Chromosome Counts of Some Veronica L. (Scrophulariaceae) Species from Iran

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Sect. Beccabunga subsect. anagalloides Keller includes
V. anagallis-aquatica L. and V. anagalloides Guss., which
presents a taxonomical problem. Sect. Alsinebe (Griseb.)
Lehm. is also an extremely confusing group and includes
32 species and several subspecies. Hybridisation occurs
frequently in this section with 3 ploidy levels, i.e. diploids,
tetraploids and hexaploids are known (Öztürk & Fischer,
1982). Some taxa examined have nearly worldwide
distribution, while V. francispetae M.A.Fischer, V.
siaretensis E.Lehm. and V. ceratocarpa C.A.Mey. are
endemic to the Elburz mountain range in Iran (Fischer,
There are several reports on the cytotaxonomy of
various species of Veronica (Fischer, 1967, 1973;
Podlech & Dieteric, 1969; Ferakova, 1976; FernandesCasas, 1977; Aryavand, 1987; Ghaffari 1987). The
present study and the previous reports of chromosome
numbers in Veronica confirm the variation in the basic
Materials and Methods
The materials used for this study were collected from
wild populations. Flower buds were fixed in Carnoy’s
mixture in the field; the ovaries were then excised and
stained in 2% aceto-orcein solution for 5 min. They were
immersed in 0.075 M KCl at room temperature for 20
min; then the ovaries were treated with enzyme solution
containing 5% cellulase and 5% pectinase and adjusted to
pH 4.0 for about 20 min at 35 °C in order to dissolve the
cell wall (Kurata & Omura, 1978). The ovaries were
squashed in 45% acetic acid and then permanently
mounted in Entellan to facilitate investigations of somatic
mitosis. Permanent preparations were made and studied
using a BH-2 Zeiss photomicroscope. Voucher specimens
are deposited in the Herbarium of Guilan University.
The chromosome numbers from 13 populations of
the 10 species examined are given in Table 1.
V. anagalloides Guss subsp. heureka M.A.Fischer
(Figure A)
This taxon occupies an area from Central Anatolia to
Pakistan. It is distinguishable from V. anagalloides subsp.
anagalloides by its wider and shorter leaves and its
suborbicular (not elliptical) and larger capsule. Our counts
of 2n = 36 agree with that given by Podlech & Dieteric
Turk J Bot
29 (2005) 263-267
Chromosome Counts of Some Veronica L. (Scrophulariaceae)
Species from Iran
Department of Biology, Faculty of Science, University of Guilan, Rasht – IRAN
Department of Agronomy and Plant Breeding, Faculty of Agricultural Science, Islamic Azad University of Rasht, Rasht – IRAN
Received: 02.09.2004
Accepted: 07.03.2005
Abstract: The chromosome numbers of 10 Veronica L. species belonging to sect. Alsinebe (Griseb.) Lehm. and sect. Beccabunga
Griseb. from northern Iran are given. The chromosome numbers for 3 of the 10 studied taxa, i.e. V. siaretensis E. Lehm., V.
ceratocarpa C.A.Mey. (2n = 14) and V. hederifolia L. (2n = 36) are presented for the first time. Our results are compared with
previous records.
Key Words: Veronica, Chromosome number, Iran, mitosis.
Research Article
(1969), who studied material from Afghanistan. Öztürk
& Fischer (1982) have reported 2n = 18 for this
subspecies based on only one collection of Turkish
material of V. anagallis-aquatica subsp. lysimachioides.
V. anagallis-aquatica L. subsp. michauxii (Lam.)
A.Jelen. (Figure B)
Some taxa of the V. anagallis-aquatica group have an
almost worldwide distribution, but its centre of diversity
is south-west Asia (Turkey to Pakistan), where there are
at least 9 subspecies or species. The tetraploid
chromosome number of 2n = 36 studied here confirms
that reported by previous authors (Öztürk & Fischer,
1982) as well as by Aryavand (1987), who examined
material from central Iran.
According to Öztürk & Fischer (1982), this species is
possibly a hybrid between subsp. michuaxii and subsp.
anagallis-aquatica, although capsules with fertile seeds
V. siaretensis E.Lehm. (Figure C)
This species is endemic to Iran, where we found it
only in one locality, Gorgan. The chromosome number of
2n = 14 is the first record for this species.
V. ceratocarpa C.A.Mey. (Figure D)
This taxon is distributed in Turkey, Iran and middle
Asia with a wide distribution in the deciduous forests of
Elburz mountain in northern Iran. There are no previous
counts for this taxon, and so the chromosome number of
2n = 14 is reported here for the first time.
Chromosome Counts of Some Veronica L. (Scrophulariaceae) Species from Iran
Table 1. Chromosome numbers of 10 Veronica species from Iran.
Taxon Pop. No. Locality 2n
V. anagalloides Guss 1 Guilan; Asalam, 50 km to Khalkhal, 1700 m, 36
subsp. heureka M.A.Fischer Saeidi 24077
V. anagallis-aquatica L. 2 Mazandaran; SW Ramsar, Javaher-Deh, 1800 m, 36
subsp michauxii (Lam.) A.Jelen. Saeidi 1505
V. siaretensis E.Lehm. 3 Gorgan; Ziarat village, 1000-1100 m, Saeidi & 14
Kaviani 1248
V. francispetae M.A.Fischer 4 Guilan; near Langroud, Talesh-Mahleh, 30 m, 14
Saeidi 24025
V. capillipes Neveski 5 Tehran; from Karaj to Chalus, near Gachsar, 28
1400 m, Saeidi 1301
V. campylopoda Boiss. 6 Gorgan; Bastam, Tash-Olia, Shavar, Saeidi 42
& Kaviani 1311
V. polita Fr. 7 Guilan; 7 km from Asalam to Talesh, 500-600 m 14
Saeidi 1315
8 Mazandaran; near Abas-Abad, 1080 m, 14
Saeidi & Asaadi 1452
V. persica Poir. 9 Guilan; Daylaman, Shah-Shahidan, 1500-2000 m, 28
Jamzad & Asri 71766
10 Mazandaran; Nowshahr, Kheyrud-Kenar forest, 28
200 m, Saeidi 1307
V. hederifolia L. 11 Tehran; from Karaj to Chalus, near Gachsar, 36
1400 m, Saeidi 1310
V. ceratocarpa C.A.Mey. 12 Mazandaran; Klardasht, 1300-1400 m, 14
Saeidi 24021
13 Guilan; Masuleh, 700-800 m, Saeidi 1197 14
V. francispetae M.A.Fischer (Figure E)
This species is endemic to northern Iran and grows
beneath trees and shrubs. The diploid chromosome
number of 2n = 14 confirms the count reported by
Fischer (1981).
V. capillipes Nevski (Figure F)
This species has a disjunct distribution in north-east
and central Iran, Pakistan and middle Asia. Meiosis
studies of this species by Aryavand (1987) resulted in a
count of n = 14, which he reported in the abstract,
although in the main text this is incorrectly given as 2n =
14. Our count of 2n = 28 for this species agrees with the
n = 14 reported by Aryavand (1978), who studied plant
material from Isfahan province in Iran.
V. campylopoda Boiss. (Figure G)
This species is an Irano-Turanian element with a wide
distribution in Iran, Turkey and middle Asia. A meiotic
count of n = 21 was reported for this species by
Aryavand (1987) and Ghaffari (1987). We found 2n = 42
for this species, which agrees with the previously
published counts.
Figures. A-K: Somatic cells of Veronica species. A. V. anagalloides subsp. heureka B. V. anagallis-aquatica L. subsp. michauxii C. V.
siaretensis D. V. ceratocarpa E. V. francispetae F. V. capillipes G. V. campylopoda H. V. polita J. V. persica K. V. hederifolia.
(Scale bar = 10 µm).
V. polita Fr. (Figure H)
This species is widespread in Asia and Europe and
extends into Africa and is also distributed in the USA
(Fischer, 1981). The diploid chromosome number of 2n
= 14 coincides with the results of previous authors
(Fernandes et al., 1977; Aryavand, 1987; Ghaffari,
V. persica Poir. (Figure J)
This species is widespread in Iran and throughout the
world. Our count of 2n = 28 agrees with those reported
by Ferakova (1976) and Fernandes et al. (1977).
V. hederifolia L. (Figure K)
This species has a relatively wide distribution in Iran,
south-west Asia, middle Asia, Turkey and Europe
(Fischer, 1981) and can be found at elevations of up to
1000 m. The count of 2n = 36 is the first record for this
species from Iran, although an earlier chromosome count
of 2n = 54 was reported from central Iran by Aryavand
(1987), which agrees with previous counts by Fernandes
et al. (1977) and Fischer (1981). Based on Danish and
Dutch material, Gadella & Kliphuis (1975, 1976)
reported 2n = 36 for this species.
The reports so far suggest that each taxon
representing a biological entity (subspecies or species) is
characterised by one ploidy level, either diploid, tetraploid
or hexaploid.
A meiotic count of n = 14 was reported for V.
capillipes by Aryavand (1987). Ghaffari, in unpublished
data (personal communication), repeated a mistake in the
text of the paper, which gives 2n = 14, and which was
published in Persian by Aryavand (1987). Aryavand
(1987) has indicated that the species is diploid, but our
studies confirm that V. capillipes is a tetraploid species.
Two different cytotypes were known for V.
hederifolia, with 2n = 54 the more common. This
cytotype is also recorded for V. hederifolia in Iran by
Aryavand (1987). The other cytotype (2n = 36),
corresponding to subsp. lucorum (Klett & Richter) Hartl.
was not previously known in Iran, but according to the
present study the cytotype of 2n = 36 suggests that this
subspecies may be present in Iran.
Fischer (1967) concluded that the 2 cytotypes are so
different that they can be regarded as separate species,
Veronica sublobata Fisch. (tetraploid) and Veronica
hederifolia L. (hexaploid), rather than as just a subspecies.
Neither Nordenstam & Nilsson (1969), nor De Jongh &
Kern (1971) accepted this conclusion. They agreed with
Fischer that the 2 taxa could be recognised, but they
disagreed with him about the species level for these taxa.
The Veronica hederifolia complex represented by material
from the Netherlands by 2 different habitat cytotypes
differ morphologically and probably do not exchange
genes (Gadella & Kliphuis, 1976). They suggested that
the morphological characters can to a certain extent vary,
but in all cases tetraploid and hexaploid plants can be
identified by means of a combination of at least 3 or 4
such characters. Gadella & Kliphuis (1975, 1976)
support Fischer’s view that these taxa should be regarded
as separate biological species. They agree that the 2
groups are distinguishable based on the colour of the
anthers and the corolla, and the form and structure of the
seeds. However, we did not find any comparable
morphological differences between them in Iran except
for the habit of the 2 cytotypes. We found that the
tetraploid plants were usually more slender and smaller
than the hexaploid plants.
Hybrids between tetraploid and hexaploid plants of
Veronica hederifolia are unknown and, considering the
subspecific rank to which the variants belong, it seems
that this subject needs to be studied in more detail before
the correct taxa can be assigned. If the view is held that
the (apparently rare) intermediate transitional forms are
so important that they blur the distinction between the 2
forms, then the tetraploid should be assigned to Veronica
hederifolia L. subsp. lucorum (Klett & Richter) Hartl. and
the hexaploid to Veronica hederifolia L. subsp. hederifolia.
Since these transitional specimens are perfectly fertile
and may occur in large proportions in Asian and European
populations, this is the main reason why we have decided
to assign the subspecific level to these 2 taxa.
These deviating chromosome numbers demonstrate
the need for further studies on the Veronica hederifolia
We are grateful to Dr. Sh. Zarre, Tehran University,
for his useful comments. The first author was supported
by project number 65 of the University of Guilan
Research Council.
Chromosome Counts of Some Veronica L. (Scrophulariaceae) Species from Iran
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