Systematic treatment of Veronica L. Section Beccabunga (Hill) Dumort (Plantaginaceae)

Abstract

Veronica species mostly occur in damp fresh water places and in the Mediterranean precipitation regime. Members of this genus grow at different altitudes from sea level to high alpine elevations. They show a high level of polymorphism and phenotypic plasticity in their responses to variations of the enviromental factors, a quality that allows them to occur over a wide range of conditions. A group with particular high levels of polymorphism is the group of aquatic Veronica L. species in V. sect. Beccabunga (Hill) Dumort. Here, we attempt to unravel some confusion in the taxonomic complexity in V. section Beccabunga. We recognize 20 taxa in V. sect. Beccabunga and explore the occurrence of V. section Beccabunga, mainly in the Mediterranean basin; especially in Egypt (Nile delta and Sinai), Turkey and Iran with each country containing 10 taxa, from a total of 20 taxa, and characterized by endemics, or near-endemic as Veronica anagalloides ssp. taeckholmiorum.The results confirmed that V. section Beccabunga is divided into three subsections Beccabunga, Anagallides and Peregrinae, which essentially can be differentiated by the absence or presence of apetiole.

Full text

eISSN: 2357-044X Taeckholmia 38 (2018): 168-183
Received on 05/10/2018 and accepted on 17/12/2018
Systematic treatment of Veronica L. Section Beccabunga (Hill) Dumort
(Plantaginaceae)
Faten Y. Ellmouni1, Mohamed A. Karam1, Refaat M. Ali1, Dirk C. Albach2
1Department of Botany, Faculty of Science, Fayoum University, 63514 Fayoum, Egypt.
2Institute of biology and environmental sciences, Carl von Ossietzky-University, 26111 Oldenburg,
Germany.
*Corresponding author: fyl00@fayoum.edu.eg
Abstract
Veronica species mostly occur in damp fresh water places and in the Mediterranean precipitation regime.
Members of this genus grow at different altitudes from sea level to high alpine elevations. They show a high level
of polymorphism and phenotypic plasticity in their responses to variations of the enviromental factors, a quality
that allows them to occur over a wide range of conditions. A group with particular high levels of polymorphism is
the group of aquatic Veronica L. species in V. sect. Beccabunga (Hill) Dumort. Here, we attempt to unravel some
confusion in the taxonomic complexity in V. section Beccabunga. We recognize 20 taxa in V. sect. Beccabunga and
explore the occurrence of V. section Beccabunga, mainly in the Mediterranean basin; especially in Egypt (Nile
delta and Sinai), Turkey and Iran with each country containing 10 taxa, from a total of 20 taxa, and characterized
by endemics, or near-endemic as Veronica anagalloides ssp. taeckholmiorum.The results confirmed that V. section
Beccabunga is divided into three subsections Beccabunga, Anagallides and Peregrinae, which essentially can be
differentiated by the absence or presence of apetiole.
Keywords: Morphological key, systematic treatment, Veronica, V. section Beccabunga
Introduction
The tribe Veroniceae, formerly part of
Scrophulariaceae, is now found within an enlarged
family Plantaginaceae (Olmstead and Reeves, 1995;
APG 1998, 2003). Veroniceae is distinguishable from
related taxa by its tetramerous corollaand capsule
dehiscence (normally loculicidal, only exceptionally
septicidal (Hong, 1984). Based on the most recent
evidence from phylogenetic analyses of Veronica and its
related genera , nine monophyletic genera are
recognizedin the tribe Veroniceae (Veronica, Scrofella,
Veronicastrum, Lagotis, Wulfenia, Kashmiria,
Picrorhiza, Wulfeniopsis and Paederota; Albach et al.,
2004a).
Veronica is the most species-rich genus of family
Plantaginaceae. It comprises about 500 species (Albach
et al., 2004a). It is distributed over most of the Northern
Hemisphere and in many parts of the Southern
Hemisphere, and is ecologically highly diverse with
species growing in different habitats and elevations,
from aquatic to even dry steppe from sea level to high
alpine regions.
Veronica in this wider sense is characterized by
several synapomorphies (mostly with several reversals
within the genus), such as four calyx and four corolla
lobes, short to absent corolla tube, subactinomorphic
flowers, two exserted stamens, notched fruit apex, much
compressed fruit, and flattened seeds (Albach et al.,
2004b). The main progressions (usually parallel) within
the genus include: life-form (subshrubby/perennial vs.
annual), position of inflorescences (axillary vs.
terminal), leaf lamina (large vs. small and linear,
undivided vs. pinnatifid), size of corolla (usually related
to breeding system), shape of seeds (flat vs. excavate
=“cymbiform”), indumentum (glandular vs. eglandular
hairs) according to Albach et al.
(2004 a, c). Albach et
al. (2008) found that most of the subgenera in Veronica
exhibit only one single basic number, i.e., x = 6, 7, 8, 9,
12, 17, or 20/21. They pointed to the putative ancestral
base number in this genus of 9, which has been reduced
several times to 8 and 7, respectively
(aneuploidy/dysploidy), often associated with transition
to annual life history.
Based on the work carried out by Albach et al.
(2004 a) and Garnock-Jones et al. (2007), genus
Veronica is, now grouped into 12 subgenera, each one
within in two to 150 species: V. subg. Beccabunga,
Veronica, Pseudolysimachium, Synthyris,
Cochlidiosperma, Pellidosperma, Stenocarpon,
Triangulicapsula, Pocilla, Pentasepalae, Chamaedrys,
and Pseudoveronica.
According to (Albach et al., 2004a) , Veronica
subg. Beccabunga includes V. sect. Beccabunga sensu
Stroh (1942) (excluding V. ciliata and V.himalensis), V.
subsect. Serpyllifolia sensu Stroh (1942) (perennials,
incl. V. serpyllifolia L. and relatives; excluding V.
aragonensis and V. chionantha) and V. subsect.

Systematic treatment of Veronica L. Section Beccabunga
Acinifolia sensu Stroh (1942) (annuals, excluding V.
macropoda and V. rubrifolia, the complete subsection
sensu Elenevsky), as well as, V. gentianoides with its
allies and the annual V. peregrina of V. subsect.
Microspermae sensu Stroh (1942).
Veronica section Beccabunga is composed of
semi-aquatic species, hygro- and hydrophytes found in
different natural zones and at very different elevations
from sea-level to alpine zones (Savinykh, 2003). Section
Beccabunga includes herbaceous perennials and
annuals, which occur primarily in the temperate regions
of all continents of the Northern and Southern
Hemispheres, except Antarctica. However, the greatest
diversity of Veronica section Beccabunga exists in
southwest Asia (Chrtek and Osbornová-Kosinová,
1981). Despite the fact that some species can live in arid
habitats, members of Veronica sect. Beccabunga are not
xerophytes.They are flowering and bearing fruit in early
spring and are rather mesophytes and ephemeral plants
(Savinykh, 2003).
Veronica section Beccabunga has been discussed
in various floras and monographic studies, such as
Krösche (1912), Römpp (1928), Schlenker (1936 a,b),
Keller (1942), Stroh (1942), Burnett (1950), Lawalrée
(1953), Khoshoo and Khushu (1966), Marchant
(1970,1967), Hartl (1968), Walters and Webb (1972),
Smejkal and Helanová-Zichová (1974), Jelenevskij
(1978), Fischer (1978), Chrtek and Osbornová-
Kosinová (1981), Öztürk and Fischer (1982), Sellers
(1983), Fischer et al. (1984), Les and Stuckey (1985),
Fischer (1989), Klinkova (1993), Tanaka (1996),
Tanaka et al. (1999), Dzhus and Dmitrieva (1999),
Öztürk and Öztürk (2000), Sánchez-Agudo (2004),
Sánchez-Agudo et al. (2011), Abd El-Ghani et al. (2010,
2011) and Ellmouni et al. (2017)
Keller (1942) divided section Beccabunga into
two separate subsections, i.e. Anagallides and
Beccabunga based on absence or presence of a petiole.
Subsection Anagallides includes species as V.
anagalloides, V. anagallis-aquatica, V. catenata, V.
scardica and V. kaiseri, which often form intermediates;
it is an extremely confusing complex of several species
and subspecies intimately connected. Veronica
beccabunga and V. americana form subsection
Beccabunga, which distinct from the above mentioned
species and form no intermediates with them.
Chrtek and Osbornová-Kosinová (1981), Öztürk and
Fischer (1982), Saeidi Mehrvarz and Kharabian (2005),
and Abd El-Ghani et al. (2010, 2011) followed Keller
(1942) in his division.
Borissova (1955) further splitted section
Beccabunga into three series, i.e. Anagallides,
Beccabungae and Oxycarpae based on the petiole and
the shape of capsule. Her series Anagallides includes
species with mostly sessile leaves (V. anagallis-
aquatica, V. anagalloides, V. anagallidiformis, V.
scardica, V. poljensis), the second series Beccabungae
includes petiolate species (V. beccabunga and V.
americana) ,while the third series Oxycarpae include
sessile or petiolate species (V. beccabungoides, V.
montioides, V. bobrovii,. V. michauxii,. V.
lysimachioides and V. oxycarpa).
Although members of V. sect. Beccabunga were
included in molecular phylogenetic analyses since the
beginning of phylogenetic analyses of the genus (e.g.,
Wagstaff and Garnock-Jones 1998), a recent study with
fairly complete taxon sampling allowed inferences
about the relationships (Ellmouni et al. 2017). However,
the complexity of molecular phylogenetic and
morphometric analyses did not allow a detailed
discussion of systematic implications.
The aim of the present study is to summarize
current knowledge of species relationships in Veronica
section Beccabunga, provide a taxonomic basis for
discussion and highlight areas of particular need for
further research. For that purpose, a systematic
treatment was conducted to include:
(1) a summary of the species recognized and reported in
the earlier workers in different floras.
(2) a key to all species of section Beccabunga.
(3) a discussion of its relationships between conspecific
species.
Material and methods
Plant material
The present study is based on specimens, both
own collections and herbarium specimens from Africa
(Egypt, Ethiopia); Europe (Turkey, Germany, Czech,
Austria); Asia (Armenia, Azerbaijan, Georgia, Pakistan,
Kyrgyzstan, Afghanistan) and North America (USA and
Canada). A total of 100 specimens representing 20 taxa
of section Beccabunga were collected and examined
with a MS003A binocular head zoom stereo
microscope. (Appendix 1).
Results and Discussion
We here compiled a summary of all species of
Veronica sect. Beccabunga recognized and reported in
the earlier studies starting with Pennell (1935) and
ending by our treatment (Table 1). In the following, we
present an artificial key for all represented taxa.
169

Faten Y. Ellmouni et al
- Artificial key to species of Veronica sect. Beccabunga based on morphological characters
1.a) Annuals, without subterranean stem; most leaf-axils flower-bearing; inflorescence "terminal or axillary"; leaves,
oblanceolate, or narrowly oblong …………………………………….….…V. peregrina (subsect. Peregrinae)
1.b) Perennials or annuals, with subterranean stems; only the upper leaf-axils flower-bearing; inflorescence terminal;
leaves, lanceolate, ovate or orbicular ………………………………………………………………………….. 2
2.a) Flowering stems procumbent or ascending; upper leaves petiolate, glabrous.
……………..…………………………………………………………….………….... 3 ( subsect. Beccabunga)
2.b) Flowering stems erect; upper leaves sessile, glabrous or hairy……………..……………. 6 (subsect. Anagallides)
3.a) Leaves lanceolate or ovate; leaf blade twice as long as wide, broadest part at base; style 2.5-3.5 mm
..…………………………………………………………………………………………..…………V. americana
3.b) Leaves obovate or broad elliptic; leaf blade as long as to twice as long as wide, broadest part near middle or above;
style 1.8-2.2 mm ………..……………………………………………………………….……..…V. beccabunga
4.a) Corolla 6-9 mm in diam.; pedicel not patent, peduncle up to 50 mm long; longer than the subtending leaf; capsule
4-6 mm long, 3-4.5mm wide………..………… ……………….………………. V. beccabunga ssp. abscondita
4.b) Corolla 4-7 mm in diam.; pedicel patent, peduncle up to 30 mm long; not longer than the subtending leaf; capsule
2.5-5.5 mm long, 3-5 mm wide ……..…………………….……….………………………………..………….. 5
5.a) Leaves 10-65 mm X 6-33 mm with clear petiole; peduncle up to 30 mm; pedicels subpatent, up to 10 mm; number
of flowers up to 22. .…………………………………………………………….V. beccabunga ssp. beccabunga
5.b) Leaves 4-25 X 3-15 mm without clear petiole; peduncle up to 20 mm; pedicel horizontally patent, up to 8 mm;
number of flowers up to 15. ….………………………………………………….….V. beccabunga ssp. muscosa
6.a) Raceme generally glandular or villous ……………………………………………………….…………...………… 7
6.b) Raceme rarely glandular, generally glabrous . …………………………………………………………….………..10
7.a) Inflorescence axis more or less villous; capsule as long as calyx or longer ……………………….……….V. poljensis
7.b) Inflorescence axis glandular-pubescent; capsule as long as calyx or shorter ………………………………………. 8
8.a) Leaves petiolate or sessile ,ovate.………………………………………………V. anagallis-aquatica ssp. michauxii
8.b) Leaves sessile, orbicular to linear. ……………………………………………………………….……………………9
9.a) Capsule elliptic ……………………………………..……………………………. ..V. anagalloides ssp. anagalloides
9.b) Capsule orbicular..………………………………………………………………………V. anagalloides ssp. heureka
10.a) Racemes alternate; lower leaves petiolate , upper subsessile or sessile..………………………………..…………11
10.b) Racemes opposite; lower leaves shortly petiolate , upper sessile …………………………………..……..………13
11.a) Leaves suborbicular to broadly elliptic; capsule with acute apex , and as long as the calyx lobes
………………………..………………..…………..…………………………………….…..………….V. kaiseri
11.b) Leaves suborbicular to rhombic; capsule with rounded apex, and slightly shorter than the calyx lobes.
..……………………..……………………………… …………………….………………………………...….12
12.a) Stems erect; length of capsule ≤ 2 mm, width of capsule < 2 mm; style ≥ 1.5 mm long ……
…………………………………………………………………………………....…… V. scardica ssp. scardica
12.b) Stems creeping or erect; length of capsule > 2 mm, width of capsule ≥ 2 mm; style < 1.5 mm long
………………………………………………………………………………….………V. scardica ssp. africana
13.a) Upper leaves ovate-lanceolate to lanceolate; capsule orbicular ……………………………….…………..……… 14
13.b) Upper leaves lanceolate to linear lanceolate or rhombic-lanceolate; capsule elliptical.……………………………21
14.a) Flower bract longer than pedicels in 1st and 2nd flower in inflorescence; bract leafy at 1st and 2nd flower and rest
lanceolate ……………………………………………………….…………...… V. catenata ssp. pseudocatenata
14.b) Flower bract shorter than pedicels, bracts lanceolate allover the inflorescence …………….…………………….15
15.a) Lower leaves shortly petiolate, leaves broader and longer, pedicel erect ……………………….…………………16
15.b) Lower leaves sessile, leaves generally broader and shorter, pedicel patent or suberect ……………………….… 19
170

Systematic treatment of Veronica L. Section Beccabunga
16.a) Capsule ovoid-deltoid, basally wider than apically ovate, apically tapering ..……………..………………………17
16.b) Capsule subglobose, basally and apically equal in width (cylindrically), apically rounded or emarginated..…… 18
17.a) Racemes up to 100 mm long, with ± lax flowers (3-4 flowers / cm); peduncle length up to 20 mm; style up to 1.5
mm long; capsule orbicular or broadly elliptic, with acute apex
……………………….………………………...………………………….… V. anagallis-aquatica var. nilotica
17.b) Racemes 150 mm long, with ± dense flowers (3-7 flowers /cm); peduncle length up to -40 mm; style to 4 mm long;
capsule ovoid-deltoid, slightly compressed, with tapering to slightly notched apex
………………………………………………………………………..…… V. anagallis-aquatica ssp. oxycarpa
18.a) Pedicel curved, at an acute angle with inflorescence axis; racemes less than 1 cm wide, usually glabrous
………...................................................................................................................................V. anagallis-aquatica
18.b) Pedicel straight, at a right angle with inflorescence axis; racemes 1–1.5 cm wide, sparsely - pubescent
.............................................................................................................................................................. V. undulata
19.a) Raceme up to 150 flowered, glabrous; capsule as long as the calyx or shorter; pedicel suberect
.............................................................................................................. V. anagallis-aquatica ssp. lysimachioides
19.b) Raceme up to 40 flowered, sparsely pubescent or glabrous; capsule as long as the calyx or longer; pedicel patent
……………………………………………………………………20
20.a) Leaves oblong-ovate to oblong-lanceolate, with entire or subentire margin; raceme with 15 - 25 flowers; style >2
mm …………………………………………………………………………………..…V. catenata ssp. catenata
20.b) Leaves ovate to ovate-lanceolate,with serrulate - subentire margin; raceme with 10-40 flowers; style < 2 mm
……..……………...…………………………………………...………….V. anagalloides ssp. heureka (Egypt)
21.a) Leaves lanceolate to linear lanceolate; bract longer than the pedicel; style 1-1.3 mm; capsule as long as calyx or
slightly longer ………………….V. anagalloides ssp. taeckholmiorum
21.b) Leaves lanceolate to rhombic-lanceolate; bract shorter than the pedicel or subequal; style 0.6-1.5 mm; capsule as
long as or twice calyx ……………………………………………..…V. anagalloides ssp. anagalloides (Egypt)
Aquatic angiosperms represent a polyphyletic
assemblage of diverse evolutionary lineages unified as a
'biological group' by their common invasion of the
aquatic environment. Water plants have long been a
fascination to naturalists, horticulturists, and aquarists;
issues of their complex transition from terrestrial to
aquatic life, however, have scarcely been addressed by
evolutionary biologists (Les and Philbric, 1993).
The taxonomic history of Veronica clearly
illustrates the difficulties arising from the recognition of
natural (monophyletic) groups within the genus based
solely on morphological traits. This is due to the fact that
many of the traditionally used taxonomic characters are
evolutionary labile and therefore not suitable for
taxonomy (Albach et al., 2004c). The taxonomic
complexity of the genus is due to several polymorphic
subsections and groups with closely related taxa (Saeidi
and Zarre, 2004).
Veronica subg. Beccabunga seems to be an early
branching clade in the genus (Albach and Chase 2001;
Albach et al. 2004b, c). Within the subgenus, section
Beccabunga appears to be sister to the rest of the
subgenus (Müller and Albach 2010).
According to Keller (1942), Chrtek and
Osbornová-Kosinová (1981), Öztürk and Fischer
(1982), Saeidi and Kharabian (2005), Abd El-Ghani et
al. (2011), Veronica section Beccabunga comprises two
subsections; Beccabunga, and Anagallides, based on
presence or absence of the petiole. In our earlier study
(Ellmouni et al. 2017), we confirmed the addition of
subsection Peregrinae to section Beccabunga as a third
subsection, congruent with Albach and Chase (2001),
Albach and Greilhuber (2004), Taskova et al.(2004),
Albach et al. (2004b), Muñoz-Centeno et al. (2006),
Albach et al. (2008), Müller and Albach (2010), and
Hassan and Abd El-Khalik (2014). Those studies have
placed V. peregrina in section Beccabunga based on
DNA data and seed morphology. The annual Veronica
peregrina L., is the only member of V. subsection
Peregrinae Elenevsky.
The main morphological character shared
between Veronica peregrina and other taxa in section
Beccabunga is the shape of leaves, which is
oblanceolate in the lower leaves and narrowly oblong in
the upper leaves according to Hong and Fisher (1998),
whereas the main difference between subsection
Peregrinae and other members of V. section
Beccabunga is appearing in the inflorescence, which in
the first is terminal, with rarely some axillary flowers
but in the others the inflorescence is axillary. An
additional shared character is the preference for at least
seasonally wet habitats. Veronica peregrina is
hexaploid based on the shared base chromosome
number x = 9 (Albach et al. 2008).
Subsection Beccabunga contains two species,
the Eurasian mainly diploid V. beccabunga and the
Americo-Beringian tetraploid V. americana (Öztürk and
Fischer, 1982). Sellers (1983) considered V.
beccabunga and V. americana as conspecific, and stated
“Although the two taxa are strikingly similar, V.
beccabunga and V. americana are better treated as
171

Faten Y. Ellmouni et al
subspecies of a single species”, due, in part, to their
geographical distribuation, and slightly difference in
both of leaf morphology, and chromosome number
(Marchant,1968; Sellers, 1983). We consider, however,
these characters to be enough to recognize them as
separate species. Veronica americana occurs primarily
in North America, throughout the Aleutian Islands, and
in East Asia along the Pacific coast and neighboring
islands; while V. beccabunga ranges mainly throughout
much of the Old World. However, isolated occurrences
of V. beccabunga in North America, are presumably
introducted. Veronica beccabunga has round or obovate
leaves with a round apices; V. americana has lanceolate
or oblong leaves with obtuse apices, whereas V.
beccabunga is diploid (x=9) while, V. americana is a
tetraploid species (Marchant, 1968; Sellers,1983;
Albach et al., 2008).
V. beccabunga and V. americana are not only
morphologically similar but, also, form bastard plants
that are sterile (Schlenker 1935) despite the fact that they
have different ploidy level. Pennell (1921) mentioned
that "V. americana appears to be only constantly
distinguishable from V. beccabunga by its leaf-form and
more erect habit".
Les and Stuckey (1985) discussed the
introduction of Veronica beccabunga into eastern North
America in the late nineteenth century as a result of
ballast disposal. The species spread westwards through
the dispersal of seeds and plant fragments. They also
mentioned that the slower migration of V. beccabunga
may be due to its affinity for undisturbed habitats where
native species such as V. americana area are already
established since V. americana and V. beccabunga are
essentially equivalent ecologically (Marie-Victorian,
1935; Les and Stuckey, 1985).
Veronica subsect. Anagallides is a taxonomically
extremely difficult group in desperate need of a DNA
based analysis to disentangle taxonomically useful
characters from those having evolved in parallel or
phenotypically plastic. Both ITS and plastid rps16-
trnK-sequences did not help much in resolving
relationships (Ellmouni et al. 2017). Species in Europe
have been studied intensively (including crossing
experiments) by Schlenker (1935) and Marchant (1970)
and seem to be distinct species. Hybrids between the
species of the subsection appear to be common (e.g.,
between V. angallis-aquatica and V. catenata),
sometimes resulting in sterile plants (V. ×
lackschewitzii) but probably also leading to fertile
backcross plants. A further difficulty is the wide
distribution and weedy nature with plants often found
along the smallest ephemeral puddles. Due to these
difficulties species delimitation is complicatedand often
based on single characters as capsule shape (e.g., V.
oxycarpa) or indumentum (e.g., V. michauxii). As
mentioned by Fischer (1981), when collecting plants
from the group many individuals in different parts of the
population should be checked for variability of the
characters, and information on sterile and injured plants
should be recorded. Flower color may be an important
character as it is often not discernible in herbarium
specimens. One character, which deserves some
attention, is the position of the inflorescence. For
Egyptian flora, some emphasis is put on this character.
The current work supports that Veronica
anagalloides, is represented by three subspecies ; V
.
anagalloides ssp. anagalloides, V
.
anagalloides ssp.
taeckholmiorum and V
.
anagalloides ssp.
heureka.These possess lanceolate to linear-lanceolate
leaves,and small elliptic capsule. However, these are
differentiated by the length of the pedicel in relation to
the bract. Veronica anagalloides ssp. anagalloides and
ssp. taeckholmiorum share lanceolate to linear
lanceolate upper leaves, and small , elliptic capsule;but
differ in the length of pedicel and bract, density of
flowers , and the size of capsule. In V
.
Anagalloides ssp.
taeckholmiorum,bracts longer than pedicel (up to 1 cm),
inflorescence dense (up to 9 flowers/ cm), and small size
of capsule; ssp. anagalloides has bracts as long as or
shorter, density of inflorescence less than in other
subspecies (up to 7 flowers / cm) and relatively large
size of capsule (Abd El-Ghan et al., 2010). The extreme
polyphyly of V. anagalloides in molecular and
morphometric analyses by Ellmouni et al. (2017) is
noteworthy and suggests that species boundaries are
either very young, hybridization occurs frequently
and/or phenotypic plasticity blurs species boundaries.
Natural hybrids are also reported to involve
Veronica anagallis-aquatica and Veronica anagalloides
(Fischer, 1981). A natural hybrid involving Veronica
poljensis and Veronica anagalloides has been reported
by Öztürk and Fischer (1982), on the basis of
morphological intermediacy.
With regards to Veronica poljensis, we do not
agree neither with Marchant (1970) who considered V.
poljensis as a part of V. anagalloides nor with
Jelenevskij (1978) who included V. poljensis in V.
anagallis-aquatica subsp. anagalloides) but agree with
Schlenker (1936) and Borisova (1955) who treated V.
poljensis as a separate species. Schlenker (1936) had
explained the relation between V. poljensis, V. anagallis
and V. anagalloides, stating that V. poljensis resembles
V . anagalloides in the narrow and linear-lanceolate
leaves and the small flowers; but differs with broader
capsule; while it shares V . anagallis-aquatica in the
suborbicular capsule but differs in its narrow leaves.
However, more detailed studies remain to test the
coherence of plants with villous capsules.
Veronica anagallis-aquatica is a polymorphic
species, a homophyllous amphibious macrophyte
(Sculthorpe, 1967). The polymorphic nature is extended
to the molecular results in Ellmouni et al. (2017), which
make it look probable that the species forms kind of
source from which other taxa originated.This species
has serrulate to subentire, narrowly ovate to lanceolate,
and sessile leaves. In some streams, V. anagallis-
aquatica occurs both as emerged and submerged forms
at low water velocities, where the substrate is a mixture
of sand and silt,while in high water velocity, V.
anagallis-aquatica occurs, only, as the submerged form
growing on gravel substrate (Boeger and Poulson,
2003). The V. anagallis-aquatica group has a nearly
172

Systematic treatment of Veronica L. Section Beccabunga
3
world-wide distribution (Chrtek &Osbornová-
Kosinová, 1981).
Chrtek and Osbornová-Kosinová (1981) and El-
Hadidi et al. (1999) mentioned the absence of V.
anagallis-aquatica var. anagallis-aquatica in Egypt, but
pointed out the presence of V. anagallis-aquatica var.
nilotica in Egypt, particularly Aswan area. In the
meantime, Abd El-Ghani et al. (2010), partially agreed
with previous authors,who confirmed the occurrence of
both taxa, V. anagallis-aquatica subsp. anagallis-
aquatica and V. anagallis-aquatica var. nilotica in
Egypt. The main differences between the two taxa are
that subsp. anagallis-aquatica has broader leaves, a
dense inflorescence; and capsules with rounded apex,
equal to or shorter than calyx, whereas V. anagallis-
aquatica var. nilotica has smaller leaves, lax
inflorescence with fewer flowers and capsules with
acute apex, equal to or slightly longer than calyx.
There are three more taxa commonly separated
from V. anagallis-aquatica as either species or
subspecies, which are V. anagallis-aquatica subsp.
michauxii, subsp.oxycarpa, and subsp. lysimachioides
(Fischer, 1978). The first one is characterized by a
glandular-villous indumentum on stem and leaves.
Specimens from this taxon consistently clustered close
in molecular and morphometric analyses by Ellmouni et
al. (2017). V. anagallis-aquatica subsp. oxycarpa is
considered transitional between subsp. anagallis-
aquatica and subsp. lysimachioides and this is also
highlighted in the morphometric analysis (Ellmouni et
al. 2017), which grouped individuals of this taxon either
with subsp. michauxii or subsp. lysimachioides.
According to Fischer (1981), it is characterized by ovate
to elliptic leaves and apically tapering capsules.
Veronica anagallis-aquatica subsp.
lysimachioides is characterized by glabrous surface,
uniformly sessile leaves, dense inflorescence, and
capsules with roundish-obtuse apex. Öztürk and Fischer
(1982) mentioned that this taxon presents considerable
problems in its delimitation from its closest relatives,
especially from subsp. oxycarpa, which is insufficiently
understood. Though quite well characterized by
Schlenker (1936 a), V. lysimachioides has been
neglected by many authors (e.g., by Marchant, 1970;
Elenevskij,1969 & 1978). Fischer (1978, 1981) has
given a full description of the taxon. It is differentiated
by the very dense raceme; all leaves sessile; the
complete absence of glandular hairs; pedicels up to 3.5
mm; and capsules shorter than 3.5 mm, with obutse
apex. These characters are not especially characteristic
and consequently specimens formed a polyphyletic
group in the morphometric analysis of Ellmouni et al.
(2017).
Veronica catenata was known earlier in Egypt
and elsewhere as V. aquatica Bernh. (Muschler,1912;
Täckholm, 1956) or as V. anagallis-aquatica var.
aquatica (Bernh.) Nyman (Täckholm, 1974). However,
the correct name for the plant, V. catenata Pennell, was
clarified by Burnett (1950). Chrtek and Osbornová-
Kosinová (1981) concluded that the Egyptian material
should be included in V. catenata, not V. anagallis-
aquatica, and gave the Egyptian material subspecific
rank as V. catenata subsp. pseudocatenata, which goes
in line with El-Hadidi et al. (1999), Boulos (1995,
2002), and Abd El-Ghani et al. (2010, 2011). The
results of our work agree with all prevoius authors in the
occurrence of V. catenata subsp. pseudocatenata in
Egypt. However, a close relationship with V. catenata
subsp. catenata is not supported by Ellmouni et al.
(2017). Abd El-Ghani et al. (2010) suggested the
occurrence of V. catenata subsp. catenata in Egypt,
based on some collected specimens showing the
characters of V. catenata subsp. catenataand differing
from V. catenata subsp. pseudocatenata in bracts that
are shorter than the pedicels. Furthermore bracts in V.
catenata subsp. catenata become less lanceolate from
the first pedicel to the last one, with the reverse case in
V. catenata subsp. pseudocatenata , where the first and
second flower pedicels have large and leafy bracts and
then become more lanceolate.
Veronica catenata subsp. catenata resembles V.
catenata subsp. pseudocatenata in the shape of upper
leaves (ovate-lanceolate to lanceolate), inflorescence lax
(with 3-7 per cm pedicel), ± straight and spreading, and
orbicular and larger capsule . On the other hand, V.
catenata subsp. catenata resembles V. anagallis-
aquatica in having lanceolate bracts shorter than
flowersand fruiting pedicels,. The two taxa differ mainly
in lower leaves sessile, leaves that are generally broader
and shorter, pedicels patent, inflorescence less dense,
capsule normally as long as the calyx or even slightly
longer in V. catenata subsp. catenata, while V.
anagallis-aquatica has shortly petiolate lower leaves,
leaves that are generally broader and longer, pedicel ±
erect, inflorescence dense, capsule often shorter than
calyx.
For Veronica scardica, Chrtek and Osbornová-
Kosinová (1981) reported another subspecies in the
Egyptian flora named subspecies africana, which is
stated to have "sessile leaves in upper part of stem
[which] often caused misidentification with V.
anagallis-aquatica, which sometimes has its lowermost
leaves shortly petiolate. Racemes of V. anagallis-
aquatica are always opposite; while alternate" in V.
scardica subsp. africana..
V. scardica subsp. scardica and V. scardica
subsp. africana possess rhombic to suborbiculate
leaves; ; capsule 2-3.5 mm long, slightly shorter than the
calyx lobes wide, and with rounded apex, possibly vary
from each other in the size of flowers and fruits. A close
association between both taxa is not suggested by results
from Ellmouni et al. (2017) and remains to be tested
more rigorously. Separate origins are likely, especially
since European plants are restricted to heavy-metal soils
(Fischer et al. 1984) other than Egyptian plants.
Marchant (1970) reported successful artificial
crosses between Veronica scardica and V. beccabunga,
although seeds were inviable. Supposed naturally
occurring hybrids of this cross are believed to represent
instead hybrids of V. scardica and V. anagallis-aquatica
(Fischer et al., 1985).
173

Faten Y. Ellmouni et al
Veronica kaiseri was described by Täckholm
(1942) based on plants from the Sinai Peninsula. Based
on more detailed comparisons of Sinai plants (Täckholm
1956, 1974),V. kaiseri is closely related to V. scardica,
differing in the apex of capsules, which are acute in V.
kaiseri ,while rounded as in V. scardica. In herbarium
specimens, V. kaiseri is sometimes misidentified as V.
beccabunga, which resembles V. kaiseri in its habit. The
differences are seen in capsules and inflorescences.
Veronica beccabunga has opposite inflorescences,
while V. kaiseri has alternate inflorescences. There are
also differences in leaves: V. kaiseri has upper leaves
sessile or very shortly petiolate, but V. beccabunga has
all petiolate leaves. Veronica kaiseri differs from V.
anagallis-aquatica in leaves (being suborbicular to
broadly elliptic), color of the corolla (bright blue) and
(ellipsoid capsule, with acute apex) while in V.
anagallis-aquatica leaves are ovate to lanceolate,
corolla violet and the capsule is orbicular with rounded
apex. It, thus, resembles mostly V. anagalloides,
especially subsp. heureka, from which it is
differentiated by the acute capsule apex, alternate
inflorescence and leaves suborbicular to broadly elliptic
with short petiole below rather than ovate leaves that are
all sessile.
Chrtek and Osbornová-Kosinová (1981) stated
that V. kaiseri was only known from the Sinai Peninsula,
especially its southern granite mountain region. Several
studies (Boulos, 1995; El-Hadidi and Fayed, 1995; El-
Hadidi et al., 1999; Boulos, 2002) agree with Chrtek and
Osbornová-Kosinová(1981) and Täckholm (1974) in
the occurrence of Veronica kaiseri only in Sinai.
However, Abd El-Ghani et al. (2010) collected some
specimens from Nile Delta (Fayoum area), which
resemble V. kaiseri in having alternate and lax
inflorescence with few flowers and capsules with acute
apex. The specimens differ from the holotype kept in
Agriculture Museum Herbarium in leaf morphology
(being elliptic-ovate in Fayoum specimens vs. ovate and
petiole length of the lower leaves (1mm long vs. 2-3
mm). More recently ,the species was discovered in
Jordan by Gregor & Albach in von Raab-Straube &
Raus (2016).
Veronica undulata is similar to V. anagallis
aquatica in their subglobose capsules, (basally and
apically equal in width), with rounded or emarginated
apex. In V. anagallis-aquatica, pedicel curved, at an
acute angle with inflorescence axis; inflorescence less
than 1 cm wide, glabrous , while pedicel straight, at a
right angle with inflorescence axis; inflorescence1–1.5
cm wide, and sparsely glandular pubescent in V.
undulata. It was not included in the analyses by
Ellmouni et al. (2017) but its distinction is supported by
the specific distribution area in Eastern Asia and the
hexaploid level not reported in any other taxon of the
subsection (Albach et al. 2008).
Acknowledgments
The authors wish to thank the Egyptian Science and
Technology Development Fund (STDF) and the
German Academic Exchange Program (DAAD) for
funding our project (GESP 5375). We thank Metin
Armagan, and curators of the herbaria HUB, VANF,
GAZI and BRUN for the provision of herbarium
specimensand Dr. Hassan Mansour, who kindly
provides the specimens of V. kaiseri from Sinai.
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(2004). Phylogeny of Veronica—A combination of
molecular and chemical evidence. Plant Biology, 6:
673–682.
Wagstaff, S.J. & Garnock-Jones, P.J. (1998).
Evolution and biogeography of the Hebe complex
(Scrophulariaceae) inferred from ITS sequences. New
Zealand Journal of Botany, 36: 425-437.
Walters, S.M. and Webb, D.A. (1972). Veronica. - In:
Tutin T. G. & al. (Ed.), Flora Europaea 3: 242-251. -
Cambridge, University Press.
Walters, S.M. (1975).Veronica L. In.C.A. Stace (Ed.),
Hybridization and the Flora of the British Isles, 371-
372. Academic Press, New York.
Williams, I.A. (1929). A British Veronica hybrid.
Journal of Botany, London, 67: 23-24.
176

Systematic treatment of Veronica L. Section Beccabunga
1
Table 1: Species recognized and reported in earlier studies. “+”in different columns 1-13 refers to presence of species in studies of the respective author, “-“
refers to absence and “X” to presence as synonym in respective study. Author name: 1- Ellmouni et al. (this publication) 2- Podlech (2012). 3- Abd El-Ghani
et al. (2010). 4- Martínez Ortega et al. (2009) 5- Hong and Fisher (1998) 6- Sellers (1983). 7- Öztürk and Fischer (1982). 8- Fischer (1981). 9- Fischer (1978).
10- Walters and Webb (1972). 11-Borissova (1955). 12- Schlenker (1936a) 13- Pennell (1935).
Three Series:
Borissova
(1955)
Two
Subsections:
Keller (1942)
Three
Subsections Species Subspecies or
variates 1 2 3 4 5 6 7 8 9 10 11 12 13
Subsection :
Peregrinae
V. peregrina V. peregrina + + - + + - - - - + - - +
Series:
Anagallides
Subsection:
Anagallides
Subsection:
Anagallides
V. anagallis-
aquatica
V. anagallis-
aquatica
+ + + + + + + + + + + + +
V. anagallis-
aquatica var.
nilotica
+ - + - - - - - - - - + -
V. anagallis-
aquatic subsp.
lysimachioides
+ - - - - - + + + - X X -
V. anagallis-
aquatic subsp.
michauxii
+ + - - - - + + + - X X -
V. anagallis-
aquatic subsp.
oxycarpa
+ + - - X - + + + - X X -
V. anagalloides
V. anagalloides
-
-
-
-
+
+
-
-
+
+
+
+
-
V. anagalloides
subsp.
anagalloides,
+ + + + - - + + - - - - -
V. anagalloides
subsp. heureka
+ + + - - - + + - - - - -
V. anagalloides
subsp.
taeckholmiourm,
+ - + - - - - - - - - - -
177

Faten Y. Ellmouni et al
* as V. beccabunga subsp. americana** partly as anagallidiformis (*)
V. anagalloides X
V. anagallis-
aquatica
+ - - + - - + - - - - - -
V. anagallidiformis
V. anagallidiformis
-
-
-
-
-
-
-
-
-
-
+
-
-
V. catenata
V. catenata,
-
-
-
+
-
+
-
-
-
+
-
-
*
V. catenata var.
catenata,
+ - + - - - - - - - - - -
V. kaiseri
V. kaiseri
+
-
+
-
-
-
-
-
-
-
-
-
-
V. X lackschewitzii
V. X lackschewitzii
+
-
-
+
-
-
-
-
-
-
-
-
-
V. scardica
V. scardica,
+
-
-
-
-
+
+
+
+
+
+
+
-
V. scardica subsp.
africana
+ - + - - - - - - - - -
V. poljensis
V. poljensis
+
-
-
-
-
-
+
-
-
-
+
+
-
V.undulata
V.undulata
+
+
-
-
+
-
-
+
-
-
-
-
Series:
Oxycarpae
V. beccabungoides
V. beccabungoides
-
-
-
-
-
-
-
+
X
-
+
-
-
V. michauxii
V. michauxii
-
X
-
-
-
-
X
X
X
-
+
+
-
V. lysimachioides
V. lysimachioides
-
-
-
-
-
-
X
X
X
-
+
+
-
V. oxycarpa
V. oxycarpa
-
X
-
-
+
-
X
X
X
-
+
+
-
Series:
Beccabungae Subsection:
Beccabungae
Subsection:
Beccabungae
V. americana
V. americana + - - - - * - - - - + + +
V. beccabunga
V. beccabunga
-
-
-
-
-
+
X
+
+
+
+
+
+
V. beccabunga var.
beccabunga.,
+ - + + - - + - - - - - -
V. beccabunga
subsp. abscondita,
+ - - - - - + + - - - - -
V. beccabunga
subsp. muscosa.
+ + - - + - - + - - - - -
Number of taxa
20
7
10
6
5
6
11
12
7
5
11
10
3
178

Systematic treatment of Veronica L. Section Beccabunga
Appendix 1 : Voucher information for taxa used in the systematic revision. A total of 100 specimens representing 20 taxa of section
Beccabunga, was used in the systematic revision
Latin Name
Voucher
Locality
Latitude
Longitude
1
Veronica sp.
Jaroslav 613771, BRNU
620815
Brehy, Czech Republic
50° 4' N
15° 34' E.
2
Veronica sp.
Jaroslav 747670, BRNU
620816
Sezemic, Czech Republic
50° 4' N
15° 51' E.
3
Veronica sp.
Jaroslav 797430, BRNU
620817
Hradišťko II, Czech Republic
50° 7' N
15° 24' E.
4
V. americana
Don Mansfield 12-431, CIC
43770
Idaho, Owyhee County,U.S.A.
42° 60' N
116° 76' W
5
V. americana
R. R. Halse 8268, SRP 38705
Oregon, Benton County, U.S.A.
44° 29 N
123° 61' W
6
V. americana
Hans Hallman 1151,
USFS0049862
Montana, Carter County,U.S.A.
45° 90' N
104° 45' W
7
V. americana
Ashley Van Hoose 19,
HPSU008353
Washington, Clark County,U.S.A.
45° 44' N
121° 37' W
8
V. americana
L. Price 37, SRP 5589
Alaska, Susitna County,U.S.A.
62°15' N
150°10' W
9
V. americana
Gerald B. Straley 4303, UBC
V193722
British Columbia, Canada
48° 51' N
123° 18' W
10
V. americana
(V.beccabunga subsp. americana )
V. Wagner 14308, ID119478
Boundary County,Idaho, U.S.A.
48° 35' N
116° 46' W
11
V. anagallis-aquatica
F. Yousef 5, Herbarium
Fayoum university 18
Al Gharaq, ATSA, El- Fayoum, Egypt
29° 7' N
30° 41'E
12
V. anagallis-aquatica
Jaroslav 613771, BRNU
620814
Brehy, Czech Republic
50° 4' N
15° 34' E.
13
V. anagallis-aquatica
Danihelka 07/034, BRNU
597098
Moravia, Czech Republic
48° 47' N
16° 40' E
14
V. anagallis-aquatica
Danihelka, BRUN 617636
Moravia, Czech Republic
48° 51' N
16° 36' E
15
V. anagallis-aquatica
Scalone 27.06
0.08, OLD
Tsagveri, Georgia
41° 47' N
43° 28' E
16
V. anagallis-aquatica subsp.
anagallis-aquatica
O. Karabacak 1018, VANF
B9, Van: Muradiye, Turkey
38°59' N
43° 46' E
179

Faten Y. Ellmouni et al
17
V. anagallis-aquatica
Ö. Eyübophe 1990, GAZI 1723
A4, Ankara, Kizilcahamam, Turkey
40°28'N
32° 38' E
18
V. anagallis-aquatica
M. Kucuk 12,5,GAZI
Kürtün, Gümüşhane, Turkey
40° 42'N
39° 05'E
19
V. anagallis-aquatica spp.
anagallis-aquatica
E.Akcicek2162, GAZI
B3, Afyon: Kumalar Dağı, Haydarlı, Turkey
38°45'N
30°33'E
20
V. anagallis-aquatica
H.Duman 1988, GAZI 2350
C6, Kahramanmaras,Turkey
37° 34' N
36° 56' E
21
V. anagallis aquatica
Uzunhisarcikli 1446, GAZI
B5 Kayseri: Korumaz Dagi,Turkey
38°45' N
35°26' E
22
V. anagallis-aquatica spp.
anagallis-aquatica
A. Güner 6535, HUB
Rize camlihemsin yaylas, Turkey
40° 57' N
41° 1' E
23
V. anagallis-aquatica
M. Vural 1985,GAZI 2484
Kars,Arpaçay, Bardaklı,Turkey
40° 49' N
43° 28' E
24
V. anagallis-aquatica spp.
anagallis-aquatica
A. Duran 1992, GAZI 1453
Çankırı, Atkaracalar Demirli,Turkey
40° 50' N
33° 03' E
25
V. anagallis-aquatica
C. Birden 1409 , GAZI
kırıkkale delice büyükavşar,Turkey
39° 53' N
32° 54' E
26
V. anagallis-aquatica
H. Pesmen 4527, HUB 25015
Kemer Kumluca yolu Ulupınar Antalya, Turkey
36°27'N
30° 25'E
27
V. anagallis-aquatica
H. Pesmen 442,3, HUB 25011
Adrasan Kumluca Antalya, Turkey
36°27'N
30° 25'E
28
V. anagallis-aquatica spp.
anagallis-aquatica
ADK 22.05.2003, HUB 1282
Duzce, Akçakoca, Karatavuk Köyü, Turkey
36°27'N
30° 25'E
29
V. anagallis-aquatica
B. Mutlu 1391, HUB 24985
Isparta şarkikaraağaç kızıldağ Milli parkı, Turkey
38°02'N
31° 21'E
30
V. anagallis-aquatica
E.Vitek 07-1096 W
Armenia (I Didn’t have photo to know the main locality)
?
?
31
V. anagallis-aquatica
Albach D3-1, OLD
Belek, Turkey
36°54' N
30°59' E
32
V. anagallis-aquatica
E. George 332, CIC 44236
Oregon, Malheur County,U.S.A.
42° 54° N
117° 16' W
33
V. anagallis-aquatica
D. Giblin 3912, SRP 47299
Idaho, Adams County, U.S.A.
45° 11 N,
116° 75' W
34
V. anagallis-aquatica
Joy Mastrogiuseppe 6350,
ID119442
Washington, Kittitas County, U.S.A.
47°10' N
120°55' W
35
V. anagallis-aquatica
Susan Bernatas 172, ID
100006, ID119451
Colorado, Teller County,U.S.A.
38° 51' N
105° 10' W
36
V. anagallis-aquatica
Terry Taylor 108, UBC
V224883
British Columbia,Canada
53° 43' N
127° 20' W
37
V. anagallis-aquatica
Neuffer, Hurka, Friesen
30.07.2004, OSBU 15652
Kirgisische Republik, Tien-shan, Schlucht kurpsi,
enrlang des Flusses, Atoinoksky Gebirgskette,
Kyrgyzstan
41° 30' N
72° 19' E
38
V. anagallis-aquatica var. nilotica
F. Yousef 6, Herbarium
Fayoum university 19
Al Gharaq, ATSA, El- Fayoum, Egypt
29° 7'56 N
30°41'E
180

Systematic treatment of Veronica L. Section Beccabunga
39
V. anagalloides
Danihelka 10/124 BRUN
617635
Moravia,Czech Republic
48° 51' N
16° 36' E
40
V. anagalloides
Danihelka 07/037 BRUN
597103
Moravia,Czech Republic
48° 51' N
16° 36' E
41
V. anagalloides
Danihelka 07/042 BRNU
59112
Moravia,Czech Republic
48° 45' N
16° 44' E
42
V. anagalloides
R. Scalone 29.06.08, OLD
S5, Tsodniskari, Georgia
41° 47' N
46° 10' E
43
V. anagalloides
R. Scalone 01.07.08, OLD
Telavi, Georgia
41°54' N
45° 22' E
44
V. anagalloides
R. Scalone 30.06.08, OLD
S5, Tsodniskari, Georgia
41° 47' N
46° 10' E
45
V. anagalloides
R. Scalone 01.07.08, OLD
Georgia
° ' N
° ' E
46
V. anagalloides
Ali A. Dönmēz 1559, HUB
25028
Turkey
° ' N
° ' E
47
V. anagalloides
Ekim 7793 GAZI
Bingöl - Elazig, Turkey
38 ° 68' N
39° 22' E
48
V. anagalloides
H.Duman 16.3.1992, GAZI
Mugla, Turkey
36 ° 58 ' N
28 ° 41' E
49
V. anagalloides
F. Özgökce 8424, VANF
Özalp, Van Province Turkey
38° 39' N
43° 59' E
50
V. anagalloides subsp.
anagalloides
F. Yousef 9, Herbarium
Fayoum university 20
Rasheed El-Mahmoudeya, Beheira,Egypt
31°19' N
30°26' E
51
V. anagalloides subsp. heureka
F. Yousef 11, Herbarium
Fayoum university 21
El- Fayoum, Egypt
29°18'N
30° 50'E
52
V. anagalloides subsp. heureka
O. Karabacak 7537, VANF
12967
Van, Ercis, Turkey
38°14'N
43°23' E
53
V. anagalloides subsp.
taeckholmiorum
F. Yousef 12, Herbarium
Fayoum university 22
El- Fayoum, Egypt
29°20'N
30°48'E
54
V. beccabunga subsp.
beccabunga.
F. Yousef 15, Herbarium
Fayoum university23
Al Siauf, Alexandria Governorate,Egypt
31°13' N
30° 0'1E
55
V. beccbunga subsp. beccabunga
R. Scalone 27.06.08, OLD
Borjomi-Bakuriani-Akhalkalaki, Georgia
41° 47' N
43°28' E
56
V. beccabunga subsp. abscondita
F. Özgökce 7252, VANF
Özalp, Van Province, Turkey
38° 39' N
43° 59' E
57
V. beccabunga subsp. beccabunga
Murat 5807, VANF 5559
Gürpınar, Turkey
38°32' N,
43°41' E
58
V. beccabunga subsp. abscondita
Murat 7343, VANF 5557
Gürpınar, Turkey
38°32' N,
43°41' E
59
V. beccabunga subsp. abscondita
Murat 1686,VANF 2932
Van muradiye, Turkey.
(Elevation 2600)
38° 59' N
43° 46' E
60
V. beccabunga subsp. abscondita
Murat 1686,VANF 2932
Van muradiye, Turkey.
(Elevation 2660,)
38° 59' N
43° 46' E
61
V. beccabunga subsp. abscondita
Murat 4158, VANF 2928
Van muradiye, Turkey
38° 59' N
43° 46' E
181

Faten Y. Ellmouni et al
62
V. beccbunga subsp. beccabunga
ü. Güler 1986, GAZI
Kirikkale, Keskin, Turkey
39° 40' N
33 ° 36 ' E
63
V. beccabunga subsp. beccabunga
A. Güner 2545, HUB 25056
Rize: Camlihemsin, Ayder, Turkey
41 ° 02 ' N
41° 01 ' E
64
V. beccbunga subsp. beccabunga
A. Güner 3686, HUB 25053
Rize: Camlihemsin, Ayder, Turkey
41 ° 02 ' N
41° 01 ' E
65
V. beccabunga subsp. muscosa
Neuffer, Hurka, Friesen
27.07.2004, OSBU 15560
Kirgisische Republik, Pamiro-Alai, Alaisky
Gebirgskette,FlussnKok-Su/ Kyrgyzstan
39° 39' ' N
72° 08' E
66
V. americana
V. Wagner 14308, ID119478
Boundary County, Idaho, U.S.A.
48° 35' N
116° 46' W
67
V. beccabunga subsp. beccabunga
E.R. Manton 1608, UBC
V238612
British Columbia, Canada
49° 93' N,
123° 36' W
68
V. catenata
Jaroslav 613771, BRNU
620813
Brehy, Czech Republic
50° 4' N
15° 34' E.
69
V. catenata
J. Danihelka 07/032, BRNU
597095
Moravia, Czech Republic
48° 45' N
16° 44' E
70
V. catenata
J. Danihelka 07/170, BRNU
596968
Kostelec, Czech Republic
49° 41' N
13° 00' E
71
V. catenata
David Giblin 2390, ID 161075,
ID119504
Washington, Adams County, U.S.A.,
46° 89' N
119.19' W
72
V. catenata
John Corman s.n., MONT
79674
Montana, Beaverhead County, U.S.A.,
45° 21' N
112.63' W
73
V. catenata
W. Owen 85023, SRP 10066
Idaho, Owyhee County, U.S.A.
42° 60' N
116° 76' W
74
V. catenata
R. Dale Thomas 9721,
WWB019512
Missouri, Shannon County, U.S.A.
37°12' N
91°26' W
75
V. catenata var. catenata
F. Yousef 1, Herbarium
Fayoum university 24
El- Fayoum,Egypt
29°18'N
30°50' E
76
V. catenata subsp. pseudocatenata
F. Yousef 4, Herbarium
Fayoum university 25
El- Fayoum, Egypt
29°18' N
30°50' E
77
V. kaiseri
F. Yousef 17, Herbarium
Fayoum university 26
El- Fayoum, Egypt
29°18' N
30°50' E
78
Veronica x lackschewitzii
J. Danihelka 07/036, BRNU
597100
Moravia, Czech Republic
48° 47' N
16° 40' E
79
Veronica x lackschewitzii
J. Danihelka 07/036, BRNU
597101
Moravia,Czech Republic
48° 47' N
16° 40' E
80
Veronica x lackschewitzii
J. Danihelka 07/036, BRNU
597102
Moravia, Czech Republic
48° 47' N
16° 40' E
81
V. anagallis-aquatica spp.
lysimachioides
O. Karabacak. 7538, VANF
12965
Van, Ercis, Turkey
38° 14' N
43° 23' E
182

Systematic treatment of Veronica L. Section Beccabunga
82
V. anagallis-aquatica spp.
lysimachioides
Murat 8926 VANF, 5562
Gürpınar, Turkey
38°32' N,
43°41' E
83
V. anagallis-aquatica spp.
lysimachioides
S. Yildirimli 2359, HUB 24992
Ovacık, Tunceli, Turkey
39° 21 ' N
39° 12' E
84
V. anagallis-aquatica spp.
lysimachioides
E.Akcicek 0638,HUB 25206
Erkenek, Turkey
37° 55 ' N
37° 56' E
85
V. anagallis-aquatica
subsp.michauxii
O. Karabacak 5487, VANF
12966
Van, Ercis, Turkey
39°09' N
43° 22' E
86
V. anagallis-aquatica spp.
michauxii
Demirkus 2152, HUB 24994
Erzurum: Oltu, Azort, Turkey
40° 32' N
41° 58' E
87
V. anagallis-aquatica spp.
michuaxii
Demirkus 2740, HUB 24995
Kars: Göle, Turkey
40° 36' N
43° 03' E
88
V. anagallis-aquatica spp.
michauxii
Demirkus 2955, HUB 24999
Kars: Posof, Turkey
41° 30' N
42°43' E
89
V. anagallis-aquatica spp.
michuaxii
A. Güner 5186, HUB 24989
Erzurum: Ispir, Ikizdere arasi Turkey
40° 28' N
40° 59' E
90
V. anagallis-aquatica spp.
oxycarpa
Murat 1786, VANF 2918
Muradiye İBesparmak, Turkey
38°52' N
43°46' E
91
V. anagallis-aquatica
subsp.oxycarpa
F. Özgökce 4851,VANF
Özalp, Van Province, Turkey
38° 39' N
43° 59' E
92
V. anagallis-aquatica
subsp.oxycarpa
M. Armagan 1973,VANF 5832
Guzeldere, Turkey
39° 61' N
41°4' E
93
V. anagallis-aquatica
subsp.oxycarpa
H. Sumbül 2529, HUB 25305
Kazancı Köyü, Turkey
41°09' N
36° 07' E
94
V. anagallis-aquatica
subsp.oxycarpa
A. Güner 4078, HUB 25000
Rize: Camlihemsin, Hisarcik-Siraköy arasi, Turkey
41 ° 02 ' N
41° 01 ' E
95
V. anagallis-aquatica
subsp.oxycarpa
S. Yildirimli 3491, HUB 25001
Tunceli ovacık, karagöl, Turkey
39 ° 21' N
39 °12 ' E
96
V. scardica
B Motto 880, HUP 25361
Isparta, Turkey
37° 47'N
30° 30' E
97
V. scardica subsp. africana
F. Yousef 13, Herbarium
Fayoum university 27
El- Fayoum, Egypt
29° 18' N
30° 50' E
98
V. poljensis
F. Öztürk 1081, VANF
Van Edremit, Turkey
38° 25' N
43° 15' E
99
V. peregrina
J. F. Smith 9734, SRP 39910
Idaho, Adams County,U.S.A.
44° 42' N
116° 32' W
100
V. peregrina
B. Ertter 19952, SRP 46473
Idaho, Ada County,U.S.A.
43° 638' N
116° 197' W
183