 30-75-flowered, becoming lax, elongated considerably and 18-24 cm long in fruit; fruiting pedicels 0.9-2 cm long, divaricate-ascending, glabrous. Sepals oblong, 2-2.5 mm long; petals pale yellow, narrowly oblanceolate, 2.5-4.5
x
0.8-1 mm; median filaments ca. 2 mm long; anthers ovate, ca. 0.2 mm long; ovules 30
-34(-
36) per ovary. Fruit pyriform, 7-7.5
x
4- 4.5 mm; valves not veined, margin strongly carinate, winged, apex acuminate, extending 0.9-1.1 mm on to stylar area; style 1.3-1.6 mm long, free portion only ca. 0.5 mm long. Seeds brown, oblong, 0.9-1.1
x
0.5-0.6 mm; seed coat minutely papillate, copiously mucilaginous when wetted.
Figure 1.
Holotype of
Camelina neglecta
.
Besancon s.n.
(MO-6869197).
The
origin of the type material is a seed collection deposited at the USDA and no original voucher is known anywhere, including BORD, long suspected to house it. As a result, a greenhouse-grown plant from the USDA seeds was pressed as the voucher and therefore is recognised as the holotype.
Camelina neglecta
is a diploid species most closely resembling the hexaploid (2
n
= 40)
C. microcarpa
DC. and the tetraploid (2
n
= 26)
C. rumelica
. Deviant counts for
C. microcarpa
are almost certainly based on misidentifications of plants of other species. For example, counts of 2
n
= 26 for
C. microcarpa
from France, Morocco and Spain (see
Warwick and Al-Shehbaz 2006
, BrassiBase) most likely belong to
C. rumelica
, a species two of the authors (MAL and TM) found to consistently have 2
n
= 26. Furthermore, diploid (2
n
= 12) counts for
C. rumelica
, from Hungary (
Baksay 1957
) and United States (
Brooks 1985
), are most likely based on plants of
C. neglecta
or another diploid species yet to be described. Critical verifications of the vouchers of these previous counts are needed to establish their identity beyond any doubt. One of the authors (IAS) examined the voucher cited in Brooks (erroneously reported as McGregor 35289 instead of 35290; Freeman, pers. com.) and it fits quite well in
C. neglecta
, based on trichome morphology and ovule number. Our count of 2
n
= 12 in
C. neglecta
(Fig.
2
) agrees with this and is based on the same seed accession as that of
Martin et al. (2017)
, misidentified as
C. microcarpa
. The present isolated occurrence of
C. neglecta
in France might appear to be odd, but with the availability of resources, a thorough search for it in eastern Europe and Southwest Asia should be made.
Figure 2.
Mitotic chromosomes of
Camelina neglecta
. Greenhouse-grown plants from seeds of
Besancon s.n.
(USDA accession 650135). Scale bar: 10
μm
.
In addition to differences in ploidy level and chromosome numbers,
Camelina neglecta
differs from both
C. microcarpa
and
C. rumelica
by having lower stems soft pilose
(
vs. hirsute) with crisped (vs. straight) trichomes not mixed (vs. mixed) with forked ones (Fig.
3
), as well as by having 30
-34(-
36) [vs. (16
-)20-24(-
26)] ovules per ovary. It further differs from the yellow-flowered
C. microcarpa
by having petals 2.5-4.5 (vs. 3.8-6) mm long petals and fruit 7- 7.5 (vs. 4-5.5) mm long. From
C. rumelica
,
C. neglecta
also differs by the smaller yellow (vs. white) petals 2.5-4.5 (vs. (5
-)6-
9) mm long and pilose (vs. hirsute) lower stems.
Figure 3.
Trichomes of lowermost part of stem in
Camelina neglecta
. Greenhouse-grown plants from seeds of
Besancon s.n.
(USDA accession 650135). Scale bar: 400
μm
.
There are two other Southwest Asian diploid species in the genus, of which
Camelina laxa
C.A.Mey. (2
n
= 12) is distributed in Armenia, Azerbaijan, Georgia, Iran and Turkey and it is unique in the genus in having strongly flexuous infructescences. The other is
C. hispida
Boiss. (2
n
= 14), a species of Iran, Iraq, Israel, Jordan, Lebanon, Syria and Turkey. The latter differs from all other species of the genus by having pubescent (vs. glabrous) middle stems and inflorescences.
The papillate seeds of
Camelina neglecta
(Fig.
4
) are copiously mucilaginous and the seed epidermis exudes the mucilage within a few seconds after soaking.
Figure 4.
SEM image of
Camelina neglecta
seed. Greenhouse-grown plants from seeds of
Besancon s.n.
(USDA accession 650135). Scale bar: 1 mm.
The native ranges of five
Camelina
species (
C. hispida
, C.
laxa
,
C. microcarpa
,
C. rumelica
and
C. sativa
) are widespread in south-eastern Europe and/or Southwest Asia (especially Turkey). Other species,
C. anomala
Boiss. & Hausskn. and
C. stiefelhagenii
Bornm., are rare in Turkey but appeared in areas outside of their known native range, with a collection of
C. anomala
from Beqaa, Lebanon (1961) and
C. stiefelhagenii
from
Dresden
, Germany (1941) and Gothenburg, Sweden (1952). It is quite possible that
C. neglecta
is more widespread in Europe and SW Asia that we currently know.
Due to the allohexaploid nature of
Camelina sativa
, there is much interest in discovering its putative diploid parents. The phylogenetic treatment of the genus (
Brock et al. 2018
) showed the relationships of diploid
Camelina
species relative to
C. sativa
and indicated a potentially shared hybridisation and polyploidisation history of the weedy
C. microcarpa
and its domesticated
C. sativa
. It is essential to identify the wild
Camelina
diploids to facilitate reconstruction of the evolutionary history of
C. sativa
and allow the potential for re-synthesis of the crop as has been done in
Brassica napus
L. (
Chen et al. 1988
).