The genus Oenothera (Onagraceae) in Belgium
† Krzysztof Rostański1 and Filip VerlooVe2
1
Faculty of Biology and Environmental Protection, Department of Plant Systematics, Silesian
University (Katowice, Poland)
2
author for correspondence: Botanic Garden of Meise, Nieuwelaan 38, B-1860 Meise, Belgium
[ilip.verloove@br.fgov.be]
Illustrations by S. Bellanger (ig. 2-5), W. Vercruysse (7-8), R. Barendse (10: lowers and stem; 11;
17), J. Pottier (10: hairs), R. Jean (21), P. Kalinowski (22) and F. Verloove (the others).
AbstrAct. – The genus Oenothera (Onagraceae) was thoroughly and critically revised in
Belgium. This revision is based on the study of several hundreds of older and more recent
herbarium specimens from all major public and some private herbaria as well as on additional ieldwork carried out since 2000. A brief history of the genus Oenothera in Belgium is
provided. Dificulties with regard to taxonomy and nomenclature are discussed. The present
treatment of Oenothera follows the ‘European school’ and is thus based on a narrow species concept for subsection Oenothera. In the present paper only species that have been
recorded since 1950 are dealt with. As such 27 species (including stabilized hybrids) have
been upheld: Oenothera biennis, O. cambrica (with two varieties), O. canovirens, O. delexa, O. depressa, O. ersteinensis, O. fallax, O. fruticosa subsp. glauca, O. glazioviana, O.
hirsutissima, O. issleri, O. lindheimeri, O. nuda, O. oehlkersii, O. paradoxa, O. parvilora,
O. pycnocarpa, O. rosea, O. royfraseri, O. rubricalyx, O. rubricaulis, O. rubricauloides, O.
stricta, O. subterminalis, O. victorinii, O. villosa and O. wratislaviensis. Nine additional species were only recorded before 1950. Oenothera is represented in Belgium by the following
sections, subsections and series: sections Gaura, Hartmannia, Kneifia and Oenothera, the
latter furthermore represented by subsections Munzia, Oenothera (with the series Devriesia,
Linderia, Oenothera and Rugglesia) and Raimannia. An identiication key and photographs
(mostly of typical and/or widespread species) are presented. Each species is concisely compared with related or similar species and its current distribution and naturalization status in
Belgium are assessed.
résumé. – Le genre Oenothera (Onagraceae) en Belgique. Le genre Oenothera (Onagraceae) fut inventorié en Belgique. Cette révision taxonomique est basée sur l’étude de plusieurs centaines d’échantillons d’herbier (anciens ainsi que plus récents) et des prospections
de terrain à partir de 2000. L’histoire du genre Oenothera en Belgique est brièvement présentée et des dificultés concernant la taxonomie et nomenclature commentées. La présente
révision du genre Oenothera suit la dite ‘école européenne’ pour la sous-section Oenothera.
Dans cet article seulement les 27 taxons (y inclus quelques espèces hybridogènes) récoltés
après 1950 sont retenus : Oenothera biennis, O. cambrica (avec deux variétés), O. canovirens, O. delexa, O. depressa, O. ersteinensis, O. fallax, O. fruticosa subsp. glauca, O. glazioviana, O. hirsutissima, O. issleri, O. lindheimeri, O. nuda, O. oehlkersii, O. paradoxa, O.
parvilora, O. pycnocarpa, O. rosea, O. royfraseri, O. rubricalyx, O. rubricaulis, O. rubricauloides, O. stricta, O. subterminalis, O. victorinii, O. villosa et O. wratislaviensis. Neuf
taxons supplémentaires ont été récoltés uniquement avant 1950. Les taxons précités appartiennent aux sections, sous-sections et séries suivants : les sections Gaura, Hartmannia,
Kneifia et Oenothera, ce dernier en plus avec les sous-sections Munzia, Oenothera (avec les
séries Devriesia, Linderia, Oenothera et Rugglesia) et Raimannia. Une clé pour l’identiication des espèces est présentée ainsi que des photos des espèces les plus caractéristiques et/ou
les plus répandues. Chaque espèce est comparée avec des espèces proches ou similaires et sa
distribution et son statut (fugace/naturalisé) sont fournis.
sAmenvAtting. – Het genus Oenothera (Onagraceae) in België. Het geslacht Oenothera
(Onagraceae) werd kritisch gereviseerd in België. Deze revisie is gebaseerd op de studie van
enkele honderden herbariumspecimens (uit de belangrijkste publieke herbaria van België
en uit enkele kleinere private herbaria), alsook op veldwerk, uitgevoerd sinds 2000. De hisDumortiera 106/2015: 12-42
12
toriek van het geslacht Oenothera in België wordt geschetst en problemen met betrekking tot
de nomenclatuur en taxonomie worden aangekaart. De huidige revisie volgt de zogenaamde
‘Europese school’ en hanteert dus een eng soortsbegrip voor de subsectie Oenothera. Alleen de 27 taxa (inclusief enkele gestabiliseerde hybriden) die na 1950 in België zijn waargenomen, worden uitgesleuteld en besproken: Oenothera biennis, O. cambrica (met twee
variëteiten), O. canovirens, O. delexa, O. depressa, O. ersteinensis, O. fallax, O. fruticosa
subsp. glauca, O. glazioviana, O. hirsutissima, O. issleri, O. lindheimeri, O. nuda, O. oehlkersii, O. paradoxa, O. parvilora, O. pycnocarpa, O. rosea, O. royfraseri, O. rubricalyx, O.
rubricaulis, O. rubricauloides, O. stricta, O. subterminalis, O. victorinii, O. villosa en O.
wratislaviensis. Negen soorten werden uitsluitend voor 1950 in België aangetroffen. Het geslacht Oenothera wordt in België vertegenwoordigd door de volgende secties, subsecties en
series: de secties Gaura, Hartmannia, Kneifia en Oenothera, deze laatste bovendien door de
subsecties Munzia, Oenothera (met de series Devriesia, Linderia, Oenothera en Rugglesia)
en Raimannia. Een determinatiesleutel en foto’s (meestal van typische en/of wijd verspreide
soorten) worden eveneens gepresenteerd. Elke soort wordt vergeleken met gelijkaardige of
verwante soorten en de huidige verspreiding en inburgeringsgraad in België worden kort
besproken.
Introduction1
The genus Oenothera L. is not native to Belgium. Although O. biennis is naturalized in this country since at
least the 18th century (Roucel 1792), the exact number
and the true identity of the other Belgian taxa always
remained unclear. This is primarily because of the very
complex taxonomy of the genus, especially with regard
to section Oenothera subsection Oenothera. For this
group, there are two very divergent taxonomical views.
According to Dietrich et al. (1997) the group is made up
of only 13 species. This concept is usually referred to as
the ‘American school’ but this is misleading. As early as
1958, the American botanist and geneticist Ruggles Gates
already presented a taxonomic concept for Oenothera that
for the greater part confers with the current ‘European’
species concept. Moreover, Dietrich is German. Rostański
(1985), who represents the so-called ‘European school’,
claimed there were 85 species and 23 hybrids. Moreover, in the past decades several additional new taxa have
been described (e.g. Jehlík & Rostański 1995, Delipavlov
1998, Rostański & Ramst 2001, Rostański et al. 2004,
Rostański 2007, Deschâtres et al. 2013).
As a result of a peculiar breeding mechanism (permanent translocation heterozygosity2) species from subsection Oenothera cross freely, producing off-spring that differs from either parent and developing populations that
consist of many different but in fact continuous genotypes.
As such, numerous, closely similar but morphologically
well-delimited entities are formed. These newly arisen
The results presented in this paper are almost exclusively based on
the determinations by the irst author. The text, however, was written
by the second author and approved by Dr. Adam Rostański, son of the
irst author.
2
Permanent translocation heterozygosity is a cytological phenomenon
in which an organism is true breeding for a set of translocations involving all or part of the chromosomes.
1
plants breed more or less completely true. They may be
intermediate but are usually nearer one parent than the
other and the degree of resemblance may vary from character to character. Some hybrids even resemble one parent
species more strongly in one stage of development and
the other parent at a later stage (Gates 1933). Also, when
Oenothera species spread to new habitats, new ‘species’
(or mere morphotypes?) adapted to new conditions, may
arise (Tokhtar & Wittig 2003). These ‘microspecies’ were
included in one of the 13 recognized species (or hybrids
of these) by Dietrich et al. (1997). As such, for example
for O. biennis, there are no less than 68 ‘synonyms’. According to Rostański, whose work was for the greater part
based on the earlier publications of Renner (1942, 1950,
1956) these mutants and hybrids are, once more or less established, preferably considered as independent species.
The choice between both concepts is not straightforward and merely a matter of opinion and there is no
consensus whatsoever. However, this species concept is
perhaps not applicable in North America, where the genetic variation is greater than in Europe. Wisskirchen &
Haeupler (1998) presented, without judgement of their
respective merits, both concepts next to one another. In
our view, we consider that Dietrich et al. (1997) presented
a too broad species concept. Under O. biennis, for example, numerous, easily differentiated species, are grouped
together (e.g. O. rubricaulis) that altogether often hardly
resemble O. biennis s.str. Other examples: O. perangusta is included with O. oakesiana, whilst the (possibly)
conspeciic O. ersteinensis is included under O. biennis.
Also, O. delexa is included in O. parvilora whilst both
are not in the least related and actually belong to different,
relatively distinct series. See also a more lengthy discussion in Rostański et al. (2010). In Europe, since Renner
(1917), who was the founding father of the European
school, a narrow species concept has been applied most of
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
13
the time although recently, in Flora Iberica, an exception
was made (not surprisingly, the genus Oenothera being
prepared by Dietrich; see Dietrich 2000). Van der Meijden
(2005a) originally also opted for this broad species concept for the 23rd edition of Heukels’ Flora (van der Meijden 2005b), yet eventually he accepted O. delexa and O.
fallax as two taxa, which were not recognized by Dietrich
et al. (1997). In the most recent edition of the Belgian
lora (Lambinon & Verloove 2012) numerous microspecies are accepted in taxonomically similar, complex genera (e.g. the apomictic genera Rubus and Taraxacum). For
these reasons, it would be at least useful and perhaps even
advisable to follow the European school, applying a narrow species concept for Oenothera in Belgium and neighbouring territories. This will allow tracking of changes
in range and frequency and to distinguish patterns and
mechanisms of distribution; application of a wide species
concept would lead to considerable loss of information
(Rostański & Karlsson 2010). Worthwhile mentioning is
that molecular phylogenetic studies have been performed
in the genus Oenothera (e.g. Krakos et al. 2014) but not
yet in subsection Oenothera.
In 2000 the irst author began a systematic revision
of the Belgian Oenothera collections, stored in the main
public herbaria (the herbarium of the Botanic Garden
Meise, BR; the herbarium of the University of Liège, LG;
and the herbarium of the University of Ghent, GENT) and
some smaller private herbaria. Since then the second author attempted to document, as much as possible, the variation of Oenothera currently encountered in favourable
habitats in Belgium (e.g. in seaports, coal mining spoil
heaps, sand pits, landills, railway infrastructure, sand depots, etc.).
In this paper, all species reliably recorded since 1950
are keyed out and briely discussed. Those species that
exclusively occurred as wool aliens in the Vesdre valley
and species that were last collected before 1950 are excluded because they are, for the present day ield botanist,
of lesser interest. These species are merely cited in the
classiication scheme (preceded by an asterisk) and their
respective records are included in the appendix of this paper.
An overview of the studies concerning Oenothera in
Belgium and Europe
As previously mentioned the genus Oenothera was not
well documented in Belgium. The majority of loras of the
19th and the irst half of the 20th century mentioned only O.
biennis and O. muricata (auct. belg. non L.) (e.g. Durand
1899), occasionally along with a few ephemeral aliens
such as O. laciniata or O. rosea. Goffart (1945) added
O. glazioviana; later, it turned out that this species was
already found in Belgium since at least 1868 (Verloove
2006a). De Langhe et al. (1967) listed the following species: O. biennis, O. erythrosepala (syn.: O. glazioviana),
O. parvilora and the ephemeral aliens O. laciniata and
O. stricta. Around the mid-seventies of the 20th century
the Belgian Oenothera collections of the herbarium of the
Botanic Garden Meise (BR) were revised for the irst time
by an expert (Jean 1975). He conirmed the reported species of De Langhe et al. (1967) and added the following:
O. ammophila, O. atrovirens (syn.: O. cruciata), O. fallax
(as O. biennis × O. lamarckiana), O. hungarica (syn.: O.
depressa) and O. longilora. Recently, it became clear that
O. ammophila as well as O. longilora were misidentiications, the corresponding collections belonging respectively to O. parvilora and O. afinis (Verloove 2006a). In the
subsequent editions of the Flora of Belgium the results of
Jean’s revision were maintained. Moreover, in Lambinon
et al. (1998) O. nuda was added, a species observed in
northern France (Jean 1990). In Lambinon et al. (2004)
some provisional results from the current study were already included (mostly based on Verloove 2002). Most
notable was that the Belgian populations of the ‘small
lowered evening primrose’ were attributed to O. delexa
and no longer to O. parvilora (see further). Verloove
(2006a) presented an overview of all observed species in
Belgium prior to 2005 and inally, in the most recent edition of the Belgian lora (Lambinon & Verloove 2012),
an extra species was keyed-out (the relatively widespread
and well deined O. fallax). The list of ephemeral adventives was also updated.
Elsewhere in Europe, in the last decades, a reasonably
good picture has been formed of the diversity and spread
of the genus (mostly as a result of the continuous revisions
by the irst author). For many European regions there are
overviews available by now. The most important (with
identiication keys and/or good illustrations) are presented in table 1, chronologically listed per country or region.
These publications are often relevant for the Oenothera
lora in Belgium as well. Especially north, central and east
European revisions usually include species that are also
found in Belgium. In contrast, for southern Europe (e.g.
Italy, Portugal and Spain) there are proportionally more
taxa that are not found in Belgium (and vice versa).
Finally, Dietrich (1999) offers an interesting and upto-date overview of the genus Oenothera in cultivation.
Problems concerning the identiication of Oenothera
Unfortunately, the complex taxonomy is not the only
problem in the identiication of Oenothera species. Some
important diagnostic characteristics disappear on drying
or become less visible: the presence or not of hairs with a
red bulbous base, the colour and the size of the petals and
their length and width, the red pigmentation of (parts of)
the stem, the colour of the sepals, the colour of the midrib
of the leaves, etc. For these reasons, it is of the utmost
importance that specimens are either identiied from fresh
material, or with the help of ield notes (especially dimensions and colour of the different plant parts). Moreover,
the identiication of many species needs to be carried out
on lowering and fruit bearing plants.
It also is important to know that the identiication of
Oenothera species only makes sense during the irst low-
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
14
Table 1. An overview of revisions of Oenothera from Europe.
Countries or regions
Publications
Europe
Rostański et al. (2010)
Germany
Hudziok (1968), Otto (1970), Gutte
& Rostański (1971), Rostański
& Schnedler (1991), Gutte (1998),
Haeupler & Mür (2000), Lang (2001),
Jäger & Werner (2005), Rostański
& Meierott (2006)
France
Rostański et al. (1994), Tison & de
Foucault (2014)
Great Britain
Rostański (1982), Sell & Murrell
(2009)
Austria
Rostański & Forstner (1982)
Eastern Europa (Baltic
States, Russia, Belarus,
Ukraine and Moldova)
Rostański et al. (2004)
Scandinavia (Denmark, Rostański (2006), Rostański (2007),
Finland, Iceland, Norway Rostański & Karlsson (2010)
and Sweden)
Czech and Slovak
Republic
Jehlík & Rostański (1979), Jehlík
& Rostański (1995), Jehlík (1997)
Serbia and Montenegro
Zlatković et al. (1998); Rakaj & Ros
tański (2009)
Spain
Dietrich (2000)
Portugal
Rostański (1991)
Italy
Soldano (1980), Soldano (1983a, b),
Soldano (1993)
ering period (for most species this is from mid-June to the
end of July). During the second lowering period, the petals are almost always (much) smaller (especially in largelowered species), the number of glandular hairs tend to
increase while the number of stiff, eglandular hairs diminishes, the branching pattern changes (i.e. development of
side branches below the main inlorescence or of oblique
stems arising from the leaf rosette) and other characters
also are often less obvious.
It is inally recommended to collect only ‘typical’ examples of a population. In many Oenothera populations
– certainly those where two or more species grow in close
proximity – plants with intermediate characteristics are
not uncommon. These locally arisen, not-ixed hybrids
will, without doubt, fail to key out properly. In this perspective, it is an illusion to think that it is possible to assign a name to each individual Oenothera plant. However,
the identiication of more or less homogeneous populations is generally feasible provided one has some ield
experience. In fact, in a mixed population of Oenothera
species the dificulty mostly is to assess which plants species are the ‘pure’ ones and which are the hybrids.
Even if the above is taken into consideration, the identiication of Oenothera species often remains a dificult
task. The correct identiication of herbarium material
without accompanying ield notes is, even for experts, not
easy. The Belgian Oenothera collections of the herbarium
of the Botanic Garden Meise have been reviewed by Raymond Jean (see Jean 1975), Werner Dietrich and the irst
author of this paper. That the conclusions of Dietrich and
Rostański differ stems (mostly) from differences in taxonomical concepts. Jean, however, followed the narrow
‘European’ species concept and yet sometimes came to
conclusions which were divergent to those of Rostański.
For instance, plants which were named by Jean as “close
to O. ersteinensis” were inally attributed to O. delexa
and O. parvilora. A collection named as “O. hungarica”
(= O. depressa) is here ascribed to O. oakesiana although
both belong to different series. Without doubt, the most
striking was that nearly all O. parvilora collections – one
of the few more or less widely spread and fully established species in Belgium – turned out to be ascribable
to (the not even remotely related) O. delexa. This important correction was already picked up by Lambinon et al.
(2004).
The repeated confusion, even amongst experts, was
clearly illustrated with respect to a collection of Oenothera oakesiana from BR (Fig. 1); this collection was
Figure 1. Identiication of species in genus Oenothera has often
been dificult, even among experts. This collection of Oenothera
oakesiana from BR was originally identiied by A. Lawalrée as
O. biennis and later annotated by Dietrich, Jean and the irst
author. It inally bears four names of species that belong to three
different series.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
15
Figure 2. Flower morphology:
A, stigma lobes;
B, anthers;
C, petal;
D, sepal;
E, sepal tip;
F, hypanthium;
G, ovary;
H, bract.
originally identiied by A. Lawalrée as O. biennis and
later annotated by Dietrich, Jean and the irst author. It
inally bears four names of species that belong to three
different series!
Diagnostic characters
Although similar, the species in subsection Oenothera are
(with some experience) easily recognized in the ield. The
diagnostic characters, such as hairiness, colour of vegetative parts and size of loral parts (Fig. 2), are strikingly
constant. However, herbarium material is frequently more
dificult to identify because some characters are lost, especially in poorly prepared material (see before; also Jean
1975).
The following characters are particularly useful for
an accurate identiication (mostly based on Rostański &
Karlsson 2010):
Mode of growth. The stem is erect in most species but
ascending in a few. The tip of the growing inlorescence is
erect in most species but in some, e.g. Oenothera parvilora and related species from series Rugglesia, it is drooping (as beautifully illustrated in Renner 1956). This is an
important character that can easily be overlooked, e.g. in
late specimens where the axis has straightened. Also, in
some species of series Rugglesia, this character is less obvious, for instance in O. parvilora (Linder 1958).
Hair types (Fig. 3). There are trichomes of three main
types: 1) short, soft, crispate hairs on the vegetative part
of the stem and, in some species, on the ovary and fruit;
2) glandular hairs, i.e. patent, glistening, short and thick
(sometimes slightly clavate) trichomes, on rachis, hypanthium, sepals, ovary and fruit in many species; and 3)
long, stiff, more or less patent, straight or slightly bent
hairs on most parts of the plant. In some species the hairs
rise from a well-delimited, multicellular base which is often an intense red (presence of anthocyanin pigment). The
shape of the thickened hair base varies. In most cases it is
conical and about as high as wide; more rarely it is wide
and low, as in O. depressa, or distinctly elongated, cylindrical and ± bent, as in O. ersteinensis.
Leaf shape. The relative leaf width varies between the
species and, to some extent, within the species. Some are
always narrow-leaved; others, like O. biennis and O. cambrica are always broad-leaved. The leaves are usually lat,
but in some species they are crinkled and sometimes have
a slightly twisted apex (e.g. in O. depressa).
Leaf hairiness. The leaves are usually sparsely hairy,
but in some species, e.g. O. hirsutissima and in series
Devriesia (e.g. O. canovirens and O. depressa), they appear grey-green and velvety to the touch, due to a fairly
dense indumentum.
Leaf colour. The midrib of the leaves is whitish in
some species, ± red in others. However, the red colour is
sometimes not developed in species normally having red
midribs, e.g. in specimens growing in shade.
Rachis colour. The main axis (rachis) of the inlorescence can be either green throughout as in O. biennis or an
intense red towards the apex as in O. ersteinensis and O.
rubricaulis. This character is more stable within species
than the colour of leaves and sepals.
Sepal colour. The sepals are green in some species,
whereas in others, e.g. O. glazioviana, they have red
streaks or a red tinge or they may be entirely deep red
(e.g. O. rubricalyx). The red colour is, however, not always developed. There are also species with a genetically
conditioned variation in this character (e.g. O. oehlkersii).
Buds (Fig. 4). The sepal-tips can be entirely separated
or pressed to each other in their full length, or together in
their proximal part but diverging distally. The length of
the sepal tips and the length of the sepals (including the
tip, and measured in fully mature buds) differ between
some species.
Flower size. The length and width of the petals, as well
as the length/width ratio, are diagnostic. The length of the
hypanthium also differs much between species; it is not
correlated with petal size. Care should be taken to study
(left) Figure 3. Hair types: A, hair with bulbous
base; B, crispate hair; C, glandular hair; D,
bristle hair.
(right) Figure 4. Flower buds and sepal tips: A,
sepal tips appressed throughout; B, sepal tips
appressed at base, separated at apex; C, sepal
tips separated (V-shaped); D, sepal tips separated (U-shaped).
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
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Figure 5. Capsule and
morphological variation
of capsule apex:
A, capsule;
B, capsule valve;
C, teeth acute;
D, teeth truncate;
E, teeth emarginate.
the plant at its peak of lowering. Later in season the lowers become notably smaller, especially in the large-lowered taxa. In some species the lowers are sometimes or
even usually cleistogamous (e.g. O. depressa).
Position of stigma. The stigma is surrounded by the
anthers at anthesis in most species, but distinctly raised
above them in e.g. O. glazioviana and O. oehlkersii.
Hairiness of capsules. The density of trichomes of different kinds on the capsules is of major diagnostic value,
but care must be taken to compare capsules at the same
height, because the hairiness varies with position (i.e.,
between lower, older capsules and upper, younger ones).
Generally, glandular hairs are more numerous on the upper, younger capsules. There are species where glandular
hairs are absent on the lower (older) capsules, but present
on the upper (younger) ones.
Capsule valves (Fig 5). The tips of the capsule valves
are rounded, truncate or more or less emarginate. Their
outline is best seen in capsules that are about to open.
Degree of naturalization of Oenothera in Belgium
In Belgium Oenothera species are mostly found in environments which have been heavily disturbed by human
activity: ports and railway grounds, sand and gravel pits,
dredge dumping areas, sand and mineral depots, coal mining spoil heaps, ruderalized sand dunes, etc. The lora of
such environments is often rather ephemeral. Although
Oenothera often occurs in great numbers, this is, in the
most of its habitats, a temporary phenomenon. Moreover,
the knowledge of the genus in Belgium is virtually nonexistent and as such only the experience of the second
author can serve as a basis. Because of this, the correct
estimation of the extent of naturalization for many species
is very dificult. Perhaps only a few species are more or
less widespread and relatively common. This corresponds
with previous indings: Mihulka & Pyšek (2001) indicated that only few species are successful in terms of increasing abundance while the majority of the species remain
rare and local.
Probably the most widespread species for the moment
is O. delexa. It also appears to be the most expansive of
all species. Locally, for example in the coastal sand dunes,
O. glazioviana is also well established as is O. fallax. The
latter is spreading vigorously, for example, in the ports of
Ghent and Antwerp. This species has also become one of
the most frequently found species in the west of Germany
(e.g. Wittig and Tokhtar 2003). At least in the area around
Ghent, O. rubricaulis is well established and in expansion. This species was already found in great numbers in
1985 (e.g. E. Robbrecht 2929 in BR) and it is now one
of the typical species for this area. The same holds true
for O. ersteinensis around Antwerp and in the Kempen.
O. biennis is a perplexing case: it is, without doubt, the
species with the longest history in Belgium but currently
appears to be, to the greater extent, replaced by related
species. Although, this species is still regularly collected,
this is much less so in comparison with the other species
cited before. Locally, in the second half of the 19th century, O. angustissima (syn.: O. muricata auct. belg. non
L.) was more or less established, for example, along a
railway track in the vicinity of Sint-Truiden (at least between 1860 and 1884). All the remaining Oenothera taxa
are either not well known enough, or are only very locally
more or less established, or (without doubt in the most
cases) strictly ephemeral.
Some of the aforementioned species (e.g. O. glazioviana) are naturalized in great number in natural habitats,
for instance in coastal dunes. In the sense of Richardson
et al. (2000) they can probably be considered as invasive
species. However, there are no indications that they replace indigenous vegetation (‘transformers’). As shown
elsewhere, species from other (sub-) sections that were
introduced relatively recently proved to exhibit a much
stronger invasive behaviour (e.g. Frean et al. 1997). However, Tokhtar et al. (2011) demonstrated that (notho-)
species from subsection Oenothera can also act as invasive species, especially if one putative parent is of North
American and the other of European origin.
Finally, an additional dificulty in assessing the degree of naturalization of Oenothera is the morphological plasticity of many populations. It often happens that
Oenothera populations change strongly during the course
of years: small but apparently more or less stabilized
populations of rare species often disappear in a hybridization swarm of commoner species. For example, a nice
population of O. victorinii very gradually disappeared
from rough ground in the vicinity of Roeselare. The few
remaining plants became less and less typical, doubtlessly
as a result of introgression with the co-existing populations of O. glazioviana and/or O. delexa, eventually disappearing altogether. The same holds true for a population
of O. hirsutissima found on a sand storage area in the canal zone of Bruges: between 2004 and 2007 a few typical plants were collected whilst from 2008 onwards only
non-deined hybrids appeared to be present. Analogously,
a population of O. wratislaviensis in the Antwerp port
area, discovered in 2007, is replaced now by plants that
exhibit characters of this species, as well as of the widely
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
17
spread O. ersteinensis. This is a well-known phenomenon
in Oenothera: Renner (1950) referred to it as “Der Kampf
zwischen der Arten”.
Geographical origin of the genus Oenothera
The origin of the genus Oenothera is quite uncertain. According to Wagner et al. (2007) its centre of diversity is in
southwestern North America and it is now widely distributed in temperate to subtropical areas in North and South
America with a few species in Central America as well. A
number of species have become naturalized nearly worldwide (Frean et al. 1997), most likely in post-Columbian
times. According to the European school, on the contrary,
several species are endemic to and arose in Europe. This
applies not only for the European mutants and hybrids
(that are unknown in America) but probably also for some
‘pure’ species such as O. biennis. Most likely, Oenothera
subsection Oenothera underwent its primary differentiation in North America but has a secondary centre of evolution in Europe (Rostański & Karlsson 2010). Quite a few
originated by hybridization in Europe and became stabilized due to the extraordinary genetic mechanism mentioned above, and have subsequently become widespread.
Species like O. biennis and O. rubricaulis have been present for a long time in Europe. Since exactly matching biotypes are not known in the wild in North America, early
immigration and speciation has been proposed (Rostański
1968, 2004), while Dietrich et al. (1997) suggest a hybridogenous origin in Europe from North American taxa
introduced in historic time.
Yet another possibility (comm. R. Jean, August 2014)
is that Oenothera is, indeed, of North American origin
but that species of subsection Oenothera already entered
the Eurasian Continent, via the Bering land bridge, at the
end of the last glacial period. This could explain why, genetically, ‘European’ species of Oenothera differ to such
an extent from ‘American’ congeners. Species from subsections Raimannia (originating in North America) and
Munzia (originating in South America), on the contrary,
were doubtlessly introduced in post-Columbian times in
Europe, have a much more limited distribution (mostly
in coastal areas, without having penetrated deeper in the
Continent) and are identical to plants found in their respective areas of origin.
Identiication key
Despite being stable, and as such occurring in the absence
of their putative parental species, Oenothera paradoxa
and O. wratislaviensis have not been included in the
identiication key. Both are rare and relatively atypical
taxa that are dificult to incorporate into the key. They
are briely described with their assumed parental species
(O. canovirens, O. depressa and O. subterminalis). Both
are hybrids between species of the series Devriesia and
Rugglesia. Their capsules display an indumentum that is
more or less intermediate, with an indeinite mixture of
short glandular and stiff, long and eglandular hairs. From
the series Devriesia and Oenothera, the unixed hybrid O.
delexa × O. depressa has also been recorded on several
occasions. As previously mentioned, it is virtually impossible to identify sporadically occurring hybrids. Plants
which either cannot or only with dificulty be keyed-out
belong in all probability to hybrid swarms.
The current study follows the so-called European
school and thus accepts a narrow species concept. Nevertheless, it seemed also useful to always refer to the corresponding name in the taxonomic concept as deined by
the so-called American school. For this reason, for every
taxon of the section Oenothera subsection Oenothera the
alternative name under the latter is indicated between
square brackets.
1
Petals yellow when fresh (rarely pale yellow) (but in
some species drying pinkish, orangish or reddish!).
Capsule cylindrical or fusiform, very rarely clavate and
abruptly constricted to base (only in O. fruticosa) .... 3
2
Capsule indehiscent. Seeds usually 34. Flowers
strongly zygomorphic ........... Oenothera lindheimeri
Capsule dehiscent. Seeds usually numerous. Flowers
actinomorphic .............................................. O. rosea
3
Flowers opening at sunrise. Capsule clavate, abruptly
constricted or cuneate to the base (apparently pedicel
late), narrowly winged .... O. fruticosa subsp. glauca
Flowers opening near sunset. Capsule cylindrical or
fusiform, without wings, not constricted to base ...... 4
4
Classiication of Oenothera in Belgium
The overview of all the taxa collected in Belgium is divided over the different sections, subsections and series
(Table 2). Taxa collected exclusively before 1950 are preceded by an asterisk (*). In the rest of this paper (except in
the appendix), no further reference is made to these taxa
(see earlier; also: Verloove et al. 2014).
This classiication is in accordance with the recent
revision by Wagner et al. (2007) with the exception of
Oenothera subsection Oenothera. This implies that the
genus Gaura L. is included in Oenothera.
Petals white or pink when fresh. Capsule clavate, ab
ruptly constricted or cuneate to the base (apparently
pedicellate) .............................................................. 2
Seeds ellipsoid to globose, not prismatic and angled.
Capsule fusiform, enlarged towards apex (24 mm
wide at base). Petals yellow, often with a red spot at
base (turning pinkish on drying) .................. O. stricta
Seeds prismatic, distinctly angled. Capsule cylindrical,
narrowed towards apex (68 mm wide at base). Petals
yellow, unspotted (exceptionally fading whitish or pink
ish on drying) ........................................................... 5
5
Stigma distinctly elevated above anthers at anthesis.
Petals always large, 3055 mm long ........................ 6
Stigma surrounded by anthers at anthesis or only
slightly exceeding anthers. Petals small or large ..... 9
6
Sepals entirely green or greenish yellow. Stem with
out hairs with red bulbous base [O. biennis × glazioviana?] ..................................................... O. oehlkersii
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
18
7
8
9
10
11
12
13
14
Sepals redstriped, suffused red or uniformly deep red.
Stem with hairs with distinct red bulbous base ........ 7
Leaf blade nearly plane, subentire at margin, up to
25 mm wide, the upper surface grayish green (in
dumentum of dense, short hairs). Hypanthium (loral
tube) without or with scattered glandular hairs. Ovary
and young capsule densely white hairy, without or with
few glandular hairs [O. elata] ........... O. hirsutissima
Leaf blade distinctly wavy, irregularly dentate at mar
gin, up to 40 mm wide, the upper surface green and
nearly glabrous. Hypanthium densely glandular hairy.
Ovary and young capsule predominantly glandular
hairy, without or with few long, white and eglandular
hairs [O. glazioviana] ............................................... 8
Sepals uniformly deep red ................... O. rubricalyx
Sepals redstriped or suffused red ..... O. glazioviana
Ovary and young capsules densely covered with long
white hairs, without or with few glandular hairs. Cauline
leaves usually lanceolate, upper surface (at least when
young) grayish green with indumentum of dense, short
hairs, velvety to the touch. Capsule teeth emarginate
or truncate at apex [O. villosa] ............................... 10
Ovary and young capsules not densely white hairy,
with a mixture of (predominantly) glandular and (scat
tered) patent eglandular hairs. Cauline leaves elliptic
to lanceolate, glabrous to slightly hairy. Capsule teeth
rounded or cuneate at apex, rarely slightly emarginate
................................................................................ 12
Stem with numerous hairs with red bulbous base.
Bracts and upper cauline leaves with distinctly wavy
margins and twisted at apex. Flowers often cleisto
gamous .................................................. O. depressa
Stem usually without or (rarely) with sparse hairs with
red bulbous base. Bracts and upper cauline leaves
with lat margins. Flowers chasmogamous, fully open
ing ........................................................................... 11
Capsule teeth truncate at apex .................. O. villosa
Capsule teeth emarginate at apex ...... O. canovirens
Inlorescence axis slightly curved before lowering
(straightening with time). Cauline leaves narrowly lan
ceolate, always with red midrib. Sepal tips not adher
ing at base, with U or V shaped sinus1. Petals small, at
most 20 mm long (series Rugglesia) ..................... 13
Inlorescence axis straight from the beginning. Cauline
leaves elliptic to lanceolate, with white or red midrib.
Sepal tips in bud adherent at base (series Oenothera)
................................................................................ 15
Sepal tips in bud 23 mm long, with U shaped sinus.
Petals 612 mm long [O. parvilora] ...... O. parvilora
Sepal tips in bud 29 mm long, with V shaped sinus.
Petals 1020 mm long ............................................ 14
Stem dark red, at least in lower half. Sepal tips in bud
49 mm long [O. parvilora] ............ O. subterminalis
Stem green or slightly suffused red in lower half. Sepal
tips in bud at most 3 mm long [O. biennis × oakesiana] ............................................................. O. issleri
The shape of the sepal tips in bud is a useful diagnostic feature, especially to distinguish between series Oenothera and Rugglesia. However,
this character should preferably be assessed on fresh material. See ig. 4.
1
15 Inlorescence axis and upper half of stem without hairs
with red bulbous base. Buds, sepals and inlorescence
axis uniformly green ............................................... 16
Inlorescence axis and upper half of stem with numer
ous hairs with red bulbous base. Buds, sepals and in
lorescence axis either green, red, or green suffused
with red .................................................................. 20
16 Inlorescence axis nearly glabrous (at most with very
scattered, long eglandular hairs) [O. biennis] .............
....................................................................... O. nuda
Inlorescence axis densely hairy, in part with short,
glandular hairs ....................................................... 17
17 Petals 2030 mm long [O. biennis] ......................... 18
Petals 1020 mm long ............................................ 19
18 Cauline leaves lanceolate. Petals about as long as
wide. Lowermost (oldest) capsules without glandular
hairs ............................ O. cambrica var. impunctata
Cauline leaves wider, elliptic to ellipticlanceolate. Pet
als obviously wider than long. All capsules glandular
hairy .......................................................... O. biennis
19 Petals 912 mm long. Hypanthium up to 30 mm long.
Capsule up to 40 mm long. Sepal tips 34 mm long [O.
parvilora] .................................................. O. delexa
Petals 1020 mm long. Hypanthium up to 40 mm long.
Capsule 3050 mm long. Sepal tips 57 mm long [O.
biennis] ................................................... O. victorinii
20 Cauline leaves with wavy margins. Sepals always red
striped. Petals 2530 mm long and slightly wider [O.
biennis × glazioviana] .................................. O. fallax
Cauline leaves plane. Sepals green or suffused with
red (redstriped in O. ersteinensis only). Petals often
smaller, 1028 mm long ......................................... 21
21 Inlorescence axis at the beginning of lowering red
(stem red or green) ................................................ 22
Inlorescence axis at the beginning of lowering green
(stem red or green) [O. biennis] ............................. 24
22 Pustulate hair base cylindrical, much longer than wide
and often slightly curved downwards. Sepals mostly
redstriped (rarely green). Stem usually entirely red [O.
biennis] ............................................. O. ersteinensis
Pustulate hair base conical, ca. as long as wide. Se
pals green. Stem green, at least in lower half ........ 23
23 Hypanthium 1525 mm long. Petals 1020 mm long
and 918 mm wide [O. biennis] ........... O. rubricaulis
Hypanthium 2535 mm long. Petals 2028 mm long
and about as wide [O. biennis?] ... O. rubricauloides
24 Petals 612 mm long ............................. O. royfraseri
Petals 1530 mm long ............................................ 25
25 Cauline leaves lanceolate, with red midrib. Petals
1525 mm long. Hypanthium 3040 mm long. Inlor
escence lax, more or less pyramidal. Stem nearly al
ways red, at least in lower half .......... O. pycnocarpa
Cauline leaves wider, ellipticlanceolate, with white or
red midrib. Petals 2030 mm long. Hypanthium 2535
mm long. Inlorescence dense, oblong. Stem usually
green throughout ........... O. cambrica var. cambrica
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
19
Table 2. Classiication of the Oenothera taxa collected in Belgium. Taxa collected exclusively before 1950 are preceded by an asterisk.
Based on Wagner et al. (2007). For subsection Oenothera the names in the second column are those of the socalled American school.
Section Gaura (L.) W.L. Wagner & Hoch
* Oenothera curtilora W.L. Wagner & Hoch (syn.: Gaura parvilora Douglas ex Lehm.)
Oenothera lindheimeri (Engelmann & A. Gray) W.L. Wagner & Hoch (syn.: Gaura lindheimeri Engelmann & A. Gray)
Section Hartmannia (Spach) W.L. Wagner & Hoch
Oenothera rosea L’Hérit. ex Ait.
* Oenothera speciosa Nuttall
Section Kneifia (Spach) Straley
Oenothera fruticosa L. subsp. glauca (Michaux) Straley
* Oenothera perennis L.
Section Oenothera
Subsection Munzia Dietrich
Series Allochroa (Fischer & Meyer) Dietrich
* Oenothera afinis Cambess. in St.Hilaire
* Oenothera indecora Cambess. in St.Hilaire
Oenothera stricta Ledeb. ex Link
Subsection Oenothera
Series Devriesia Rostański
Oenothera canovirens Steele
Oenothera depressa E. Greene
Oenothera paradoxa Hudziok
Oenothera villosa Thunb.
Oenothera wratislaviensis Rostański ex Rostański
O. villosa
O. villosa
O. biennis
O. villosa
?
Series Linderia Rostański
Oenothera hirsutissima (A. Gray ex S. Watson) de Vries
O. elata
Series Oenothera
Oenothera biennis L.
Oenothera cambrica Rostański
Oenothera delexa Gates
Oenothera ersteinensis Linder & Jean
Oenothera fallax Renner emend. Rostański
Oenothera glazioviana Micheli
Oenothera nuda Renner ex Rostański
O. biennis
O. biennis
O. parvilora
O. biennis
O. biennis × O. glazioviana
O. glazioviana
O. biennis
Oenothera oehlkersii Kappus ex Rostański
Oenothera pycnocarpa Atkinson & Bartlett in Bartlett
Oenothera royfraseri Gates
Oenothera rubricalyx Gates
Oenothera rubricaulis Klebahn
Oenothera rubricauloides Rostański
Oenothera victorinii Gates & Catches.
O. biennis × O. glazioviana?
O. biennis
O. biennis
O. glazioviana
O. biennis
O. biennis?
O. biennis
Series Rugglesia Rostański
* Oenothera angustissima Gates
* Oenothera cruciata Nuttall ex G. Don
Oenothera issleri Renner ex Rostański
* Oenothera oakesiana (A. Gray) J.W. Robbins ex S. Watson & Coulter
Oenothera parvilora L.
Oenothera subterminalis Gates
O. parvilora
O. parvilora
O. biennis × O. oakesiana
O. oakesiana
O. parvilora
O. parvilora
Subsection Raimannia (Rose) Dietrich
* Oenothera laciniata Hill
Notes on the species recorded in Belgium after 1950
For each of the species included in the key, additional
information is provided with respect to its identiication
and differentiation from related or similar species, current and/or historical distribution in Belgium, degree of
naturalization, etc. The most important synonyms have
also been added. This is useful since several species were
initially recognized under other names in Europe. And,
inally, the probable area of origin of each species is also
provided (see before).
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
20
Figure 6. Oenothera biennis. Brugge, E-side of
Boudewijnkanaal, sand storage area, August 2007.
Characteristic features are the entirely green stem
and sepals and the medium-sized petals.
After having compared specimens from America and
Europe, Rostański (1985) came to the conclusion that ten
Oenothera species described from Europe were in fact already previously described from North America. However, part of this synonymy was rejected subsequently (see
for instance under O. cambrica and O. ersteinensis). It
may well be that none of the species described from Europe exactly match and that they are best referred to using
their ‘European’ names (comm. R. Jean, August 2014).
• Oenothera biennis L., Sp. Pl. 1: 346. 1753.
Figure 6.
Probable area of origin: Eurasia?
Oenothera biennis doubtlessly was the irst introduced
species of the genus in Belgium. Several sources conirm
its presence at the end of the 18th century when the species
was already locally abundant according to Crépin (1860):
“(…) dès 1792, Roucel la signalait déjà comme abondante
aux environs de Bruxelles, Gand et Termonde.” Yet, in the
last few decades, it has been less often observed and it is
even uncertain if it can really be considered as naturalized
at present. From the recent distribution map of Flanders
(Verloove 2006b) it does appear to be slightly expanding
but it is doubtful if this relects reality.
Oenothera biennis is easily recognizable by the medium sized petals that are wider than long, the total absence
of hairs with red bulbous base, the uniformly green-colored sepals and lower buds, etc. Oenothera delexa and O.
victorinii are more or less similar but have clearly smaller
petals. Most similar probably is O. cambrica (especially
var. impunctata). The latter has slightly narrower leaves,
petals that are as long as wide and at least the lower capsules are eglandular.
At the coal mining spoil heap of Grande Machine à
Feu near Dour existed, for years, a population with pale
yellow petals (last conirmed in 2007). Such plants belong to f. sulphurea de Vries. At the same site, it was
also observed that this taxon formed hybrids with the coexisting population of Oenothera fallax. At various locations where O. biennis and O. delexa occur sympatrically
occasional hybrids between these species have also been
observed.
• Oenothera cambrica Rostański, Fragm. Florist. Geobot.
23(3-4): 285. 1977.
Figure 7.
Probable area of origin: Europe?
Oenothera cambrica is a rare but possibly overlooked species (see before for the differentiation with the related O.
biennis). For a long time it was known only from the British Isles (from where it was initially described; Rostański
1977) but more recently it has also been observed in the
northwest of France (comm. D. Mercier). The typical variety (var. cambrica) has a red punctuate stem whilst var.
impunctata Rostański has a uniformly green stem. In the
last few years both varieties have been collected, in Roeselare (2001) and in Zwijnaarde (2007, 2014).
For some taxa which were described in Europe, it inally turned out that these were documented before in
America. Rostański (1985) cited ten such cases, including
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
21
= O. velutinifolia Hudziok, Verh. Bot. Vereins Prov.
Brandenburg 105: 103. 1968.
Probable area of origin: North America.
Oenothera canovirens is a very rare species in Belgium.
It was last collected in 1959 from a sand raised site in
Antwerpen-Linkeroever. Along with O. depressa, it belongs to the series Devriesia. Typically, these species
have dense, stiff white hairs (with or without a few, barely
visible glandular hairs) on the young fruits and leaves are
densely hairy and velvety to the touch. Moreover, the upper leaves are often conspicuously falcate. Confusingly,
they easily form hybrids with species of the series Oenothera; these hybrids display a much less typical assemblage of characters. The putative hybrid of O. canovirens
and O. subterminalis (= O. wratislaviensis), a taxon that is
also found in areas where one or both parental species are
absent, has recently been collected on several occasions,
including, for example, the port of Antwerp, in the Baai
van Heist and at an abandoned coal mining site in Hensies. This hybridogenous species differs from O. canovirens by the clearly red striped lower buds, the red midrib
of the leaves and by petals being broader than long. O.
canovirens is best differentiated from O. depressa by the
lattened or slightly V-shaped bracts and leaves (without
wrinkled margins), the chasmogamous lowers and by the
complete or almost complete absence of hairs with a red
bulbous base.
Figure 7. Oenothera cambrica (var. cambrica). Zwijnaarde
(Gent), roadside, August 2014. This taxon resembles O. biennis
but the stem is clearly red-punctate.
Oenothera cambrica. From this, the correct name for O.
cambrica appeared to be O. novae-scotiae Gates. However, according to Dietrich (1991) the type of this species
is certainly not identical with O. cambrica, nor with O.
biennis and probably more closely related to O. parvilora. Whichever taxonomical concept is used, the name O.
novae-scotiae cannot be used for O. cambrica.
• Oenothera canovirens Steele, Contr. U.S. Natl. Herb.
13(10): 365-366. 1911.
= O. renneri H. Scholz, Wiss. Z. Pädagog. Hochschule
Potsdam, Math.-Naturwiss. Reihe 2: 206. 1956.
• Oenothera delexa Gates, Philos. Trans., Ser. B 226:
332. 1936.
= O. lipsiensis Rostański & Gutte, Ber. Arbeitsgem.
Sächs. Bot. n.s. 9: 69. 1971.
Figure 8.
Probable area of origin: North America.
Oenothera delexa is, at present, perhaps the commonest species in Belgium. The irst record dates from 1915
(Verloove 2006a) but it has only been spreading vigourously in recent decades (see current distribution map for
Flanders in Verloove 2006c). This species was for a long
time confused with O. parvilora, a species which does
not resemble it at all, except that they both have small
petals. O. delexa resembles, for the most, O. biennis and
O. victorinii (all with green sepals and lower buds, absence of hairs with red bulbous base, etc.). It differs from
O. biennis by its much smaller petals that are not broader
than long. O. victorinii, in turn, has petals, hypanthium
and capsules that are slightly larger.
Gutte & Rostański (1971) described Oenothera lipsiensis from Germany as a new species. Later, it became
evident that Gates had already earlier described the same
species on the basis of American plant material (as O.
delexa) (Rostański 1985), the latter binomial evidently
having priority.
Although this species is currently, without doubt, the
most commonly found species in Belgium, it is elsewhere
in Europe much less common. Rostański (2006) concluded that it was more or less widespread in Germany,
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
22
leaves that are velvety to the touch make it a striking and
easy to recognize plant. In 2007, it was observed at an
abandoned coal mining site in Hensies. An earlier Belgian
record for Brussels (Jean 1975) now appears to be referable to O. oakesiana.
Just as with Oenothera canovirens, O. depressa easily
forms hybrids with species from the series Oenothera and
Rugglesia. At a coal mining spoil heap in Beringen-Mijn
Figure 8. Oenothera delexa. Zwijnaarde (Gent), roadside, August 2014. This species is recognized by its small-sized petals
and the absence of hairs with a red bulbous base.
whereas in Sweden its occurrence was considered as very
rare. According to ield observations carried out by the
second author, it is also to be found in the Netherlands and
France, close to the Belgian frontiers.
At a sand storage area in the canal zone of Bruges, a
hybrid with Oenothera fallax has also been found growing among the parental species. Likewise, in the port of
Bruges and at several other locations, putative (unstabilized) hybrids of this species with O. biennis have also
been collected.
• Oenothera depressa E. Greene, Pittonia 2: 216. 1891.
= O. hungarica (Borbás) Borbás, Magyar Bot. Lapok 2:
246. 1903.
= O. salicifolia Desf. ex G. Don, Gen. Hist. 2: 685. 1832,
nom. inval.
Figure 9.
Probable area of origin: North America.
This species belongs, just as Oenothera canovirens, to
the characteristic series Devriesia (see earlier). It differs
mostly from the latter by the striking wavy edges of the
leaves and bracts, the presence of numerous hairs with a
red-bulbous base and the usually cleistogamous lowers.
This combination of characteristics as well as the numerous white, eglandular hairs on the young fruits and the
Figure 9. Oenothera depressa. Hensies (Les Sartis), abandoned
coal mining site, July 2007. This species is softly grey-hairy
throughout, its leaves have wavy margins and lowers are often
cleistogamous.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
23
a reasonably large population of O. paradoxa is found, at
least since 2001. This is a probable (stable) hybrid of O.
depressa and O. subterminalis (or O. parvilora). It has
striking lower buds that are only suffused with red for the
uppermost third but for the remainder it is atypical and
dificult to insert in the identiication key. It is moreover
unclear if (at least part of) the current concept of O. paradoxa is in agreement with the original concept of this species. [Compare for example Jäger & Werner (2005): “Stg.
und Bstandachse nicht getupft” with Hudziok’s original
description (Hudziok 1968): “Caulis … rubropunctulatus
…”] In addition to O. paradoxa, a probable hybrid of O.
delexa and O. depressa has recently been collected in
Zeebrugge and Zwijnaarde.
In large parts of Central and Eastern Europe, Oenothera
depressa is one of the most common species (e.g. Jehlík
& Rostański 1979, Rostański et al. 1994, Zlatković et al.
1998, Rostański 2006). It is found especially in dried out
riverbeds (e.g. Deschâtres 1954; sub O. strigosa) and similar habitats that are, for the most part, not found in Belgium. It is possible, that it has been overlooked along the
River Maas, where it could ind a suitable habitat. However, O. depressa obviously becomes much rarer towards
Western Europe (compare also with Rostański 1982).
This species is often referred to as Oenothera salicifolia Desf. ex G. Don (1832). This is an invalid name
since the epithet was already given to a different species
by Lehmann (1824). The oldest valid name for this taxon
appears to be O. depressa.
• Oenothera ersteinensis R. Linder & R. Jean, Bull. Soc.
Bot. France 116: 523. 1970.
?= O. perangusta Gates, Canad. Field-Naturalist 64: 142.
1950.
Figure 10.
Probable area of origin: North America.
Oenothera ersteinensis is one of the relatively easy-toidentify species in Belgium. It is unmistakable due to
its striking stem hairs: the thickened red hair base is not
conical-shaped but cylindrical (much longer than broad)
and often slightly curved. It is further characterized by its
(usually) red striped sepals and lower buds, the relatively
small petals and the (mostly) deep-red stem. Since 1971,
it has, on two occasions, been collected in the vicinity
of Liège (Ougrée and Chenée). However, it is most well
known in the port of Antwerp where, since 2001, it has
been found on a regular basis. At present (2014) it has
become one of the most widely spread species in the port
area. Also in the Kempen it is not rare at all (comm. R.
Barendse).
Linder & Jean (1969) described Oenothera ersteinensis as a new species from northeastern France. Rostański
Figure 10. Oenothera ersteinensis. Balen
(Wezel), August 2010 (lowers and stem)
and Grobbendonk, Albertkanaal, July
2013 (hairs). This is a characteristic species with a deep-red inlorescence axis,
red striped sepals, medium-sized petals
and hairs with a red bulbous base that is
longer than wide. In the similar O. rubricaulis, sepals are green and the hair base
is conical.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
24
(1985) commented that the species had already been earlier described by Gates on the basis of American plant material (as O. perangusta). Noteworthy, is that Dietrich et
al. (1997) considered O. perangusta as a synonym for O.
oakesiana, whilst O. ersteinensis was considered as belonging to O. biennis. Both are indeed probably distinct,
the Belgian plants belonging to O. ersteinensis, not to O.
perangusta. Also, in Rostański et al. (2010) both were
eventually accepted as separate species.
• Oenothera fallax Renner emend. Rostański, Fragm.
Flor. Geobot. 11: 507. 1965.
Figure 11.
Probable area of origin: Europe.
This stable hybrid of Oenothera biennis (♂) and O.
glazioviana (♀) parentage is one of the strongest spreading species in Belgium (see also Wittig & Tokhtar 2003).
It was already recorded at the end of the 19th century (valley of river Bocq) but its genuine naturalization probably
started in the irst half of the 20th century when it began to
occur in coastal dunes where O. glazioviana and O. biennis were already naturalized. It is now locally common,
especially at sites in the port of Antwerp and Ghent but
also in numerous other localities throughout the country.
After O. biennis, O. fallax is the most numerous species in
the Rhine valley in Germany (Wittig et al. 1999).
Oenothera fallax is readily recognizable by its showy
red-striped lower buds and its medium sized petals and
cannot be confused with any other species. It is mostly
found in populations with both parental species but is also
found without them. Moreover, this hybridogenous species is also cultivated in gardens.
At a sand depot in the canal zone of Bruges, a (nonstabilized) hybrid has been found with Oenothera delexa.
• Oenothera fruticosa L., Sp. Pl. 1: 346–347. 1753.
subsp. glauca (Michx.) Straley, Ann. Missouri Bot. Gard.
64(3): 403. 1977 [1978].
= O. tetragona Roth, nom. inval.
Probable area of origin: North America.
This ornamental was recorded only once, in 1998, near a
waste recycling factory in Ghent. According to Dietrich
(1999) this is one of the most frequently cultivated species but this does not seem to be the case in Belgium. It is
a very attractive garden plant with its club shaped fruits,
day lowering and for being a perennial (not biennial, the
most typical life cycle found in the other taxa occurring
in Belgium). For a long time, O. fruticosa was accommodated in a distinct genus (Kneifia Spach).
This taxon has long been referred to as Oenothera tetragona, an invalid name. The correct name, according to
Wagner (2014), is O. fruticosa subsp. glauca.
Figure 11. Oenothera fallax. Grobbendonk, Albertkanaal, July
2013. Typical features of this species are the red striped sepals,
the medium-sized petals and the presence of numerous hairs
with red bulbous base on the stems.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
25
Figure 12. Oenothera glazioviana. Kortrijk, August
2014. Very characteristic species with large petals,
sepals that are suffused with red and stems that are
red punctate. Compared with O. hirsutissima, leaves
are wider, less hairy and with distinctly wavy margins.
• Oenothera glazioviana Micheli, Fl. Bras. 13(2): 178179. 1875.
= O. erythrosepala Borbás, Magyar Bot. Lapok 2: 245.
1903.
Figure 12.
Probable area of origin: Europe.
Oenothera glazioviana is one of the few genuinely established species in Belgium. Initially published as O.
lamarckiana (misapplied), subsequently as O. erythrosepala (synonym), this species was already well-established
in the irst half of the 20th century according to Goffart
(1945), especially in the coastal dunes: “se propage au littoral où il abonde dans les dunes (…)”. In 1920, it was
already found in the vicinity of the coastal towns of Koksijde and Nieuwpoort. It appears that it is still spreading,
as shown on the distribution map for Flanders in Verloove
(2006d), with a concentration of its expansion clearly located on the Belgian coast. In Belgium it can only be confused with O. hirsutissima (see earlier), O. rubricalyx (see
further) and O. oehlkersii, all with large petals and with
stigma branches elevated high above the anthers. From
the latter it is best differentiated by the presence of hairs
with red bulbous base on the stems and the sepals that are
suffused with red.
Oenothera glazioviana probably arose in Europe as a
result of hybridization or mutation. It is unknown in North
America, at least as a wild species.
• Oenothera hirsutissima (A. Gray ex S. Watson) de Vries,
Mutationstheorie 1: 327. 1901.
≡ O. elata Kunth subsp. hirsutissima (A. Gray ex S. Watson) W. Dietrich.
Figure 13.
Probable area of origin: North America, Mexico.
Oenothera hirsutissima is a rare and ephemeral species
in Belgium. Up to the present it has always been found
on sand: in 1961 in the sand dunes near to Nieuwpoort
(Belgian coast) and, more recently, at a sand storage area
alongside the canal zone of Bruges (between 2004 and
2007) and at a sand raised site located at the Kluizendok
in the Ghent port area (2007). This species most resembles
O. glazioviana (large petals, stigma branches elevated
high above the anthers, presence of hairs with red bulbous
base, etc.). It is nevertheless differentiated by the typical
appressed whitish hairs (absence of glandular hairs) on
young leaves and capsules and the narrower, nearly entire
leaves that are velvety to the touch. At the site in Bruges
it has been observed that O. hirsutissima gradually dimin-
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
26
Figure 13. Oenothera hirsutissima. Brugge, E-side of Boudewijnkanaal, sand storage area, July 2007. This species is much
reminiscent of O. glazioviana but it has softly hairy upper leaf
surfaces. Its petals often turn orangish.
ished, apparently as a result of introgression or hybridization with the accompanying species (especially O. biennis, O. delexa, O. fallax and O. glazioviana).
Oenothera hirsutissima is possibly overlooked in Europe. This species is also cultivated but is probably not
fully hardy in Western Europe (Dietrich 1999). Also in
Germany, it was recently found in the wild (Rostański &
Meierott 2006).
The systematic position of O. elata, O. hirsutissima
and O. hookeri Torr. & A. Gray, the latter two considered
as subspecies of the former by Dietrich et al. (1997), is
very unclear. Rostański (1985) assigned O. elata s.str. to
the series Devriesia, whilst he included both other taxa in
the series Linderia. From this complex O. hirsutissima is
the most widely spread taxon (Dietrich et al. 1997).
• Oenothera issleri Renner ex Rostański, Fragm. Florist.
Geobot. 11: 514. 1965.
Figure 14.
Probable area of origin: Europe.
Oenothera issleri is a very rare and ephemeral species in
Belgium and has been collected only twice so far. It was
seen along a railway track in Jambes (Namur) in 1955
and more recently, in 2014, on bare sandy ground in an
Figure 14. Oenothera issleri. Beveren (Verrebroek), bare sandy
area, August 2014. Typical features
of this species are its oblique stem
and lower buds with sepal tips with
V-shaped sinus.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
27
Figure 15. Oenothera lindheimeri. Vlissingen (Netherlands), dry
grassland, September 2009. Unmistakable by its zygomorphic lowers and previously accommodated in a separate genus (Gaura).
industrial area in Verrebroek (Beveren). It is a stable hybrid of O. biennis and O. oakesiana parentage (series Oenothera and Rugglesia respectively). Oenothera issleri
is also known from a single population on a coal mining
spoil heap in northwestern France, relatively close to the
Belgian frontiers (Jean & Delay 2008).
Figure 16. Oenothera nuda. Blaton, sand pit, July 2007. Fairly
characteristic in being virtually glabrous throughout.
• Oenothera lindheimeri (Engelm. & A. Gray) W.L. Wagner & Hoch, Syst. Bot. Monogr. 83: 213. 2007.
≡ Gaura lindheimeri Engelm. & A. Gray, Boston J. Nat.
Hist. 5: 217. 1845.
Figure 15.
Probable area of origin: North America.
Oenothera lindheimeri is at present often cultivated as
an ornamental in Belgium (‘prachtkaars’). It is unmistakable with its numerous white (or rose) lowers borne on
an inlorescence, which is long and drawn out. In recent
years it has, now and then, been found on waste ground
or levelled soil, often in places where garden waste has
been disposed. In Vlissingen (the Netherlands) it has been
recorded in dry grassland in the port area, close to the
railway station where it persists in small number since its
initial discovery in 2009.
This species was until recently included in the genus
Gaura but molecular phylogenetic analysis has demonstrated that this genus is better included in Oenothera
(Wagner et al. 2007).
• Oenothera nuda Renner ex Rostański, Nordic J. Bot.
27(2): 138 (-139). 2009.
Figure 16.
Probable area of origin: Europe.
Oenothera nuda is distinctive with its stem, loral tube,
capsule and inlorescence that are virtually glabrous. For
the rest, it is particularly reminiscent of O. biennis. This
species has been found earlier on a few occasions in Wallonia (Houx and Huy). More recently, O. nuda has also
been collected from a sand pit near to Blaton and at an
abandoned coal mining site in Hensies.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
28
• Oenothera parvilora L., Syst. Nat. (ed. 10) 2: 998. 1759.
Probable area of origin: North America.
In Belgium, Oenothera parvilora is rare and declining.
Nearly all references cited in the literature are in fact referable to O. delexa (see earlier). O. parvilora belongs
(along with O. subterminalis) to the series Rugglesia.
This group is characterized by a slight to clearly curved
inlorescence just before lowering and the sepal points
which are separated from the base. Unfortunately, both
characteristics are only really clear in fresh material.
Moreover, this species is also differentiated from O. delexa by the frequent presence of hairs with a red bulbous
base. O. parvilora has, since 1884, been collected on 15
occasions, especially around Brussels. The majority of the
inds date from the 1950s and 1960s. Yet more recently, it
has also been collected in Halle (near to Brussels) (1978),
Heist-aan-Zee (Belgian coast) (2000) and Jemeppe-sur
Meuse (1984).
Figure 17. Oenothera oehlkersii. Brugge, rough ground, August
2014. This species is much reminiscent of O. glazioviana but it
has green sepals and stems.
• Oenothera oehlkersii Kappus ex Rostański, Feddes
Repert. 96 : 9. 1985.
Figure 17.
Probable area of origin: Europe.
Oenothera oehlkersii is a striking hybridogenous species which cannot be confused with any other species in
Belgium. It resembles for the most O. glazioviana (large
petals, stigma branches elevated high above the anthers,
etc.). It differs from the latter by its uniformly green sepals and lower buds and by the total absence of hairs with
a red bulbous base.
Oenothera oehlkersii was already collected in Liège
in 1884 but all of the remaining inds are (very) recent.
Since 2001, this species has been found fairly often (e.g.
in Balegem, Bruges, Ghent, leper, Mouscron, Roeselare,
Zonhoven, Zwijnaarde), sometimes in large amounts (for
example, at an old dumping site alongside the E40 motorway in the vicinity of Zwijnaarde). Currently, it is a fairly
regularly found species, partially due to the fact that it is
easy to identify. O. oehlkersii has been claimed as a hybrid of O. glazioviana and O. suaveolens Desf. parentage.
However, the presence of a long style is a recessive character that rather suggests a mutant (translocation) of O.
glazioviana, not a hybrid (comm. R. Jean, January 2009).
• Oenothera pycnocarpa Atkinson & Bartlett in Bartlett,
Rhodora 15: 83. 1913.
= O. chicaginensis de Vries ex Renner & Cleland, Z. Indukt. Abstammungs- Vererbungsl. 66: 275. 1933.
Figure 18.
Probable area of origin: North America.
This species is a recent newcomer to Belgium (the oldest inding dates from 1992 in Queue-du-Bois). It was
regularly found, especially since 2007, at several different sites in the port of Antwerp as at a demolition site in
the port of Bruges. The largest population (at least 1000
plants) was, also in 2007, discovered at an excavated
coal mining spoil heap in Gilly near to Charleroi. At this
site it appeared that Oenothera pycnocarpa was more or
less established; however, three years after its discovery
it completely disappeared, despite its habitat remaining
unchanged (comm. M. Lannoy, August 2014). This species bears some resemblance to O. biennis (medium sized
petals, green sepals) but differs from it by the presence of
hairs with a red bulbous base, the stem which in the lower
half is colored red (also often peach-colored) and in the
upper half green, at least at the beginning of lowering.
Moreover, O. pycnocarpa lowers rather late, from the
second half of July to the beginning of August.
In the west of Germany it is the 3rd most numerous
species (e.g. Wittig et al. 1999, Wittig & Tokhtar 2003).
Bearing this in mind, it is possible that it has been overlooked in the east of Belgium.
• Oenothera rosea L’Hér. ex Ait., Hort. Kew. 2: 3. 1789.
Probable area of origin: Central America.
This unmistakable species, which at irst sight resembles
more an Epilobium (on behalf of its relatively small, rose
coloured petals), was collected on several occasions in
the 19th century in Wallonia. Around 2000, it was also
observed at an abandoned coal mining site near to Saint
Ghislain, where it did not persist (comm. P. Dupriez).
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
29
Figure 18. Oenothera pycnocarpa. Brugge, industrial area, demolition site, August 2007. This species is related to O. biennis but
differs from it by its red-punctate stem that is always reddish in the
lower half.
Figure 19. Oenothera rubricalyx. Rekkem, LAR-transportzone, grassland, July 2014. This species differs from O.
glazioviana by its entirely blood red sepals.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
30
In the extreme Southwest of France, Oenothera rosea
is commonly naturalized.
• Oenothera royfraseri Gates, Philos. Trans., Ser. B 226:
285, ig. 30, 31. 1936.
= O. turoviensis Rostański, Fragm. Florist. Geobot. 11:
514. 1965.
Probable area of origin: North America.
In common with other species, including Oenothera delexa and O. parvilora, this species also has small petals.
It can be differentiated from O. delexa by the presence of
hairs with a red bulbous base on the stem. By the absence
of a curved inlorescence as well as the basally contiguous
sepal tips it can be differentiated from O. parvilora.
Oenothera royfraseri was found in large numbers at
a demolition site in the port of Antwerp in 2007 but not
conirmed subsequently.
• Oenothera rubricalyx Gates, Rep. (Annual) Missouri
Bot. Gard. 20: 133. 1909.
≡ O. glazioviana f. rubricalyx (Gates) Lambinon, Nouv.
Fl. Belg., Grand-Duché Luxemb., Nord France, 5th ed.:
1048. 2004.
Figure 19.
Probable area of origin: North America.
A beautiful population of this species was discovered in
dry grassland (former sand raised site) in the Lauwe/Aalbeke/Rekkem transport zone (LAR) in 2014. The species
occurs in fairly large amounts and looks more or less established.
Oenothera rubricalyx is a mutant of O. glazioviana
with entirely deep red sepals. It is striking and garden
worthy. It seems to be rare in the wild in Europe and was
previously only reported from coastal dunes near Etaples
in northwestern France (Rostański et al. 1994, Lambinon
& Verloove 2012).
• Oenothera rubricaulis Klebahn, Jahrb. Hamburg. Wiss.
Anst. 31: 12. 1914.
Figure 20.
Probable area of origin: Europe.
Although this species was already collected in 1887 (Antwerp) it has only become more or less widespread in recent years. Especially around the city of Ghent (e.g. in the
port area, where it has been observed at least since 1985
and where it is irmly established now) Oenothera rubricaulis has become a reasonably common species. Recently, it has also been seen more often around Antwerp.
It differs from most other species by its small to medium
sized petals and the deep red inlorescence axis. It is similar to O. ersteinensis; however, the latter has red-bulbous
hairs with a cylindrical (not conical) base and mostly redstriped sepals. However, O. rubricaulis most resembles
O. rubricauloides (see under).
Elsewhere in Europe, Oenothera rubricaulis is also
a rapidly expanding species. It is even the second most
numerous Oenothera species in Scandinavia (Rostański
2006) and in Eastern Europe, it is considered – along with
O. biennis and O. depressa – the species with the highest
invasion rate (Tokhtar & Groshenko 2014).
Figure 20. Oenothera rubricaulis.
Zeebrugge port area (near Baai van
Heist), ruderalized sand dunes, July
2007. Characteristic features of this
species, compared with the similar
O. ersteinensis, are the entirely green
sepals and the stem hairs that have a
conical base.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
31
• Oenothera rubricauloides Rostański, Ann. Bot. Fenn.
44: 395. 2007.
= O. rubricaulis var. longistylis Gutte & Rostański, Ber.
Arbeitsgem. Sächs. Bot. 11: 187. 1981.
Probable area of origin: Europe.
This hybridogenous species was collected in 2007 on two
occasions around Ghent, each time at locations where
(among others) Oenothera glazioviana as well as O. rubricaulis were found. Rostański (2007), who recently described this taxon, did not cover its possible relationship.
A cross between the two mentioned species appears quite
possible: O. rubricauloides is in all loral characteristics
more or less intermediate between both and the stigma
branches are somewhat elevated above the anthers (hence
the variety name if considered an infraspeciic taxon of O.
rubricaulis). Alternatively, Rostański & Karlsson (2010)
suggested a hybrid formula O. muricata × O. biennis.
In Scandinavia this taxon is reasonably widespread and
for this reason it is considered as an independent species.
• Oenothera stricta Ledeb. ex Link, Enum. Hort. Berol.
Alt. 1: 377. 1821.
Probable area of origin: South America.
At the end of the 19th century (at least between 1878 and
1900), Oenothera stricta was more or less established at
the old city walls of Antwerp (“station abondante”). Now
and then it was also collected elsewhere (e.g. De Panne,
Fleurus, Wilsele, etc.) but perhaps always as an ephemeral
alien. In the 20th century only two collections are forthcoming: Bellecourt (around 1950) and Angleur (1984).
Elsewhere in Europe, O. stricta is, in many places, established, mostly in coastal sand dunes (e.g. France, Great
Britain, Portugal, etc.). It obviously is conined to the climatologically milder parts of Europe and may perhaps be
not fully hardy in Belgium.
Typical for this species are its nearly hemispherical
seeds, the capsules which are gradually narrowing towards base and the petals which are turning reddish or
pinkish after lowering.
• Oenothera subterminalis Gates, Philos. Trans., Ser. B
226: 278, Fig. 26, 27. 1936.
= O. silesiaca Renner, Ber. Deutsch. Bot. Ges. 9: 455.
1942.
Figure 21.
Probable area of origin: North America.
Just like Oenothera parvilora, this species belongs to the
series Rugglesia and thus combines the drooping tip of
the inlorescence (just before lowering) with the sepal
tips which are separated from the very base. The petals are
mostly larger, the stem has a deeper red color and sepal
tips are longer (see key).
Oenothera subterminalis is one of the characteristic species found in the coal mining basins of northwest
France. Strangely, it is virtually absent at identical habitats found just over the border in Belgium (Borinage) (see
Figure 21. Oenothera subterminalis. Hénin-Beaumont (France),
coal mining spoil heap, July 2009. This species has narrowly
lanceolate leaves, a deep red stem and relatively small petals.
Before lowering its inlorescence axis is slightly curved.
also Jean 1975). In 1979, it was collected on a single occasion at a spoil heap in Châtelineau. In 2004, it was also
collected near the Zeebrugge port area (Baai van Heist)
together with O. wratislaviensis, its possible hybrid with
O. canovirens (see earlier).
This species was described by Renner on the basis
of European material (as Oenothera silesiaca). As in
similar cases, later it appeared that it was already earlier
described in America (as O. subterminalis) (Rostański
1985), although both may be distinct as well (comm. R.
Jean, August 2014).
• Oenothera victorinii Gates & Catches., J. Linn. Soc.,
Bot. 49: 182. 1933.
= O. nissensis Rostański, Fragm. Florist. Geobot. 11: 508.
1965.
= O. rostanskii Jehlík, Folia Geobot. Phytotax. 20(4): 439.
1985.
Figure 22.
Probable area of origin: North America.
Oenothera victorinii mostly resembles O. biennis and
O. delexa (see earlier). This species was only collected
in Belgium for the irst time in 1992 (Lasne). In the last
few years it has certainly been observed more often, for
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
32
Figure 22. Oenothera victorinii. Osłowo, Podlaskie
region, Siemiatycze commune (Poland), July 2011.
This species much resembles O. delexa but it has
slightly larger petals.
Figure 23. Oenothera villosa. Brugge,
E-side of Boudewijnkanaal, sand storage area, July 2014. This species shares
the soft greyish pubescence with O. depressa but its stem is hardly (or not at
all) red-punctate, leaf margins are not
crispate and lowers are chasmogamous.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
33
instance in Gentbrugge (1998-2004) and Roeselare. At
both sites it persisted over the course of several years but
eventually disappeared. It was also recently collected at
several locations around Antwerp.
Just as with the previous species, it turned out only recently that this species was already earlier known in North
America and thus was needlessly described from Europe
(Rostański 1985).
• Oenothera villosa Thunb., Prodr. Pl. Cap. 1: 75. 1794.
Figure 23.
Probable area of origin: North America.
Oenothera villosa was discovered in 2007 in a sand storage area alongside the Boudewijnkanaal in Bruges and
regularly conirmed subsequently (ca. 5-10 plants still
present in 2014).
This species from section Devriesia is much reminiscent of Oenothera canovirens (both with a velvety leaf indumentum, lat leaf margins, hairs with red bulbous bases
absent or scarce on the stem, etc.). It is distinguished from
the latter by its capsule teeth that are truncate to nearly
round at apex instead of emarginate.
Acknowledgements. – Raymond Jean (France) and
Adam Rostański (Poland) are thanked for their general
contribution to this paper. Kevin Balkwill (South Africa),
Werner Dietrich (Germany) and Warren Wagner (U.S.A.)
kindly provided relevant literature. Michel Lannoy and
Rutger Barendse provided chorological data for particular
species. Sven Bellanger (Meise) prepared the line drawings. Finally, Jacques Lambinon (Liège) kindly assisted
with data from the LG herbarium, and Steven Janssens
(Meise) provided information on genetics.
References
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Appendix: overview of specimens examined
In this overview all species of Oenothera ever recorded
in Belgium (including those recorded exclusively prior
to 1950) are alphabetically listed. Records are provided
chronologically per province. Records from the current
provinces Flemish Brabant and Walloon Brabant are
grouped together.
Oenothera afinis
• Liège: Vovegnez, graviers, 09.1909, P. Halin s.n. (LG); Vovegn
ez (Wegnez), berge de la Vesdre, 12.09.1909, M. Halin s.n. (BR).
Oenothera angustissima
= O. rubricuspis Renner ex Rostański
• Brabant: Héverlé, talus, 08.1864, O. De Dieudonné s.n. (BR);
Tirlemont, talus du chemin de fer, 08.08.1864, H. Vandenborn
s.n. (LG); Dieghem, vias ferreas, 1877, C. Baguet s.n. (BR); entre
Groenendael et La Hulpe, ligne du chemin de fer, 08.1877, O.
Hecking s.n. (BR);
• Limburg: StTrond, talus du chemin de fer, 30.07.1860, H.
Vandenborn s.n. (LG); StTrond, bord de chemin, 24.07.1861, A.
Thielens s.n. (BR); StTrond, 07.1862, A. Thielens s.n. (LG); St
Trond, talus du chemin de fer, 05.08.1862, H. Vandenborn s.n.
(BR); StTrond, 1864, Vandenborn s.n. (BR); StTrond, talus du
chemin de fer, 08.08.1864, Vandenborn s.n. (BR, LG); StTrond,
talus du chemin de fer, 22.07.1865, H. Vandenborn s.n. (LG);
StTrond, talus du chemin de fer, 22.07.1865, H. van Heurck &
A. Martinis s.n. (BR, GENT); StTrond, talus du chemin de fer,
08.1867, H. Vandenborn s.n. (BR); StTrond, bords des chemins,
cette plante n’existe en Belgique que sur les talus d’un chemin
de fer à StTrond, où elle tend à se naturaliser, 05.08.1867, A.
Thielens & A. Devos s.n. (BR, GENT, LG); StTrond, talus du
chemin de fer, 1869, J.E. Bommer s.n. (BR); StTrond, 10.1870,
P. de Chestret 154 (BR); StTrond, fortiications, 07.1884, P.J.
Delrez s.n. (LG).
Oenothera biennis
• Antwerp: Anvers, endroits frais, bassin de natation, 06.1855,
Tosquinet s.n. (BR); Anvers, Kiel, digue de l’Escaut, 08.1860,
A. Vanheurck s.n. (LG); Tongerlo, sous les murs de l’abbaye,
07.1862, Carnoy s.n. (BR); Tongerloo, 07.1863, A. Thielens s.n.
(BR); Lierre, bords des chemins, 11.07.1874, L. Piré s.n. (BR);
Anvers, fortiications près de la porte de Wilrijk, 23.06.1878, H.
Vandenbroeck s.n. (BR); Calmpthout, terrain inculte, 08.07.1882,
J. Hennen s.n. (BR); Brasschaet, au Peerdsbosch, station de
Vieux Dieu, 07.1896, J. Spas s.n. (BR); Anvers, fortiications
à Berchem, 07.1900, R. Godding s.n. (BR); Wilryck, fortise 6,
09.1903, C. Picquet s.n. (BR); Emblehemsas, helling van den
aquaduc, 06.07.1913, E. van Rompaey GIII/1436.1 (BR); He
renthals, colline, 25.07.1920, V. Lambert s.n. (BR); St. Anna,
chemin (naturalisé), 15.08.1934, G. De Gottal s.n. (BR); Ant
werpen, Zuidervesting, grazige plaats, 17.08.1942, Frison s.n.
(GENT); Malines, Coloma, talus du chemin de fer, 15.07.1949, N.
Cnops 49.118 (BR); Deurne, op droge grond, 19.07.1951, H. Mer
vielde s.n. (BR); Brecht, le long de voie ferrée, 07.1953, J. Plancke
104 (BR); Kruisschans, zandgrond, 12.07.1953, E. Jacques 341
(BR) ; Rijmenam (IFBL D5.31.24), zandig terrein, 13.07.1956,
A. Jans 127/56 (BR); Postel, Moeren, rudéral, bord de route,
02.09.1956, A. Lawalrée 8078 (BR); Postel, chemin sablonneux
près de la tourbière de Postel, 21.09.1956, J. Lambinon (LG);
AntwerpenWest, zandgrond, 11.07.1957, Léothade De Ruyver
s.n. (BR); SintAnna (IFBL C4.26.14), opgespoten gronden,
28.06.1959, E. van Rompaey GIII/1436.1 (BR); Postel, digue d’un
étang, 03.08.1965, A. Lawalrée 13185 (BR); MolPostel, langs
waterkant, 03.08.1965, S. Peeters 123 (BR); MolMaat, langs het
kanaal aan sluis, 08.1967, H.G. Rabijns 1465 (BR); Turnhout,
weg naar BaarleHertog, grazige ruigte, niet afgemaaide plaats,
04.08.1974, J. Aerts 74/57 (GENT); Turnhout, Kempenlaan, tus
sen Gierledreef en Antwerpsesteenweg, opgehoogde braak
liggende grond langs de weg, 09.07.1976, J. Aerts 76/38 (GENT);
• Brabant: KesselLoo, chemin de fer (talus), 07.1855, C. Baguet
s.n. (BR); près de Bruxelles, naturalisé à Groenendaal, 08.1856,
Martinis s.n. (BR); Groenendael, 18.07.1861, L. Piré s.n. (BR);
Uccle, sur les hauteurs, 09.1862, L. Van Den Borren s.n. (BR);
Betekom, bords du Demer, 20.07.1863, C. Baguet s.n. (BR);
Louvain, parc, talus du chemin de fer, 07.1865, C. Baguet s.n.
(BR); Forest, talus sablonneux, 06.07.1866, L. & V. Coomans s.n.
(BR); Auderghem, 1868 (BR); Forest, champ sablonneux, L. & V.
Coomans s.n. (BR); Forest, lieux incultes, 25.07.1875, A. Gravis
s.n. (LG); St. JossetenNoode, talus du chemin de fer, 18.06.1883,
L. Guelton s.n. (BR); Ottignies, 05.1885, H. Vindevogel s.n. (BR);
Louvain, talus du chemin de fer, 02.06.1887, A. Busschodts s.n.
(BR); Parck, talus du chemin de fer, près de l’abbaye, 06.1888,
C. Piquet s.n. (BR); Blauwput, 29.08.1894, E. Michel s.n. (BR);
Wilsele, lieux incultes, 07.1896, C. Baguet s.n. (BR); Uccle (Crab
begat), 06.09.1918, F. Stockmans s.n. (BR); Forest, derrière le
Parc Duden, carrières de sable, 10.09.1919, F. Stockmans s.n.
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
36
(BR); entre Vilvorde et Haeren, talus du chemin de fer, <1920,
Vermoesen & Guns s.n. (BR); Louvain, talus du chemin de fer en
tre les 2 lignes de Malines et d’Aerschot, 30.08.1920, Vermoesen
s.n. (BR); Louvain, Blauwput, voie du chemin de fer vers Malines,
30.08.1920, Vermoesen s.n. (BR); EtterbeekCinquentenaire,
cendrée de la station, 14.07.1923, E. Michel s.n. (BR); Haeren,
talus le long des voies du chemin de fer de l’état, 26.09.1923,
V. Lambert s.n. (BR); Uccle, <1929, J. Henry s.n. (BR); Vilvorde,
remblai du chemin de fer, 02.07.1930, De Vits s.n. (BR); Vilvorde,
remblai du chemin de fer, 28.07.1930, V. Lambert s.n. (BR, LG);
Louvain, canal, 07.07.1933, R. Mosseray & F. Vandevelde s.n.
(BR); Aarschot, terrain inculte le long du chemin de fer de cein
ture AnversDiest, 29.07.1937, E. Michiels s.n. (BR); Parc (lez
Louvain), talus de la tranchée de la voie ferrée, 07.07.1941, A.
Lawalrée 720 (BR); Genval, décombres, 29.06.1946, J.E. Rogis
ter s.n. (BR); Schaarbeek, gare de formation, 07.1947, E. Michel
s.n. (BR); Bruxelles, jonction nordmidi, décombres, 08.1947, E.
Michel s.n. (BR); Woluwé, bois, 06.07.1950, M. Couteaux s.n.
(BR); entre OudHeverlee (gare) et CorbeekDijle, décombres au
bord du chemin, 01.07.1951, A. Lawalrée 3619 (BR); KesselLo,
Blauwput, puinen, 12.07.1951, J. Pelgrims s.n. (BR); Anderlecht,
décombres, 08.1954, s.c. (BR); Heverlee, wegkant, 01.11.1954,
M.T. De Roo s.n. (BR); Auderghem, coin du Boulevard du Souve
rain et de l’Avenue HermannDebroux, terre remuée cette année
pour les travaux du viaducautoroute, 23.09.1973, A. Lawalrée
18221 (BR);
• East Flanders: Aeltere (Aalter), Craenepoel, s.d., s.c. (GENT);
Aeltere, sapinières, 1851, Coemans s.n. (BR) + Gand, chemin
de remblais dans les prés aux moines (semences probable
ment apportées avec les décombres), 1851, Coemans s.n. (BR);
Gand St.Pierre, 08.1854, Scheidweiler s.n. (BR); Tronchiennes
(Drongen), 07.1855, s.c. (GENT); Gand, 1871, s.c. (BR); Bellem,
station, 07.1873, E. Vandermeersch s.n. (GENT); Bellem, bois
(coupes), naturalisé, 19.07.1883, L. Magnel s.n. (BR); Waasmun
ster, Uilenkasteel, 21.07.1903, Guns s.n. (BR); Assels, oostzijde
spoorweg, 15.09.1931, s.c. (GENT); SintAmandsberg, spoor
berm, 23.07.1966, J. Van Den Haute 164 (BR); Gent, even voor
bij Meulestedebrug, driehoekig terrein tegenover Café “Les Rout
iers”, kruidenrijke vegetatie met Malva sylvestris, 06.07.1974, E.
Robbrecht 638 (BR); Gent, RUG, Plantentuin Maton, K.L. Lede
ganckstraat 35, braakliggend terrein, zonnig, droog, opgehoopt,
05.08.1976, H. Viane 62 (GENT); SintDenijsWestrem, station,
in verwaarloosd terrein, vroeger opslagplaats van goederen en
kolen, 03.08.1978, D. Duytschaever 6061 (GENT); Zelzate,
weg GentZelzate via Linkeroever, graafwerken ten N van ex
ecutieoord Rieme (IFBL C3.33.34), ruigte, 17.09.1982, E. Rob
brecht 2530 (BR); Appels (IFBL D3.28.24), gestorte grond rand
weg, 16.09.1992, K. Thielemans s.n. (BR); Harbour of Ghent,
Singel near Farmanstraat (IFBL D3.13.11), ruderal gravelly road
verge, 24.06.2000, F. Verloove 4409 (BR); Gent, Hundelgem
sesteenweg near river Schelde (IFBL D3.23.33), waste ground,
27.06.2004, F. Verloove 5714 (BR); Belzele (NGent), N of Ring
vaart near junction with Brugse Vaart (IFBL D3.11.21), stony bor
der of canal, common, 24.06.2007, F. Verloove 6777 (BR); Kerk
brugge (NGent) (IFBL C3.52.24 + 53.13), former railway yard,
24.06.2007, F. Verloove 6778 (BR);
• Hainaut: BraineleComte, tunnel, 05.07.1949, S. Depasse
88/22 (BR); Quivrain, railway yard, gravelly soil, 04.07.2004, F.
Verloove 5726 (BR); Frameries, bas du terril du PASS, 08.2006,
D. GeerinckCoutrez 9701 (BR); Dour, La Grande Machine à
Feu (terril) (IFBL G3.53.1221), coal mine heap, 07.07.2007, F.
Verloove 6872 (BR); Mons, Mont Ostène, terril du Pass (IFBL
G3.45.41), terril, zone en combustion, 25.07.2007, F. Verloove
6866 (BR);
• Liège: Rochefort, s.d., F. Crépin s.n. (BR); Tilff, 06.1871, H.
Donckier s.n. (LG); Fraipont, rocailles au bord de la Vesdre, 1872,
s.c. (GENT); entre Trooz et Fraipont, lieu rocailleux, 1872, Strail
s.n. (BR); Ougrée, bord de la Meuse, 30.06.1872, J.L. Wathelet
s.n. (BR); entre Limbourg et [...], le long de la […], 08.1874, O.
De Dieudonné s.n. (BR); environs de Visé, 1876, A. Hardy s.n.
(BR); Flaire, 09.09.1883, P.J. Delrez 450 (LG; mixed collection
with O. cruciata); Soiron, […], 09.09.1883, P. Delrez s.n. (LG);
Modave, jardin, 04.08.1884, G. Evrard s.n. (BR); Esneux, 1888,
C. Sladden s.n. (BR); ValDieu (Charneux), bord d’un chemin,
07.1890, M. Halin s.n. (BR); Tilff, bords de l’Ourthe (il n’y en
avait que 2 ou 3 pieds, j’ai remarqué qu’il était plus abondant
près d’Esneux), 26.06.1892, C. Sladden s.n. (BR); Comblainau
Pont, 29.07.1902, H. de Luesemans s.n. (BR, LG); Lambermont,
1905, P. Doubleman s.n. (LG); Esneux, sous la Roche aux Fau
cons, sentier broussailleux, 10.07.1906, A. Maréchal s.n. (LG);
Vovegnez (Wegnez), berge de la Vesdre, 07.1907, M. Halin s.n.
(BR); Seraing, décombres, 09.08.1907, H. Henin s.n. (LG); Val
Benoit, terrain vague, 13.07.1908, M. Halin s.n. (BR); Theux,
13.07.1908, s.c. (BR); Liège, 08.1908, P. Doubleman s.n. (LG);
ViersetBarse, gare de Régissa, 08.07.1919, A. Charlet s.n. (LG);
Vesdre, 12.09.1920, A. Visé s.n. (LG); Ensival, Vesdre, 07.1940,
A. Visé s.n. (LG); Fond des Cris (Chaudfontaine), 09.1946, L. Re
nard s.n. (LG); Tilff, quai Ste. Anne, sentier longeant la rive droite
de l’Ourthe, 18.08.1962, A. Lawalrée 11711 (BR);
• Limburg: Hasselt, derrière le cimetière, 1877, O. Geraets s.n.
(BR); Wychmael (Beverloo), 16.07.1911, M. Guns s.n. (BR);
CurangeHasselt, voies du chemin de fer, aux environs des at
eliers de réparation, assez répandu, 07.08.1920, Vermoessen
s.n. (BR); Leopoldsburg, sables, 07.1956, J. Lebeau s.n. (BR);
Genk (IFBL D7.51.22), port, 26.08.1956, A. Lawalrée 8032, 8034
(BR); GenkWaterschei, déblais d’usine, 21.07.1958, A. Lawal
rée 10159 (BR); Bree (‘t Hasselt), langs brug, oever ZuidWil
lemsvaart, 17.07.1962, J. Pelgrims 1454 (BR); Leopoldsburg,
langs een weg, 07.1967, H.G. Rabijns 1462 (BR); Zolder, <1998,
L. Janssen s.n. (BR);
• Luxembourg: Orval, abbaye, 26.07.1856, Gravet s.n. (BR);
Forêt d’Orval, coteau, 08.1882, E. Chapuis s.n. (LG); Virton, à la
grande route vers Rabais, 08.08.1949, A. Lawalrée 2417 (BR);
Lahage, halte, près de la halte de la voie ferrée, 04.07.1963, A.
Lawalrée 12499 (BR); limite des communes Mirwart, Arville et
Smuid, bord de la voie ferrée, 09.09.1964, A. Lawalrée 12864
(BR); Buzenol, gare, 27.07.1968, M. Lambert s.n. (LG); entre Or
val et […], route, 24.07.1974, D. Geerinck 782 (BR);
• Namur: Anseremme, bords de la Lesse, 07.1867, C.L. Guilmot
s.n. (BR); Han, bois à l’entrée de la grotte, 06.1882, P. de Ches
tret s.n. (BR); Eprave, abondant à la lisière d’un bois, 07.1886,
F. Pietquin s.n. (BR); Belvaux, 1889, H. Verheggen s.n. (BR); La
Plante, naturalisé, 06.1890, Leclerc s.n. (BR); Belvaux, coteau
près du Gouffre de Belvaux, 09.1936, O. Gras s.n. (BR); Eprave,
route de Han, 08.1938, O. Gras s.n. (BR); Anseremme, décom
bres, 06.09.1942, O. Gras s.n. (BR); Houyet, talus du chemin de
fer vers Hout, 07.1943, J. Legrain s.n. (BR); Durnal, Pipeti, Vallée
du Bocq, rive droite, talus dans une chênaie silicicole, 17.07.1977,
E. Sérusiaux 77/B/434 (LG); Namur, chemin du Pont de Bricques
à Jambes (IFBL G5.47.11), 10.07.1986, D. Geerinck 3728 (BR);
environs de Sosoye, long de l’ancienne voie ferrée, ballast de la
voie ferrée, 13.10.1990, D. Geerinck 1569 (BR); vallée de la Moli
gnée, près de l’abbaye de Maredsous, ballast de la voie ferrée,
13.10.1990, G. Bruynseels 1662 (BR);
• West Flanders: NieuportBains, lieux incultes, 23.07.1892, C.
Baguet s.n. (BR); La Panne, lieux incultes, 27.07.1892, C. Baguet
s.n. (BR); Duinbergen, dunes, 07.1929, E. Dewildeman s.n. (BR);
Wenduine, 07.1946, L. Renard s.n. (LG); Comines, bord de la Lys,
28.06.1947, J. Baily s.n. (BR); Adinkerke, sable au bord d’une
avenue, 03.09.1954, J.L. De Sloover 2581 (BR); De Panne,
dunes, 08.1956, J. Lebeau s.n. (BR); Avekapelle (IFBL D1.11.11),
braakliggende akker, 06.10.1997, H. Ruysseveldt 2007 (BR); Gits
(IFBL D1.38.13), ruigte bij het station, 27.08.1987, F. Verloove
6 (BR); Nieuwpoort (IFBL C1.41.11), open plek geixeerd duin,
07.08.1995, H. Ruysseveldt s.n. (BR); Waregem (Eertbrugge)
(IFBL E2.26.13), grazige vegetatie berm, brug over autosnel
weg, 24.06.2003, H. Ruysseveldt 3459 (BR); Brugge, Eside of
Boudewijnkanaal (IFBL C2.11.24), sand deposit, 08.08.2007, F.
Verloove 6853 (BR).
Oenothera cambrica var. cambrica
• East Flanders: Zwijnaarde (Gent), Ringvaart (Wside), close to
E40 motorway (IFBL D3.32.21), workedup roadside, 17.08.2014,
F. Verloove 10953 (BR).
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
37
• West Flanders: Roeselare, Bruggesteenweg x Leopold IIIlaan
(IFBL D1.48.31), ruderal area, 21.07.2001, F. Verloove 4993 (priv.
herb. F. Verloove).
• Oenothera cambrica var. impunctata
• East Flanders: Zwijnaarde (ZGent) (IFBL D3.32.24), road
verge, along river Schelde, one specimen, 10.07.2007, F. Ver
loove 6809 (priv. herb. F. Verloove).
Oenothera canovirens
• Antwerp: St.Anna (IFBL C4.26.14), opgespoten gronden,
28.06.1959, E. Van Rompaey 14362 (BR);
• Liège: Entre Ensival et Pepinster, graviers de la Vesdre,
09.1901, P. Halin s.n. (LG).
Oenothera cruciata
• Brabant: Forest, 06.1870, Coomans s.n. (BR);
• Liège: Flère, Gomélèvai, 09.09.1883, P. Delrez 450 (LG; mixed
collection with O. biennis).
Oenothera delexa
• Antwerp: AnversOuest (rive gauche), 18.08.1955, L. De Ruyver
s.n. (BR); Koersel, bord d’un sentier, 23.09.1962, E. Hostie s.n.
(GENT); BalenNete, route de Postel, bord de la route, 03.08.1965,
A. Lawalrée 13157 (BR); AntwerpenNoord (IFBL C4.16.33),
opgespoten gronden ten W van Oosterweel, 04.08.1965, E. Van
Rompaey 14362 (BR); Hoboken, langs weg, 05.10.1965, M. Van
den Wijngaert s.n. (BR); Turnhout, Dombergstraat, tussen Wa
tertappingstraat en pannenfabriek Tuca, inz. noordelijke boord
van de weg, 22.09.1969, J. Aerts s.n. (GENT); Turnhout, kanaal,
noordelijke oever nabij de grens TurnhoutBeerse, 25.07.1974,
J. Aerts 74/48 (GENT); MechelenBattel, oude Gentse Steenweg
nabij Heffen, Heembeemd (IFBL D4.27.13), verwilderd langwer
pig perceel, droog, spoorwegsintels, 14.08.1974, L. Vanhecke
4260 (BR); Lillo (IFBL B4.54.21), opgespoten gebied langs
de Galgeschoor, droge open zandgrond, pioniersbegroeiing,
11.09.1974, L. Vanhecke 4318 (BR); Turnhout, Everdongenlaan,
enkele meters ten zuiden van de brug over de Aa, op de rand van
de weg, 09.07.1976, J. Aerts 76/37 (GENT); Turnhout, kanaal,
noordelijke dijk, nabij de grens met Beerse, op de oever van het
kanaal, 22.07.1976, J. Aerts 76/48 (BR, GENT); Turnhout, werf
De Clercq (IFBL B5.48.11), tussen steenslag, 1 eks., 22.07.1979,
A. Vermeijen 79/75 (BR); Mechelen, Mechels Veld, terrain vague
près Du Pont de Nemours, 06.07.1980, R. Wechuysen 1023 (BR);
Turnhout, Nieuwe Kaai (IFBL B5.47.42), wegenbouw Van Gorp,
aardhopen grondwerken, 05.09.1981, J. Aerts 81/79 (GENT);
Burcht (comm. actuelle Zwijndrecht) (IFBL C4.25.44), terrain
vague sabloargileux près de l’Escaut, 24.09.1990, J. Lambinon
90/B/504 (BR, LG); Harbour of Antwerp, between 4e and 5e Ha
vendok (IFBL C4.16.32), road verge, along railway track, 1 ex.,
29.07.2007, F. Verloove 6870 (BR);
• Brabant: Anderlecht, talus de la voie ferrée à la Petite Ile,
09.1938, G. André s.n. (BR); Evere, terrains vagues, 17.09.1944,
E. Hostie s.n. (GENT); Evere, Kerkebeek, puinen van verwoeste
huizen bij de kerk, 23.10.1948, E. Michiels s.n. (BR); JetteDiegem,
woest terrein bij den Paelboschwegel, 23.07.1949, E. Michiels
s.n. (BR); KesselLo, Blauwput, stort van puinen, 28.08.1951, J.
Pelgrims s.n. (BR); Anderlecht, remblai chemin de fer, 09.1951, A.
Lefebvre s.n. (BR); Nodebais, décombres, 27.09.1954, L. Muyl
dermans 858 (BR); Leuven, Keizersberg, wegkant, 15.10.1954,
M.T. De Roo s.n. (BR); KesselLo, Martelarenlaan, braakliggend
terrein langs de spoorweg, 25.07.1955, J. Pelgrims 109 (BR,
GENT, LG); Hofstade, prairie boissée près grand lac, sol sablon
neux, 24.09.1955, R. Wilczek 1532 (BR); Auderghem, Parc des
Princes, Avenue Leemans, terrain vague, 22.07.1960, A. Lawal
rée 11030, 11032 (BR); KesselLoo, terrain vague, 31.07.1962,
J. Bouharmont 1383 (BR); Ixelles (Boondael), Chaussée de
Boitsfort, le long du chemin de fer, plante très abondante partout,
08.1965, L. Dubois 1647 (BR); Auderghem, à l’angle de l’Avenue
HermannDebroux et du Boulevard du Souverain, terre remuée il
y a quelques mois à peine lors de la construction de l’autoroute en
viaduc, une colonie abondante et uniforme, 23.09.1973, A. Lawal
rée 18225 (Soc. Ech. Pl. Vasc. Eur. Bass. Médit.: BR, GENT, LG);
Auderghem, Rue de la Vignette (IFBL E4.36.24), terrain rudéral
isé sur sable, 08.1977, D. GeerinckCoutrez 1686 (BR); Kessel
Lo, Koning Albertlaan, straatberm, 08.08.1978, s.c. (BR); Haacht,
station WespelaarTildonk, rond « La Corbeille » (IFBL D5.42.41),
20.07.1980, R. Wechuysen 1060 (BR); Uccle, Rue de Bourdon,
terrain sableux pauvre, pelouse ouverte et rase, 01.08.1980, M.
Tanghe s.n. (BR); Anderlecht, Rue Bollinckx, sur terre de remblai,
limoneuse, mêlée, 23.07.1981, M. Tanghe s.n. (BR); Strombeek
Bever, chantier du Ring de Bruxelles, croisement avec le chemin
MeiseHeysel, buttes de sol amené, groupement riche en an
nuelles, 23.09.1981, E. Robbrecht 1592 (Soc. Ech. Pl. Vasc. Eur.
Bass. Médit.: BR, GENT); Boitsfort, Avenue de la Forestière (In
ternational School), pelouse à saules, 04.10.1982, M. Tanghe s.n.
(BR); WatermaelBoitsfort, terrain rudéralisé dans le quartier des
Pêcheries, 09.1984, D. GeerinckCoutrez 3380 (BR); Woluwe
SaintLambert (IFBL E4.27.31), prairie près du Cora, 08.1987,
D. GeerinckCoutrez 4372 (BR); Diest (IFBL D6.31.44), oud stort
langs de Demer, 07.07.1990, E. Jacques 17572 (BR); Waterma
elBoitsfort (IFBL E4.46.42), terrain rudéralisé de la Foresterie,
08.1992, D. GeerinckCoutrez 6716 (BR, LG); Wemmel, Allée des
Peupliers, fasciatie, 1995, Verheyden s.n. (BR);
• East Flanders: Langerbrugge, 1932, […] (GENT); Denderleeuw,
Grote Steenweg, op uiterwaarden, zandgrond, 22.08.1971,
J. Van den Haute 605 (BR); Eke (IFBL D3.41.14), 14.07.1973,
D. Duytschaever 7 (GENT); Oudegem, spoorwegbrug over
de Dender, berm van de spoorwegbrug (grazig en kruidenrijk),
18.07.1976, E. Robbrecht 1055 (BR, GENT); Denderleeuw (IFBL
E3.28.21), recent opgehoogd terrein, 07.08.1976, H. Ruysseveldt
1504 (BR); WelleWellemeersen (IFBL E3.18.23), braakliggende
akker, 11.09.1980, H. Ruysseveldt 1503 (BR); Gent, westzijde
Handelsdok, bij douane (IFBL D3.12.44), grazige braakliggen
de terreinen (opgevuld voormalig dok), 10.1987, E. Robbrecht
3539 (BR); Appels (IFBL D3.28.24), grazige wegkant op dijk,
15.07.1992, A. Van Den Bergh s.n. (BR); Asbeek (IFBL E4.12.14),
braakliggende akker, 22.07.1992, W. De Schrijver s.n. (BR);
Denderleeuw (IFBL E3.28.21), verruigd hooiland, 14.07.1993,
H. Ruysseveldt s.n. (BR); Bornem (Mariekerke) (IFBL D4.12.44),
bij een klein kanaal naar de Schelde, 06.09.1998, D. Geerinck
Coutrez 9319 (BR); Harbour of Ghent, Singel near Farmanstraat
(IFBL D3.13.11), ruderal gravelly road verge, 24.06.2000, F. Ver
loove 4410 (BR); Gent, border of river Schelde at Gentbrugge
brug (IFBL D3.23.13), demolition area of former steelfactory,
abundant, 08.07.2002, F. Verloove 5120 (priv. herb. F. Ver
loove); Gent, river Schelde near Rijmstraat, crack in pavement,
27.06.2004, F. Verloove 5715 (BR); Gent, Voorhavenlaan (IFBL
D3.12.24), ruderal area, 27.06.2004, F. Verloove 5716 (BR); Zwi
jnaarde (ZGent) (IFBL D3.32.24), former dump site near river
Schelde and E40 motorway, 10.07.2007, F. Verloove 6873 (BR);
Zwijnaarde (ZGent), border of river Schelde (IFBL D3.32.24),
ruderal road verge, 10.07.2007, F. Verloove 6874 (BR); Harbour
of Ghent, Sside of Kluizendok (IFBL C3.43), artiicial sandscape,
10.07.2007, F. Verloove 6875 (BR); Zwijnaarde (ZGent), border
of river Schelde (IFBL D3.32.24), road verge, 10.07.2007, F. Ver
loove 6876 (BR);
• Hainaut: MarchiennesauPont, cendrées le long de la voie fer
rée, 08.1949, J. Duvigneaud s.n. (BR); Mons, bord d’un chemin,
10.1949, F. Buxant s.n. (BR); Jamioulx, talus chemin de fer,
08.1954, J. Lebeau s.n. (BR); Viesville, terrain vague, 29.07.1956,
P. Demalsy s.n. (BR); Soignies, carrière de Perlonjour, chantier,
19.07.1969, A. Lawalrée 15859 (BR); ChapellelezHerlaimont,
terril n° 7, pente schisteuse, M. Tanghe s.n., 13.08.1970 (BR);
Cuesmes, terrain rudéral, 03.10.1976, C. Vanden Berghen s.n.
(BR); Baudour, 24.07.1983, S. Depasse 88/102 (BR); Soignies
(IFBL F3.58.32), terrain rudéralisé sur la pierre blanc de la carrière
de Hainaut, 14.09.1985, D. GeerinckCoutrez 3618 (BR); Boussu,
terril SaintAntoine, coalmine heap, 04.07.2004, F. Verloove 5719
(BR); Quivrain, railway yard, gravelly area, 04.07.2004, F. Ver
loove 5727 (BR); Blaton, réserve naturelle « La Grande Bruyère
» (IFBL G3.21.41), ancienne sablière, très commun, 25.07.2007,
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
38
F. Verloove 6869 (BR); Quiévrain, gare (IFBL G3.41.44), along
railway track, 12.07.2007, F. Verloove 6877 (BR); Hensies (Les
Sartis) (IFBL G3.31.44), coal mine heap, 12.07.2007, F. Verloove
6879 (BR); Hornu, terril du Sept (IFBL G3.44.31), coal mine heap,
25.07.2007, F. Verloove 6867 (BR);
• Liège: Dison, terrain vague, 06.1915, P. Doubleman s.n. (LG);
ViersetBarse, gare de Régissa, 08.07.1919, A. Charlet s.n. (BR);
Ensival, Vesdre, 07.1940, A. Visé (BR); Chaudfontaine, carrières,
16.08.1943, A. Isaäcson s.n. (BR); Fond de Cris (Chaudfontaine),
terrain vague, 16.08.1943, A. Visé s.n. (LG); Vallée de la Vesdre,
bord de la route LiègeVerviers, 26.09.1960, N. Cnops 60.163
(BR); Angleur, rive gauche de l’Ourthe, en aval de l’île Rous
seau, terres remaniées, 29.08.1973, J. Duvigneaud 73/B/1131;
Ougrée, ancien crassier des usines métallurgiques, bois Saint
Laurent, 22.07.1974, J. Duvigneaud 74/B/953 (LG); Angleur, gra
viers déversés dans la plaine alluviale de l’Ourthe, 30.07.1974, J.
Duvigneaud 74/B/1013 (LG); Amay, graviers le long de la Meuse,
30.09.1975, W. Bellotte 75/707 (LG); Angleur, rive gauche de
l’Ourthe, 20.10.1975, Hechtermans & Monfort 75/752 (LG); Her
stal (IFBL F7.24.33), décombres d’une vieille usine au nord de
la gare, 09.08.1980, R. Wechuysen 1118 (BR); Jemeppesur
Meuse, terril houiller du Bas Laveu, friche en bordure d’une zone
en combustion, 25.08.1984, J. Lambinon 84/B/640 (LG); Hamoir
(IFBL G7.42.32), au S de l’agglomération, le long de la vallée de
l’Ourthe, rive gauche, site rudéral, 13.08.1985, A. Lawalrée 25745
(BR); Angleur (IFBL F7.43.22), pied du remblais du chemin de fer
près de l’ancienne gare, 27.07.1986, J. Lambinon 86/B/208 (LG);
Angleur (IFBL F7.43.21), pied de mur près du canal de l’Ourthe,
à proximité du Pont Marcotty, 03.08.1988, J. Lambinon 88/B/265
(LG); QueueduBois, parking usine Aldi, 04.09.1992, J. Beau
jean 92/86 (LG); Chênée, décombres au Piedrou, 30.09.1992, J.
Beaujean 92/131 (LG);
• Limburg: Genk, Waterschei, Horenszee, bord de route,
26.08.1956, A. Lawalrée 8010 (BR); Hasselt, rudéral, 07.1960,
L. Delvosalle s.n. (BR); Stokrooi, langs kanaal, 08.07.1960, H.
Vannerom s.n. (BR); Genk, aan de spoorweg van de Kolenhaven,
21.07.1967, J. Rammeloo 309 (GENT); GenkLangerloo, zan
dige wegberm, 22.07.1967, L. Vanhecke 67/4 (BR); Genk, terrain
vague, 02.08.1979, P. Claes s.n. (LG);
• Luxembourg: entre Chantemelle et Châtillon, dans une sablon
nière, 13.08.1968, V. d’Ansembourg 3737 (BR); Forrières, bord
de la voie ferrée vers Jemelle, association rudérale, plusieurs
petites colonies, au total +/ 50 pieds, 04.09.1998, A. Lawalrée
26229 (BR);
• Namur: Namur, terrain vague, 22.09.1954, J.L. De Sloover
2581 (BR); Jambes, bord de la voie ferrée près des usines Ma
terne, 09.1955, J. Lambinon s.n. (LG); Lustin, bord de la voie fer
rée près de la gare, 04.09.1956, J. Lambinon s.n. (LG); Godinne
(îles), colonisatrice de pierres, 06.08.1972, E. Sérusiaux 72/B/38
(LG); Dorinne, rive gauche du Bocq (IFBL H5.28.14), ancienne
carrière de Famennien, 20.08.1978, J. Duvigneaud 78B603 (BR);
Rochefort, carrière abandonnée, 28.08.1978, B. Bastin s.n. (BR);
Namèche (IFBL G6.31.13), friche, 09.08.1983, J. Saintenoy
Simon s.n. (BR); NovillesurMéhaigne (IFBL F5.47.14), gare
désaffectée, 10.08.1983, J. SaintenoySimon s.n. (BR); Grands
Malades (IFBL G5.37.14), friche, 21.08.1983, J. SaintenoySimon
s.n. (BR);
• West Flanders: Brugge, SintMichiels, spoorwegberm,
29.07.1973, R. Viane 93 (GENT); bos van Houthulst, ruigte,
11.09.1977, L. Vanhercke s.n. (BR); Haven Roeselare, Beurt
kaai (IFBL D1.48.43), wegberm, achter een bermmarkering,
04.07.1992, F. Verloove 383 (BR); Rekkem (IFBL E1.58.22),
wand greppel langs maïsveld, 29.07.2000, H. Ruysseveldt 2643
(BR); Roeselare, near the railway station (IFBL D1.48.34), rail
roadbank, several specimens, established, 12.07.2001, F. Ver
loove 4992 (priv. herb. F. Verloove); Zedelgem, ongebruikte
spoorweg, 11.09.2001, H. Ruysseveldt 3009 (BR); Ingelmunster,
railway track, 03.07.2004, F. Verloove 5717 (BR); Kortemark,
railway yard, gravelly area, common, 08.07.2004, F. Verloove
5721 (BR); RumbekeRoeselare (IFBL D1.58.32), former claypit
Ostyn, dump area, 18.07.2004, F. Verloove 5731 (BR); Brugge,
port (IFBL C2.22.11), ruderal area, gravelly soil, 28.06.2007, F.
Verloove 6878 (BR, LG) (hybrid); Zeebrugge (portarea), SE of
Distrigasplant, A. Ronsestraat (IFBL B2.42), sandy ruderal area,
+/ 10 ex., 18.07.2007, F. Verloove 6859 (BR); Heist, Baai van
Heist (IFBL B2.32.33), seadunes, +/ ruderalized, 18.07.2007, F.
Verloove 6860 (BR); Brugge, Pathoekeweg, W of Nijverheidsdok
(IFBL C2.21.22), ruderal area (demolition site), small population
of uniform plants, 08.08.2007, F. Verloove 6856 (BR).
Oenothera depressa
• Hainaut: Hensies, Les Sartis (IFBL G3.31.44), coal mine heap,
scattered specimens, 12.07.2007, F. Verloove 6801 (priv. herb. F.
Verloove, BR);
• Liège: Horto Leod., 1828, R. Courtois s.n. (LG).
Oenothera ersteinensis
• Antwerp: Harbour of Antwerp, Eerste Havendok (IFBL
C4.16.42), sandy, workedup road verge, nice population, >100
specimens, 22.07.2001, F. Verloove 4991 (priv. herb. F. Ver
loove, BR, LG); Harbour of Antwerp, SWside of Hansadok (IFBL
C4.15.23), along railway track, 2 ex., 29.07.2007, F. Verloove
6831 (priv. herb. F. Verloove); Retie (IFBL C6.13.21), omgewerkte
berm, 08.07.2009, R. Barendse s.n. (BR); Brecht, Groot Schiet
veld (Brechtse Baan) (IFBL B4.48.24), zandige braakgrond bij in
gang schietstand, talrijk, 11.07.2010, F. Verloove 8131 (BR); Ant
werp (Luchtbal) (IFBL C4.17.13), ground heaps, by railway track,
03.08.2014, F. Verloove 10951 (BR);
• Liège: OugréeSart Tilman, domaine universitaire, en bordure
de la route près de l’Institut de Botanique, 13.08.1971, P. Auquier
1566 (BR); Chenée, décombres au […], 30.09.1992, J. Beaujean
92/131 (BR);
• Limburg: LommelBalendijk (IFBL C6.25.24), in redelijke ho
eveelheid (met O. glazioviana, O. biennis), 07.08.2010, R.
Barendse s.n. (BR).
Oenothera fallax
• Antwerp: Harbour of Antwerp, SWside of Delwaidedok (IFBL
B4.45.33), sandy border of a railwaytrack, numerous specimens,
22.07.2000, F. Verloove 4627 (priv.herb. F. Verloove);
• Brabant: Bruxelles, chantiers de la jonction NordMidi, 09.1951,
A. Lawalrée 3790 (BR); Bruxelles, chantier de la jonction Nord
Midi, association rudérale, 04.08.1956, A. Lawalrée 7775 (Soc.
Fr. Ech. Pl. Vasc. n° 2982; BR); Auderghem, Avenue Chaudron,
terrain vague, remblai, 11.07.1958, A. Lawalrée 10037 (BR);
Rijmenam (IFBL D5.31.22), langs de baan aan ‘t Zwart Water,
16.08.1967, N. Cnops 67.567 (BR); SintJansMolenbeek, braak
liggend terrein, 22.06.1971, J. Loots 105 (BR; mixed with O.
glazioviana); Haeren, près de Bruxelles, gare de triage, talus ru
déralisé, 29.08.1987, G. Bruynseels 1257 (BR);
• East Flanders: Haven van Gent, GentZelzate via Linkeroever,
graafwerken ten N van executieoord Rieme (IFBL C3.33.34), ruig
te, 17.09.1982, E. Robbrecht 2529 (BR); Port of Ghent, Farman
straat near Kennedylaan (IFBL D3.13.13), roadsides, railways and
rough places, ca. 15 individuals in old railway bed, 19.08.1986,
E. Robbrecht 3122 (BR); Zwijnaarde (ZGent), langs Ringvaart
(IFBL D3.32.24), voormalige zandopspuiting, enkele planten met
O. biennis, 23.06.2001, F. Verloove 4822 (BR); SintDenijsWes
trem, oprit E40 richting Oostende, zandig braakliggend terrein,
talrijk, 21.06.2004, F. Verloove 5655 (BR);
• Hainaut: Dour, terril Grande Machine à Feu, coalmine heap,
04.07.2004, F. Verloove 5720 (BR); Hensies (Les Sartis) (IFBL
G3.31.44), coalmine heap, 04.07.2004, F. Verloove 5718, 5722,
5723 & 5725 (BR);
• Liège: Neupré, RotheuxRimière, jonction des routes de Marche
et d’Esneux, bord rudéralisé de la route, 20.07.1979, P. Auquier
5069 (LG);
• Limburg: Waterschei, ancien […], près de […], 09.1956, J.
Lambinon s.n. (BR);
• Namur: vallée du Bocq, bords des eaux, 08.1889, Dens s.n.
(BR); between Freyr and Waulsort along the river Meuse,
06.09.1991, R. Govaerts 33 (BR);
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
39
• West Flanders: Middelkerke, dune, 25.07.1934, P. [...] s.n. (BR);
Le Coq, wegkanten, 14.08.1939, L. Standaert s.n. (BR); Koksijde,
dans les dunes, très répandue partout, 07.1966, L. Dubois 1646
(BR); Koksijde, près du moulin, 20.08.1970, R. Boutique 808 (BR);
StIdesbald, dune rudéralisée, 28.07.1974, B. Bastin s.n. (BR);
Knokke, Duinbergen, restje duin tussen villa’s langs Kon. baan,
01.11.1978, R. Viane 697 (GENT); KnokkeHeist, dunes rudérali
sées, 07.1982, D. GeerinckCoutrez 688 (BR); Nieuwpoort (IFBL
C1.41.11), open plekjes geixeerd duin, 07.08.1995, H. Ruyssev
eldt s.n. (BR); Wervik (IFBL E1.46.24), tussen resten afgebroken
fabrieksgebouw, 01.08.2000, H. Ruysseveldt 2647 (BR); Brugge,
Pathoekeweg, W of Nijverheidsdok (IFBL C2.21.22), ruderal area
(demolition site), 1 ex., 08.08.2007, F. Verloove 6855 (BR).
Oenothera fruticosa subsp. glauca
• East Flanders: GentWest, Vinderhoute, Vinderhoutse Meersen
(IFBL D3.11.22), ruderale inrijlaan van een afvalverwerkend bedri
jf langs de R4, samen met andere tuinvluchtelingen, 05.07.1998,
F. Verloove 3381 (BR).
Oenothera glazioviana
• Antwerp: Hoboken, braakland aan de staalgieterij, 02.07.1937,
E. Frison s.n. (GENT); Wildert, barre grond, zand, 01.08.1953,
E. Jacques 354 (BR); Essen, ’t Hoefken, adventief in Arnosere
toScleranthetum, 09.10.1956, J. Traets 520 (BR); Duffel (IFBL
D4.18.13), op spoorwegdam, 02.09.1957, N. Cnops 57.313
(BR); Antwerpen (IFBL C4.36.23), afgegraven vestingwal, ruig
te, 18.08.1967, E. Van Rompaey GIII/14351 (BR); Antwerpen
(IFBL C4.36.23), afgegraven vestingwal, ruigte, 21.08.1967,
E. Van Rompaey GIII/14351 (GENT); Duffel, bord de la route
LierreMalines, à l’ouest de la voie ferrée AnversMalines (IFBL
D4.18.13), talus, 02.07.1970, A. Lawalrée 16128 (BR); Turnhout,
Smalvoortstraat (IFBL B5.47.32), bouwwerf, 21.07.1979, J. Aerts
79/25 (GENT); ‘s Gravenwezel, oude baan (IFBL C4.28.22), aan
de voet van de oprit van de brug over het Albertkanaal, zandige
wegkant, 21.07.1986, E. Jacques 15852 (BR); Haven van Ant
werpen, NDelwaidedok (IFBL B4.45.13), spoorwegemplace
ment, 08.10.2001, E. Molenaar s.n. (BR);
• Brabant: ParkHeverlee, spoorweg naar Waver, 01.08.1909,
E. Michiels s.n. (BR); Keerbergen, marécage, 12.08.1920, E.
Hostie s.n. (GENT); Wilsele, talus du chemin de fer, 19.07.1923,
J. Lebrun s.n. (BR); Uccle, sablière, 07.1928, J. Lebrun 2680
(BR); Aarschot, woest terrein achter het station, 30.07.1937, E.
Michiels s.n. (BR); Louvain, MontCésar, pelouses, 29.09.1940,
A. Lawalrée 525 (BR); Aarschot, woest terrein achter het sta
tion, 02.07.1941, E. Michiels s.n. (BR); Uccle, talus de la voie
ferrée, 18.07.1951, G. André s.n. (BR); Bruxelles, chantier jonc
tion NordMidi, rudéral, 04.08.1956, A. Lawalrée 7779 (BR);
KesselLoo, terrain vague, 31.07.1962, J. Bouharmont 1382
(BR); KesselLo, spoorwegberm, 18.06.1964, J. Pelgrims 1822
(BR); SintJansMolenbeek, braakliggend terrein, 22.06.1971, J.
Loots 105 (BR; mixed with O. fallax); Auderghem, Jardin Massart,
grande parcelle expérimentale, 03.08.1976, M. Tanghe s.n. (BR);
Auderghem, Chaussée de Wavre (IFBL E4.37), bord de route,
07.1977, D. GeerinckCoutrez 1591 (BR); près de Haren, gare
de formation, terrain rudéralisé, 01.09.1987, G. Bruynseels 1247,
1552 (BR); Ramsberg (ten E van Tienen) (IFBL E6.21.21), stortp
laats tussen maïsakkers en laagstamboomgaarden, 02.10.2002,
L. Vanhecke s.n. (BR);
• East Flanders: Haven van Gent, GentZelzate via Linkeroever,
graafwerken ten N van executieoord Rieme (IFBL C3.33.34),
ruigte, 17.09.1982, E. Robbrecht 2528 (BR); Haven van Gent,
GentZelzate via Linkeroever (km 7.0), bij executieoord (IFBL
C3.33.44), grote zandhopen, 16.08.1984, E. Robbrecht 2829
(BR); Gent, Dampoort, railway yard, now demolished, 10.07.2004,
F. Verloove 5728 (BR);
• Hainaut: MontignysurSambre, colonise certains terrils de
charbonnages, 05.08.1930, Culot s.n. (BR); Jamioulx, le long du
chemin de fer, terrain vague, 07.1954, J. Lebeau s.n. (BR) ; Beau
vechain, friche, 01.10.1954, L. Muyldermans 841 (BR);
• Liège: Visé, bord de la Meuse, 08.1868, E. Marchal s.n. (BR);
Vierset, bois de Faqueval, 08.1889, A. Charlet s.n. (LG); Lam
bermont, décombres, 07.1908, P. Halin s.n. (LG); Theux (Hod
bomont), 09.08.1948, J.M. Warlet 5382584 (BR); Angleur, do
maine universitaire du Sart Tilman, le long de la route axiale, bord
de la route, accotements remués, 06.07.1971, A. Lawalrée 16675
(BR); Ougrée, Sart Tilman, domaine universitaire, en bordure de
la route près de l’Institut Botanique, 13.08.1971, P. Auquier 1566
(LG); Angleur, talus en bord de route, 04.07.1973, J. Beaujean
167 (LG); Tilff, route aux abords du château de Colonster, bord
de route, 09.08.1973, J. Duvigneaud s.n. (LG); JupillesurMeuse,
bord de route, 14.09.1979, J. Beaujean 79/146 (LG); BenAhin
(IFBL G6.24.14), milieu rudéralisé, 01.09.1983, J. Rousselle
s.n. (LG); Gué (?) [BenAhin], IFBL G6.24.14, terrain vague,
01.09.1983, J. SaintenoySimon s.n. (BR);
• Limburg: Bokrijk, deux pieds dans un lieu cultivé, 28.07.1889,
H. Dupont s.n. (LG); Berlingen, bij ‘t Kasteel van Rullingen, gras
boord, 02.08.1934, E. Michiels s.n. (BR); Genk (IFBL D7.51.22),
port, 26.08.1956, A. Lawalrée 8033 (BR); Waterschei, ancien
crassier près d’un charbonnage, 09.1956, J. Lambinon s.n. (LG);
Wellen, wegboord, 18.07.1958, E. Michiels s.n. (BR); Beringen,
vaartkant, 27.08.1959, E. Jacques 3939 (BR) ; Beringen, langs
een weg aan de mijnterril, 07.1967, H.G. Rabijns 1464 (BR);
Genk, aan de spoorweg van de kolenhaven, 21.07.1967, J. Ram
meloo 310 (GENT); Genk, Kolenhaven, langs de spoorlijnen,
abondant in droge zandige vegetatie, 22.07.1967, H. Stieperaere
s.n. (GENT); Genk, terrains vagues, 17.08.1980, P. Claes s.n.
(LG); Lanaken (Neerharen), Hochter Bampd (IFBL E7.15.11),
grindbank langs de Maas, 02.10.2010, F. Verloove 8289 (BR) ;
• Luxembourg: Poncelle (Tintigny), prairie aride (sable), assez
abondant, 20.08.1945, V. d’Ansembourg 599 (BR); Buzenol,
jardin près de la voie ferrée, 08.1949, J. Duvigneaud s.n. (BR) ;
Poncelle, terrain sec, 13.09.1946, J. Legrain s.n. (BR); Torgny,
20.09.1959, A. Louette s.n. (LG);
• Namur: Lustin, bord de la voie ferrée, 04.09.1956, J. Lambinon
s.n. (BR); Andenne, dans des décombres, 1962, F. Siebertz s.n.
(LG);
• West Flanders: Nieuport, ruines, 1920, A. Navez s.n.
(BR); Houthulst, forêt, 01.07.1920, Guns s.n. (BR); Coxyde,
dunes, 10.08.1920, L. Magnel s.n. (BR); Clemskerke, pâtur
ages, 31.08.1923, A. Isaacson s.n. (BR); Westende, bord de la
route, 07.09.1924, J. Lebrun s.n. (BR); Koksijde, dunes ixées,
20.08.1928, A. Maréchal s.n. (LG); Duinbergen, 08.1931, E.
Dewildeman s.n. (BR); Middelkerke, 07.1946, J. Duvigneaud
s.n. (BR); Wenduine, 07.1946, L. Renard s.n. (LG); Ostende,
partout, 06.07.1947, A. Visé s.n. (BR); StIdesbald, dunes vois
inages des villas, 02.09.1947, V. d’Ansembourg 1025 (BR);
Oostduinkerke, 08.1948, L. Renard s.n. (LG); Groenendijk,
dunes herbeuses, 12.08.1949, A. Maréchal s.n. (LG); Koksijde,
duinen, 14.07.1952, J. Pelgrims s.n. (BR); StIdesbald, dunes ru
déralisées, 15.08.1953, J. Duvigneaud s.n. (BR); Koksijde, ten
W van de Normandie, rand weg, 17.07.1957, E. Jacques 3080
(BR) ; Coxyde (SaintIdesbald), dunes, 06.07.1958, A. Lawalrée
9991 (BR); CoxydeBains, près du moulin, dunes rudéralisées,
16.07.1960, A. Lawalrée 11078 (BR); Nieuport, dunes, 10.1961,
P. Auquier 462 (LG); Le Coq, dunes, 05.08.1962, J.E. Rogister
s.n. (BR); Nieuport, sables +/ rudéralisés, 07.1963, L. Delvosalle
s.n. (BR); SintIdesbald, dunes près de la localité balnéaire,
08.07.1965, A. Lawalrée 13112 (BR); Koksijde, dans les dunes,
très répandue partout, 07.1966, L. Dubois 1648 (BR); De Panne,
droge wegkant aan het standbeeld Leopold I, 29.07.1967, E.
Robbrecht 67.13 (BR); SaintIdesbald (sud du village), panne
rudéralisée, 09.07.1968, P. Auquier 1142 (LG); Koksijde, Doorn
panne, begroeide duinpanne, 10.07.1973, M. Lejeune 84 (BR) ;
Hoeke, Steenoven, wegberm in zware kleigrond, 15.07.1973, R.
Viane 94 (GENT); Oostduinkerke, duinen, 07.07.1974, J. Loots
571 (BR); De Panne, verkaveling “Westhoek”, zuidelijke weg
(IFBL C0.56.41), verruigde Salix repensRosa pimpinell.zoom
aan de rand van Ligustrumstruweel met Euonymus europaeus,
03.09.1977, P. Goetghebeur 2790 (GENT); De Panne (IFBL
C0.56.44), dunes, 01.11.1978, P. Sotiaux s.n. (BR); Coxyde,
dunes rudéralisées, 20.09.1979, W. Debaisieux s.n. (LG); Kok
sijde, wegkant, 23.07.1980, R. Vanquathem 110 (BR); Oost
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
40
duinkerke, Doornpanne, dépression dunaire, 14.10.1982, M.
Tanghe s.n. (BR); Coxyde, dune rudéralisée, 27.07.1983, J.
Beaujean 83/10 (LG); HeistDuinbergen, Zeegrasstraat (IFBL
B2.32.43), in verruigde duinen, braakland, 21.08.1983, J. Van
Den Haute 1455 (BR); NieuwpoortBad, duinen, 08.09.1984, J.
Loots 1066 (BR); Oostende, Fort Napoleon (C1.14.42), talus sa
blonneux en contrebas de dunes rudéralisées, 09.1991, R. Fabri
1524 (Soc. Ech. Pl. Vasc. Eur. Occ. Bassin. Médit. n°15322: BR,
LG); Diksmuide, industrieterrein (IFBL D1.23.32), droge grond,
21.06.1992, F. Verloove 286 (BR); Kortemark (IFBL D1.26.42),
spoorwegemplacement, op grintpuin, 24.06.1992, F. Verloove
311 (BR); N382 tussen Roeselare en Izegem, ter hoogte van
kanaal naar Leie te Kachtem (IFBL D2.51.13), wegberm, onder
de vangrail, 18.07.1998, F. Verloove 3184 (BR); Adinkerke, road
towards BrayDunes (IFBL D0.16.11), seadunes, near railway
track, locally abundant, 26.06.2000, F. Verloove 4624 (priv.herb.
F. Verloove); N382 RoeselareIzegem, near drivein E403 (IFBL
D2.51.11), ruderal area, levelled soil (former dumping place),
09.07.2000, F. Verloove 4626 (priv.herb. F. Verloove).
Oenothera hirsutissima
• East Flanders: Harbour of Ghent, Sside of Kluizendok (IFBL
C3.43), artiicial sandscape, few specimens, 24.06.2007, F. Ver
loove 6769 (priv. herb. F. Verloove);
• West Flanders: Nieuport, dunes, 10.1961, P. Auquier 462 (BR);
Brugge, Eside of Boudewijnkanaal, sand depot, 17.07.2004, F.
Verloove 5734, 5735 & 5736 (priv.herb. F. Verloove, BR); Brugge,
Eside of Boudewijnkanaal (IFBL C2.11.24), sandy ruderal area,
few specimens, known since several years (+ hybrids with several
other taxa), 28.06.2007, F. Verloove 6768 (priv. herb. F. Verloove);
Brugge, Eside of Boudewijnkanaal (IFBL C2.11.24), sand depos
it, 18.07.2007, F. Verloove 6863 (BR) (hybrid); Brugge, Eside of
Boudewijnkanaal (IFBL C2.11.24), sand deposit, 18.07.2007, F.
Verloove 6865 (BR) (hybrid).
• Liège: ValBenoit, décombres, 12.07.1923, A. Maréchal s.n.
(LG);
• Limburg: StTrond, bords de chemin, 05.08.1867, A. Thielens
(BR).
Oenothera oehlkersii
• East Flanders: Gent, Sneppebrug (IFBL D3.22.13), road verge,
near railway track (disturbed soil), 24.06.2007, F. Verloove 6774
(BR); Zwijnaarde (ZGent) (IFBL D3.32.24), voormalige stortp
laats langs de E40, 10.07.2007, F. Verloove 6810 (BR, LG);
• Liège: Angleur, fortiications, 07.1884, P.J. Delrez s.n. (BR);
• West Flanders: Roeselare, Bruggesteenweg (IFBL D1.48.13),
demolition area of former CocaColafactory, several specimens,
21.07.2001, F. Verloove 4994 (priv. herb. F. Verloove, BR); Rum
bekeRoeselare, former claypit Ostyn, dump, 18.07.2004, F. Ver
loove 5732 (BR); Ieper, railway yard, gravelly area, 27.07.2004,
F. Verloove 5730 (BR).
Oenothera paradoxa
• Limburg: BeringenMijn (IFBL D6.15.13), Sside of coal mine
heap, common, 08.07.2001, F. Verloove 4848 (priv. herb. F. Ver
loove); BeringenMijn (IFBL D6.15.31), coal mine heap, common,
01.07.2007, F. Verloove 6812 (priv. herb. F. Verloove).
Oenothera parvilora
• East Flanders: Vrasene, Aven Ackers (industrial area) (IFBL
C4.13.31), bare sandy area, 31.08.2014, F. Verloove 11203 (BR).
• Namur: Jambes, Materne, bord de la voie ferrée, 09.1955, J.
Lambinon s.n. (BR).
• Brabant: Dieghem, talus du chemin de fer, 1884, A. Douret s.n.
(BR); BruxellesNord, chantier de la jonction NordMidi, secteur
Botanique, 21.06.1947, A. Lawalrée 1627 (BR); Bruxelles, plaine
de l’Exposition de 1935, à 100 m du Planetarium, 08.08.1948,
E. Michel s.n. (BR); Bruxelles, chantier jonction, 19.08.1958, M.
Coûteaux 742 (BR); Bruxelles, terrain vague, 19.09.1972, A. La
walrée 17972 (BR); Halle, gare, 02.09.1978, S. Depasse 88/85
(BR);
• Hainaut: Viesville, près du canal, rudéralia, 10.08.1955, A. La
walrée 6870 (BR); BraineleComte, talus SNCB, 22.07.1957, S.
Depasse 88/26 (BR); Tertre, 05.08.1967, S. Depasse 88/69 (BR);
• Liège: JemeppesurMeuse, terril houillers du Bas Laveu, friche
en bordure d’une zone en combustion, 25.08.1984, J. Lambinon
84/B/640 (BR);
• Luxembourg: Halte de Rossart, chantier de bois, 30.08.1965, V.
d’Ansembourg 3372 (BR);
• West Flanders: Baai van Heist (OZeebrugge) (IFBL B2.32.33),
ruderal sandy area, near the sea, abundant, 08.09.2000, F. Ver
loove 4623 (priv.herb. F. Verloove).
Oenothera laciniata
Oenothera perennis
• Brabant: Wilsele, lieux incultes, 07.1893, C. Baguet s.n. (BR);
Wilsele, dépendances de l’usine Bodart, 07.1894, C. Baguet
s.n. (BR, GENT, LG); Schaarbeek, environs de la gare, 1907,
A. Isaacson s.n. (BR); Anderlecht, immondices, 05.07.1920, E.
Michel s.n. (BR);
• Liège: Juslenville, voie ferrée, 27.09.1903, M. Halin s.n. (BR);
Vovegnez, graviers, 12.09.1909, P. Halin s.n. (LG); Vovegnez,
berge de la Vesdre, 12.09.1909, M. Halin s.n. (BR); Liège, subs
pontané dans le jardin du collecteur, 1924, A. Maréchal s.n. (BR);
• West Flanders: Coxyde, entre les ils de fer barbelés au pied
du Hoogenblikker près du nouveau chemin empierré vers Oost
duinkerke, un seul pied, 28.09.1919, L. Magnel s.n. (BR).
• Antwerp: Capellenbosch, zandgrond, 20.06.1943, L. Standaert
s.n. (BR).
Oenothera indecora
• Brabant: CorbeekLoo, champ de trèles, 08.1895, C. Baguet
s.n. (BR).
Oenothera issleri
Oenothera pycnocarpa
• Hainaut: Hensies, Les Sartis (IFBL G3.31.44), coal mine heap,
12.07.2007, F. Verloove 6880 (BR) (hybrid); Blaton, sablière le
long du canal (IFBL G3.21.42), sur sable, 25.07.2007, F. Verloove
6868 (priv. herb. F. Verloove);
• Liège: Huy, terrain vague, 16.08.1956, A. Lawalrée 7784 (BR);
• Namur: Houx, taillis, 01.10.1933, R. Mosseray s.n. (BR).
• Antwerp: Harbour of Antwerp, Sside of 5th Havendok (IFBL
C4.16.33), road verge, talus, few specimens, 29.07.2007, F.
Verloove 6834 (BR); Harbour of Antwerp, SWside of Hansadok
(IFBL C4.15.23), along railway track, 1 ex., 29.07.2007, F. Ver
loove 6835 (BR); Waaslandhaven, Kallo, SintJansweg (IFBL
C4.14.12), ruderal sandy area, in front of Sagrex sand storage,
very numerous, 27.07.2014, F. Verloove 10925 (BR);
• Hainaut: Gilly, au NO du cimetière (IFBL G4.48.11), le long
d’une route, creusée dans les lancs d’un ancien terril, peuple
ment très important (1000 pieds au moins), 07.2007, M. Lannoy
s.n. (priv. herb. F. Verloove 6854, BR, LG);
• Liège: QueueduBois, parking usine Aldi, 04.09.1992, J. Beau
jean 92/86 (BR);
• West Flanders: Brugge, Pathoekeweg, W of Nijverheidsdok
(IFBL C2.21.22), waste ground (demolition area), locally abun
dant, 08.08.2007, F. Verloove 6852 (priv. herb. F. Verloove, BR).
Oenothera oakesiana
Oenothera rosea
Oenothera nuda
• Brabant: Bruxelles, travaux de
26.06.1945, A. Lawalrée 1303 (BR);
la
jonction
NordMidi,
• Hainaut: Ath (Tournai), s.d., Ronday s.n. (BR); Ath, 1874, Ron
day s.n. (BR);
K. Rostański and F. Verloove, The genus Oenothera in Belgium [Dumortiera 106/2015: 12-42]
41
• Liège: Vervian, Leodis, sponte in oleraciis, s.d., R. Courtois s.n.
(LG).
Oenothera royfraseri
• Antwerp: Harbour of Antwerp, angle of Amerika and Albertdok
(IFBL C4.16.41), ruderal area, bare and gravelly, very common,
29.07.2007, F. Verloove 6833 (priv. herb. F. Verloove).
Oenothera rubricalyx
• West Flanders: Rekkem, LARtransportzone (IFBL E2.41.32),
grassland, several populations, 11.07.2014, F. Verloove 10892
(BR).
Oenothera rubricaulis
• Antwerp: […], bords de chemin, 07.1887, P. Halin s.n. (BR);
Harbour of Antwerp, Sside of Delwaidedok, railway track,
15.08.2004, F. Verloove 5748 (BR);
• Brabant: Aarschot, woest terrein achter het station, 06.07.1939,
E. Michiels s.n. (BR);
• East Flanders: Haven van Gent, Meulestedekaai, aan Meulest
edebrug (IFBL D3.12.22), kanaalberm met naakte min of meer
dichtgeslagen zandbodem, massale populatie, 05.07.1985, E.
Robbrecht 2929 (BR); Gent, spoorwegberm tussen Ringvaart
en Watersportbaan (IFBL D3.21.24 + D3.22.13), grindrijk terrein,
zeer talrijk, 09.10.2005, F. Verloove 6230 (BR); GentWest, Ring
vaart x railway track (IFBL D3.21.24), railway track, formerly a
massive population, now partly demolished after infrastructural
works, 24.06.2007, F. Verloove 6775 (BR); Harbour of Ghent,
Singel (E of Grootdok) (IFBL D3.13.11), ruderal area, abundant,
24.06.2007, F. Verloove 6779 (BR);
• West Flanders: Heist, Baai van Heist (IFBL B2.32.33), sea
dunes, +/ ruderalized, 18.07.2007, F. Verloove 6861 (BR).
Oenothera victorinii
• Antwerp: Harbour of Antwerp, right bank of river Schelde, SW
of 5th Havendok (IFBL C4.15.43), worked up road verge, several
specimens, 29.07.2007, F. Verloove 6830 (BR);
• Brabant: Lasne, rue du Double Ecot, carrière abandonnée,
12.07.1992, F. Billiet & B. Jadin 2331 (BR);
• East Flanders: Gentbrugge, Hundelgemsesteenweg (IFBL
D3.23.33), braakgrond, nabij de oevers van de Schelde,
02.08.1998, F. Verloove 3357 (BR); Gent, Hundelgemsesteenweg
near river Schelde (IFBL D3.23.33), ruderal area, 10.07.2004, F.
Verloove 5729 (BR);
• West Flanders: N382 RoeselareIzegem, near drivein E403
(IFBL D2.51.11), ruderal area, levelled soil (former dumping
place), several specimens, along with O. glazioviana, 09.07.2000,
F. Verloove 4625 (priv. herb. F. Verloove); Roeselare, N382 at
drivein E403 (A17) (IFBL D2.51.11), levelled soil, persistent, irst
recorded in 2000, 13.07.2001, F. Verloove 4996 (LG).
Oenothera villosa
• West Flanders: Brugge, Eside of Boudewijnkanaal (IFBL
C2.11.42), sand deposit Hanson, one specimen, 18.07.2007,
F. Verloove 6816 (priv. herb. F. Verloove); Brugge, Eside of
Boudewijnkanaal (IFBL C2.11.42), sand storage, +/ 5 specimens,
21.07.2014, F. Verloove 10917 (priv. herb. F. Verloove, BR).
Oenothera wratislaviensis
• Antwerp: Harbour of Antwerp, Sside of 5th Havendok (IFBL
C4.16.33), road verge, talus, abundant, 29.07.2007, F. Verloove
6832 (priv. herb. F. Verloove);
• Hainaut: Hensies, Les Sartis (IFBL G3.31.44), coalmine heap,
04.07.2004, F. Verloove 5724 (priv. herb. F. Verloove);
• West Flanders: Heist, Baai van Heist, sandy grassland near the
sea, 17.07.2004, F. Verloove 5733 (priv. herb. F. Verloove, BR).
Oenothera rubricauloides
• East Flanders: GentWest, Ringvaart x railway track (IFBL
D3.21.24), railway track, 24.06.2007, F. Verloove 6776 (BR); Zwi
jnaarde (ZGent) (IFBL D3.32.24), former dump site, 10.07.2007,
F. Verloove 6811 (priv. herb. F. Verloove).
Oenothera speciosa
• Hainaut: SaintGhislain, s.d., Van Bastelaer s.n. (BR).
Oenothera stricta
• Antwerp: Anvers, fortiications près de la porte de Wilrijk, 23.05
& 23.06.1878, H. Vandenbrouck s.n. (BR); Anvers, fortiications,
station abondante, 23.06.1878, H. Vandenbrouck s.n. (BR); An
vers, fortiications, 08.1882, H. Vandenbrouck s.n. (BR); Anvers,
remparts, 31.08.1882, J. Hennen s.n. (BR); Anvers, fortiications,
07.1884, P.J. Delrez 451 (LG); Anvers, remparts, 29.07.1884, J.
Hennen s.n. (BR); Anvers, remparts, 07.1887, J. Hennen s.n.
(BR, LG); Anvers, fortiications à Berchem, 07.1900, R. Godding
s.n. (BR);
• Brabant: Wilsele, lieux incultes, 07.1893, C. Baguet s.n. (BR);
• Hainaut: SaintAmandlezFleurus (IFBL G5.21), 06.1883,
Grosse s.n. (BR); Bellecourt (IFBL G4.23), sur les talus du chemin
de fer, plante naturalisée, <1951, A. Briart s.n. (BR);
• Liège: Angleur, friche en bordure d’une zone en combustion,
25.08.1984, J. Lambinon 84/B/640 (LG);
• West Flanders: La Panne, 05.06.1881, W. Rouseau s.n. (BR).
Oenothera subterminalis
• Hainaut: Châtelineau, Châtelet, terril, face sud SW, 11.06.1979,
JC Moniquet s.n. (BR);
• West Flanders: Heist, Baai van Heist, seadunes, +/ ruderal
ized, 02.08.2004, F. Verloove 5738 (BR).
Non-stabilized hybrids
Cf. Oenothera biennis × O. delexa
• Antwerp: Harbour of Antwerp, Vosseschijnstraat, E of 2e Ha
vendok (IFBL C4.16.42), railway track, 29.07.2007, F. Verloove
6871 (BR);
• East Flanders: Gent, Leon Tertzweillaan (IFBL D3.23.13),
omgewoelde, eroderende, zandige, grintrijke spoorwegberm, pio
niervegetatie met veel adventieven, 06.09.1977, P. Goetghebeur
2802 (GENT); Handzame (IFBL D1.26.33), hoop gestorte grond
langs wegkant, 28.08.1999, H. Ruysseveldt 2389 (BR);
• West Flanders: Brugge, harbour, E of Groot Handelsdok (IFBL
C2.22.11), former railway track + ruderal area, 18.07.2007, F. Ver
loove 6864 (BR).
Oenothera biennis × O. glazioviana
• Luxembourg: Buzenol, coupe dans le bois de la Ferme de
Buzenol, le long du ruisseau, 08.1949, J. Duvigneaud s.n. (BR).
Cf. Oenothera delexa × O. fallax
• West Flanders: Brugge, Eside of Boudewijnkanaal (IFBL
C2.11.24), sand deposit, 08.08.2007, F. Verloove 6857 (BR).
Cf. Oenothera depressa × O. delexa
• East Flanders: Zwijnaarde (ZGent) (IFBL D3.32.24), entrance
of former dump area, one specimen, 10.07.2007, F. Verloove
6808 (priv. herb. F. Verloove);
• West Flanders: Zeebrugge (portarea), N of Verbindings
dok (IFBL B2.41.42), artiicial sandscape near Tropicanaplant,
18.07.2007, F. Verloove 6858 (BR); ZeebruggeStrand, Wside of
port (IFBL B2.41.24), sandy ruderal soil, near railway station, 2
ex., 08.08.2007, F. Verloove 6851 (BR).
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