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1869
Danihelka & al. • Reject Viola montana and V. persicifoliaTA XO N 59 (6) • December 2010: 1869–1878
INTRODUCTION
Viola subsect. Rostratae (Kupffer) W. Becker (= V. sect.
Trigonocarpea Godron) is represented in Europe by five aro-
sulate species, often referred to as V. “ser. Arosulatae Borbás”
(Van den Hof & al., 2008). Viola canina L. (2n = 40), the com-
monest one, has a wide distribution range reaching from the
Iberian Peninsula in the west to Lake Baikal in the east. It is
extraordinarily morphologically variable, and its intraspecific
classification is still in dispute. Viola lactea Sm. (2n = 58), in
contrast, is a strongly oceanic species confined to the British
Isles, the northern parts of the Iberian Peninsula, western France,
and Belgium. The three remaining species, in recent literature
known as V. elatior Fr. (2n = 40), V. p umila Chaix (2n = 40), and
V. stag nina Kit. ex Schult. (or V. persicifolia Schreb.; 2n = 20),
have wide distribution ranges reaching from the British Isles and
eastern France eastwards to western or central Siberia. In central
Europe they are often confined to the floodplains of the large
lowland rivers. The taxonomy and ecology of the three f lood-
plain violets in Central Europe was recently reviewed by Eck-
stein & al. (2006). In the course of our studies, we have encoun-
tered nomenclatural difficulties that will be dealt with herein.
VIOLA MONTANA
Herbarium specimens of V. e l a t ior collected in the late
18th and early 19th centuries have been frequently identified as
V. montana L. (Sp. Pl. 2: 935. 1753), which is in c onf lict with t he
prevailing current use of this Linnaean name for certain mor-
photypes of V. canina. These different interpretations can be
traced back to a putative redefinition of V. m o n t a n a in the sec-
ond edition of Flora suecica (Linnaeus, 1755) and subsequently
in the second edition of Species plantarum. The use of the name
V. montana has been repeatedly discussed. Some authors have
suggested that the name V. m ontana originally referred mainly
to the species currently known as V. elatior (e.g., Neilreich,
1859: 774; Borbás, 1892; Wilmott, 1916; Lindberg, 1958). Ni-
kitin (1988) reviewed the nomenclatural history of V. m ontana
and proposed a lectotype (Herb. Linn. No. 1052.13, LINN)
referable to V. elatior. This lectotypification is in accordance
with the protologue and should not be overturned. However,
only a few authors apart from Nikitin seem to have accepted
its consequences and replaced V. elatior by V. montana (e.g.,
Chen & al., 2007), while many other national checklists and
floras published after 1988 preferred nomenclatural stability
and clarity to correctness, and continued using V. elat ior. The
replacement of a well-established name by another name that
was only rarely used in its original sense after the 1820s is
undesirable and would destabilise nomenclature. Therefore we
have decided to propose V. montana for rejection, as already
announced by Kirschner & Skalický (1989).
Viola montana L., Sp. Pl. 2: 935. 1753, nom. utique rej. prop. (Van
den Hof & al. in Taxon 59: 1900 – this issue). Ind. loc.: “Hab-
itat in Alpibus Lapponiae, Austriae, Baldo.” – Lectotype
Viola montana and V. persicifolia (Violaceae): Two names
to be rejected
Jiří Danihelka,1,2 Kevin van den Hof,3 Thomas Marcussen4 & Bengt Jonsell5
1 Department of Botany & Zoology, Masaryk University, Kotlářská 2, 611 37 Brno, Czech Republic
2 Institute of Botany, Department of Vegetation Ecology, Academy of Sciences of the Czech Republic, Lidická 25/27,
602 00 Brno, Czech Republic
3 Netherlands Centre for Biodiversity Naturalis – National Herbarium of the Netherlands, Leiden University branch,
P.O. Box 9514, 2300 RA Leiden, The Netherlands
4 Department of Plant and Environmental Sciences, University of Gothenburg, Box 461, 405 30 Göteborg, Sweden
5 Konsumvägen 20B, 756 45 Uppsala, Sweden
Author for correspondence: Jiří Danihelka, danihel@sci.muni.cz
Abstract
The taxonomic and nomenclatural histories of Viola elatior Fr. (1828), V. pumi l a Chaix (1785) and V. stag n i n a Kit.
ex Schult. (1814) in central and western Europe are discussed. The names V. s t agnina and V. e l a t ior are lectotypified with
specimens corresponding to the current use of these names. The neglected lectotypification in 1988 of V. m o ntana L. (1753)
with a specimen referable to V. e latio r is brief ly reviewed. The name V. p ersic ifol ia Schreb. (1771), used in some floras instead
of V. s tagnin a , is analysed in detail, and we conclude that this name should be interpreted as referring also to V. e latio r. In
addition the types of V. hornemanniana Schult. (1819) and V. stipu lacea Hartm. (1820) are conspecific with V. elat ior. The use
of V. persi c ifol ia and V. mont a n a, representing earlier legitimate names for the species widely known as V. ela t i o r, has been
notoriously confused for t wo centuries, and we herein recommend these two names for rejection in order to assure nomenclat ural
clarity and stability. For similar reasons, we recommend conservation of V. e la tior against V. hornemanniana and V. st ip ul acea .
Keywor ds
nomenclature; typification; Viola elatior; V. stag nina ; Viola subsect. Rostratae
1870
TA XO N 59 (6) • December 2010: 1869–1878Danihelka & al. • Reject Viola montana and V. persicifolia
(vide Nikitin in Bot. Zhurn. 73: 1541. 1988): “Viola 10 /
montana” (Herb. Linn. No. 1052.13, LINN, vide http://www
.l i nnean-online.or g /11110 / ).
VIOLA PERSICIFOLIA
Taxonomic hi sto ry. —
The name Viola persicifolia was
published by Schreber (1771: [163]) with a reference to a descrip-
tion in a pre-Linnaean flora of Leipzig (Boehmer, 1750) (Fig.
1). However, this publication remained neglected for long or the
name was considered not validly published due to the putative
lack of description. For these reasons or due to contemporary
nomenclatural practice, the name was ascribed to later authors,
initially to Roth (1789: 271; e.g., Schultes 1814: 427; Mertens &
Koch, 1826: 268; Reichenbach 1823: 88) and later to Schkuhr
(1803: 187; e.g., Reichenbach 1832: 708, 1839–1840: 45). Even
now, more than two centuries after its publication, the name
V. persicifolia is subject to controversy: in some floras, mainly
those from western Europe, it has been used for the species
otherwise known as V. stagnina (Valentine & al., 1968; Guino-
chet & Vilmorin, 1982; Stace, 1997; Haeupler & Wisskirchen,
1998; Elven, 2005; Van der Meijden, 2005), while others argued
that it refers to the species known as V. e l a t i o r Fr. and should
be proposed for rejection in the terms of the Code to prevent
further confusion (Mansfeld, 1939; Hylander, 1945; Rauschert,
1973; Kirschner & Skalický, 1990; Eckstein & al., 2006).
A simplified survey of taxonomic and nomenclatural treat-
ments of Viola elatior and its relatives in Central European
floras and some other monographs is given in Table 1. As can
be seen from the table, the interpretation of the name V. p e r -
sicifolia has been connected with difficulties from the very
begin ning. Bot h Roth (1789: 270 –272) and Schkuh r (1803: 187 )
recognised two arosulate Viola species from th is g roup, V. per-
sicifolia and V. m o n t a n a, the latter in its original concept and
including at least partly V. elatior. Still, Schkuhr (l.c.) clearly
Fig. .
Validating description of Viola persicifolia Schreb. (Boehmer,
1750).
Tab le .
Taxonomy and nomenclature of floodplain violets and V. lactea in major Central European floras and monographs between 1771 and 1917
Currently accepted species
Author Viola elatior Viola stagnina Viola pumila Viola lactea
Schreber, 1771 V. persicifolia
Roth, 1789 V. persicifolia (V. montana)
Willdenow, 1798 V. montana
Schkuhr, 1803 V. persicifolia (V. montana)
Schultes, 1814 V. persicifolia “Roth”
V. montana?
V. stagnina V. lactea
Reichenbach, 1823 V. persicifolia “Roth” V. lactea
Mertens & Koch, 1826 V. persicifolia “Roth” V. stagnina V. pratensis V. lactea (syn. V. lancifolia)
Fries, 1828 V. elatior V. “persicaefolia” Schreber V. lactea
V. l. var. humilior, V. l. var. pratensis
Reichenbach, 1832 V. persicifolia “Schkuhr” V. lactea
(different from V. lancifolia)
Koch, 1836 V. elatior V. stagnina V. pratensis V. lactea (syn. V. lancifolia)
Reichenbach, 1839–1840 V. persicifolia “Schkuhr” V. lactea
(incl. V. l. var. humilior but different from V. lancifolia)
Koch, 1843 V. elatior V. stagnina V. pratensis
Uechtritz, 1871 V. elatior V. stagnina V. pumila V. lactea
Borbás, 1892 V. montana V. persicifolia “Roth” V. pumila V. lactea (syn. V. lancifolia)
Becker, 1910 V. elatior V. stagnina V. pumila V. lactea
Becker, 1917 V. elatior V. persicifolia “Roth” V. pumila V. canina subsp. lactea
1871
Danihelka & al. • Reject Viola montana and V. persicifoliaTA XO N 59 (6) • December 2010: 1869–1878
expressed his uncertainty about their delimitation (see also p. 6
of Nachtrag), and his contemporary Willdenow (1798: 1165)
did not consider them different at all (though he referred to the
second edition of Flora suecica ; Linnaeus, 1755) and treated
them collectively as V. montana. However, it is probable that the
description of another arosulate violet, V. lactea (Smith, 1798:
445), provided an incentive for Willdenow to later recognise
more than one species in this group. To our knowledge this
was never published by Willdenow himself, who died in 1812,
but his herbarium (Röpert, 2000–) contains one folder labelled
V. montana and a second labelled V. lactea. His delimitation
of the two, however, makes no sense in our view. The former
folder contains two sheets of V. ela t ior and one of V. p u mil a ,
and the latter folder two sheets of V. elatior, three of V. p um ila,
and two of V. stagnina.
Schultes (1814: 425–427) was the first to recognise more
than two species of floodplain violets. His three species were
V. lacte a, based on a specimen collected by P. Kitaibel in Hun-
gary and whose description roughly corresponds to V. p umil a
(represented by two Kitaibel specimens in herbarium Will-
denow; see Röpert, 2000–), V. persicifolia “Roth”, referable
to V. elatior, and V. s t a g n i n a that he described as a new spe-
cies based on a specimen collected by P. Kitaibel in Croatia.
In addition to these three species, Schultes (l.c.) further kept
V. montana (with a question mark and only general information
about its distribution and habitat); the fifth species, V. lanci-
folia, reported from the surroundings of Berežany in south-
western Ukraine, is difficult to interpret but it may refer to a
specimen of V. canina.
Nine years later, Reichenbach (1823) paid a great deal of
attention to Viola. B ased o n Wahle nberg’s opi nion, he c oined the
concept of V. montana as redefin ed by Lin naeus (1755: 305–30 6)
in the second edition of Flora suecica, i.e., as a species similar
to V. c a n in a (Reichenbach, 1823: 84–85). The taxonomy of the
floodplain violets was discussed on pp. 86–88 as comments on
pls. XCIX (Viola lactea. Sm.) and C (Viola persicifolia. Roth.)
(Fig. 2). Reichenbach recognised two species, V. lactea, with
all leaves glabrous and oblong-lanceolate (sometimes ovate-
or cordate-lanceolate), and V. persicifolia (ascribed t o Roth),
with ovate-lanceolate leaves, pubescent when young. The latter
clearly corresponds to V. elatior; a chasmogamously flowering
specimen (figure 209) and upper part of a fruiting specimen with
capsules from cleistogamous flowers (figure 210) were drawn
after plants collected in Leipzig (“Specimina Lipsiae legi in loco
Ruppiano.”; p. 88). Here and in the synonymy Reichenbach ex-
plicitly refers to a violet treated in two pre-Linnean f loras as oc-
curring near Leipzig (Ruppius, 1726: 233, 1745: 289; Boehmer,
1750: 190). He also explained that Roth, as not being familiar
with this violet, created a new name based on Ruppius’s phrase
name (Reichenbach 1823: 88). Also V. l ac te a in Pl ate XCI X was
drawn after a plant collected in Leipzig (“Lipsiae legi.”; p. 86).
In his comments Reichenbach (p. 86) stated that it is a wide-
spread species collected from a major part of Europe, but at the
same time much confused. Variation in leaf and stipule shape
was, according to Reichenbach, merely plastic responses to
differences in humidity and soil conditions, and thus not worth
recognition. To prove his point, he drew along with the whole
plant a series of laminas and stipules as figures c–n of Plate
XCIX. In our opinion, while the whole plant is clearly referable
to V. s t a g n i n a , the detailed leaf and stipule drawings belong to
V. pumila. Reichenbach also associated V. pu mila “Vill.” with
his V. lactea, but with some degree of uncertainty (see the syn-
onymy in p. 86), while the choice of the younger name V. l ac te a,
based on British plants, was supported by comparison of his
specimens with the drawing in Smith (1798: pl. 445).
The second volume of Röhlings Deutschlands Flora
(Mertens & Koch, 1826) brought important novelties. Its
authors accepted V. persicifolia “Roth” as circumscribed by
Reichenbach (1823) but almost excluded V. lactea Smith (with
V. lancifolia Thore, Essai Chloris: 357. 1803, as synonym; pp.
264–265) from the f lora of Germany, referring only to a sin-
gle specimen collected by C.F.W. Wallroth near Wendelstein
in Thuringia. F.K. Mertens and W.D.J. Koch were the first to
recognise that Reichenbach’s V. l a c t e a consisted of two spe-
cies, V. s t a g nina and the newly described V. p ratensis Mert. &
W.D.J. Koch (Deutschl. Fl. 2: 267. 1826), i.e., V. p umil a . The
characters given in the descriptions (pp. 265–268) delimitate
the two from each other as well as from V. canina s.l. They
further discussed the appearance of plants with capsules and
cleistogamous flowers and also noted, in the synonymy of
V. stag nina (p. 266), that plants identified as V. persicifolia by
J.C.D. Schreber in his herbarium correspond to V. s t a g n i n a .
Two years later, Fries (1828: 274–279) also accepted three
species using similar concepts, but under completely different
names. First he argued (p. 275) that the plant found by Ruppius
(1745: 289) near Leipzig was V. st a g nina rather than V. elatior,
referring also to the description in Haller (1768: 241–242, spe-
cies no. 562), and that the description provided by Roth (1789:
270–272) would apply better to V. stagnina than to V. e l a t i o r
(treat ed by Rot h under V. montana). For these reasons, Fries used
the name V. persicifolia (“V. persicaefolia”) for V. stagnina and
proposed a new name, V. elat ior, to replace V. persicifolia as used
es pecia lly by Roth (1789) a nd afte rwa rds . In contra st to Merte ns
& Koch (1826), he kept the name V. lactea (instead of V. praten-
sis) for V. pu mila, based on the opinion of O. Swartz, who had
declared Fries’s specimens to be the genuine V. lactea of Smith.
Reichenbach (1832: 707–708) may be understood as a po-
lemic with Mertens & Koch (1826). He insisted that only two
species of floodplain violets should be recognised in Germany,
i.e., V. la c te a, consisting of our V. pu mila and V. s t a g n i n a , and
V. persicifolia “Schk.”, corresponding to V. ela tior. He further
argued that “V. stagnina Kit. nil est nisi status post f lorescen-
tiam” of his V. lactea. However, he accepted Fries’s opinion that
“V. persicaefolia” of Ruppius, Schreber, and Roth be conspe-
cific with his V. lactea and not with V. persicifolia as described
and drawn by Schkuhr (1803: 187 & pl. CCLXIX). Instead of
accepting V. elatior as correct name, he kept V. persicifolia
and ascribed it to Schkuhr (p. 708). This may have been in
accordance with contemporary nomenclatural practice but it
only further deepened the nomenclatural confusion.
In the first edition of the Synopsis, Koch (1836: 85–86)
kept the concept of the three species as proposed ten years
earlier (Mertens & Koch, 1826) but, following Fries (1828), he
replaced the name V. persicifolia with V. e la tior. The diagnoses
1872
TA XO N 59 (6) • December 2010: 1869–1878Danihelka & al. • Reject Viola montana and V. persicifolia
were precise and distinguished well among the three. Referring
to plate XCIX in Reichenbach (1823), Koch assigned the main
figure to V. s t a g n i n a but the leaf drawings c–f to his V. pra-
tensis. He further definitely excluded V. l actea (as V. lancifolia)
from the flora of Germany.
Reichenbach returned to the topic with two plates (Re-
ichenbach 1838–1839: pl. XVI & XVII) and a long accompany-
ing text (Reichenbach, 1839–1840: 38–48). He was very critical
about the treatment of floodplain violets in the Synopsis (Koch,
1836) and used strong words bordering to personal attacks. Like
in his earlier work, Flora germanica excursoria (Reichenbach,
1832: 707–708), he recognised only two species, V. persicifolia
“Schkuhr” and V. lactea. The latter consisted of populations
classified now as V. p u m ila and V. s t a g ni n a , and Reichenbach
considered them one taxon conspecific with the British popu-
lations of V. lacte a (but different from V. lancifol i a described
Fig. .
Plate C (Reichenbach,
1823) depicting Viola persicifo-
lia “Roth” drawn by Reichen-
bach himself after plants
collected near Funkenburg in
Leipzig.
1873
Danihelka & al. • Reject Viola montana and V. persicifoliaTA XO N 59 (6) • December 2010: 1869–1878
from nor thwestern Fra nce). He repeated his arguments again st
the species rank of V. stagnina and V. praten si s, at the same
time recognizing as a separate taxon 4507b V. lactea var. hu-
milior Fr. (with V. pratensis in synonymy); the corresponding
figure in pl. XVII (labelled as 4507.b. pratensis M.K) repre-
sents a typical V. p u mi l a. However, Koch (1843) apparently
ignored Reichenbach’s strong criticism and only added a few
reasons for not using the names V. lactea and V. persicifolia.
Uechtritz (1871) adopted the same taxonomy as proposed
by Koch (1836, 1843). However, he was probably among the
first to replace V. pratensis by the earlier name V. pumila. He
interpreted V. persicifolia as originally referring to V. stag-
nina but recommended to “remove” this notoriously misap-
plied name. Borbás (1892), adopting the same classification,
paid considerable attention to nomenclature: he suggested a
return to the original Linnaean concept of V. m o n t a n a and
recommended the use of this name instead of V. elatior, and,
based on the description by Roth (1789: 271–272), he replaced
the name V. stag nina with V. persicifolia “Roth”.
Becker (1910) accepted the taxonomy developed by his
immediate predecessors but preferred to use the unambiguous
name V. s t ag n i n a instead of V. persicifolia. However, in his
monograph on Asian and Australian species (Becker, 1917),
he reintroduced V. persicifolia “Roth” to replace V. s t a g ni n a .
Becker’s last important monograph seems to have inf luenced
the interpretation of the name V. persicifolia until the present.
Becker’s reasoning reads as follows (p. 394):1
Ich habe für diese Art die Bezeichnung „V. persicifolia
Roth“ wieder verwandt, da es keinem Zweifel unterliegt,
daß Roth unter diesem Namen obige [= V. stag nina]
Pflanze verstanden hat. Roth hat die Art nach der Phrase
Ruppius’ in der Fl. Jenens. (1726) p. 233 (Hallers Ausg. a.
1745, p. 289) benannt: „Viola palustris, angustis Persicae
foliis mucronatis et serratis, nondum descripta“. Rupp
gibt seine Art von Sumpfwiesen „bei Leipzig, nicht weit
von der Funkenburg“ an. Roth zitiert nicht nur die Rupp-
sche Pflanze, sondern auch Boehmer Fl. Lipsiae indigena
(1750) p. 190 n. 456: […], welcher auch als Standort die
„Funkenburg“ angibt und gut beschreibt. Hier kam die
Art, die von Rupp l. c. als häufig bezeichnet wird, noch
zu Reichenbachs Zeiten vor (Rchb. Dtschl. Flora nr. 4507,
a. 1839, p. 39).
1 I used again the name “V. persi c ifol ia Roth” for this species as
there is no doubt that Roth understood under this name the above
mentioned plant [= V. stagni n a]. Roth named this species after
Ruppius’s phrase-name in the Fl. Jenens. (1726) p. 233 (Haller’s ed.
a. 1745, p. 289): “Viola palustris, angustis Persicae foliis mucronatis
et serratis, nondum descripta”. Ruppius reports his species from
swamp meadows near Leipzig, not far f rom Funkenburg. Roth cites
not only Ruppius’s plant, but also Boehmer Fl. Lipsiae indigena
(1750) p. 190 n. 456: […]; Böhmer also reports and describes well
the “Funkenburg” site. Here the species, which was indicated
by Ruppius l.c. as frequent, still occurred in Reichenbach’s time
(Rchb. Dtschl. Flora nr. 4507, a. 1839, p. 39).
However convincing this may sound, it is, in fact, incor-
rect. Although it is true that Reichenbach (1839–1840: 39) dis-
cussed the identity of the Funkenburg violet and attributed it
to V. lactea (i.e., V. stagnina or V. pumila), the re is no evidence
that any of the plants depicted as “4507. Viola lactea Smith”
in Icones (Reichenbach, 1839–1840: pls. XVI & XVII) were
collected near Funkenburg. Actually, Reichenbach published
a drawing made after the Funkenburg plant 16 years earlier in
the Plantae criticae (Reichenbach, 1823: pl. C, figs. 209 & 210
and p. 88; see also above), and the drawings unambiguously
represent V. e l a t io r. Already Gerstlauer (1943: 46) pointed
to this error but this publication has been neglected by some
botanists.
Nomenclatural analysis. —
The name Viola persicifolia
was published with an extremely short protologue (Schreber,
1771: [163]). It consisted solely of a number (456), in the right
column, representing a reference to species 456, Viola caule
erecto, foliis ovato lanceolatis, serratis, in Flora Lipsiae indi-
gena (Boehmer, 1750: 190; cf. Schreber, 1771: [172]). This has
to be considered indirect reference to a previously published
description as described in Art. 32.6, required for valid publi-
cation of a name by Art. 32.1.(c) of the ICBN (McNeill & al.,
2006). The treatment of the species in Boehmer (Fig. 1) consists
of a phra se name, a nother phrase na me use d in the thi rd edition
of an earlier flora of Jena and its surroundings (Ruppius, 1745:
289), locality information, and a description. As no herbarium
specimens collected by G.R. Boehmer or H.B. Ruppius are
known to be extant (Stafleu & Cowan, 1976, 1983), those four
elements are the only base for the interpretation of the name. In
principle, V. persicifolia could refer to any or all of V. elatior,
V. pumila, and V. stag ni na because all three are known to have
occurred in the surroundings of Leipzig at least until the 1850s
(Reichenbach, 1823; Petermann, 1838; Hardtke & Ihl, 2000;
P. Gutte, in litt.; also numerous specimens from herbarium
Reichenbach fil. at W).
Boehmer’s description is rather ambiguous and contains
little i nformation. “St ipulae duae m inore s” may be interpreted
as a character of V. st a g nina or merely as a comparison to the
size of the lamina and petiole. The erect stem is typical of
V. elatior, while pale corolla (in comparison with V. odorata)
applies better to V. e l a t i o r and V. stagnina than to V. pu m ila.
However, the fact that the species was cultivated in gardens (as
ornamental plant?) applies best to V. elat ior, which is relatively
undemanding in cultivation and certainly has an interesting
habit and some decorative value, and probably also to V. pu mila.
The treatment in Ruppius (1745: 289) is even shorter—“Viola
palustris, angustis Persicae foliis mucronatis, & serratis, non-
dum descripta. Ist häuffig auf sumpfigten Wiesen bey Leipzig,
nicht weit von der Funcken-Burg, floret Aprili.”—and does not
offer much additional information. In general, the informative
value of such old diagnoses should not be overestimated: in this
case, the phrase names from Ruppius and Boehmer are also
cited in the validating description of a species in the V. c a n i n a
group, V. ruppii All. (Haller, 1768: 241; Dandy, 1970; see also
Fries, 1828: 274). Further, Haller (l.c., species 562), editor of
the third edition of Ruppius’s flora (Ruppius, 1745), mentioned
that he collected it in Jena, in Suevia [= Swabia/Schwaben,
1874
TA XO N 59 (6) • December 2010: 1869–1878Danihelka & al. • Reject Viola montana and V. persicifolia
Germany], and not far from Scaphusia [= Schaffhausen], but
not in Leipzig. His gatherings, now preserved at P, correspond
to neither V. stagnina nor V. elatior, but to V. r uppii as under-
stood today, i.e., a taxon close to or conspecific with V. canina
(Kirschner & Skalický, 1989). This indicates that he had him-
self not seen the Funkenburg violet. In contrast to some other
violet species in Ruppius’s flora, he did not add any comments
behind the species treatment adopted from the second edition,
which further supports this assumption.
Both Boehmer (1750: 190) and Ruppius (1745: 289) re-
ferred to the same site, variously spelled as Funckenburg bei
Gonnewitz or Funcken-Burg, now part of the city Leipzig and
not far from its centre. Adjacent to Funkenburg, hard-wood
forest (Leipziger Auenwald) and wet meadows were found in
the 19th century. Leipzig floras from this period (P. Gutte, in
litt.; e.g., Petermann, 1838: 192–193; see Reichenbach 1823:
88) reported only V. e l a t ior from this site but not V. pumila
or V. st a g nina. The former presence of V. elatior at this site
is confirmed by an undated specimen from the herbarium
Reichenbach fil. “Funkenburg Lips.” (sine coll.) now deposited
at W as no 1889/305915 (Fig. 3). Still, we cannot rule out that
also V. pu mila and/or V. st ag nina occurred there as well, but we
have not seen any corresponding specimens. The probability
that f urther gatherings from Funkenburg will be discovered is
very low because the Leipzig university herbarium was com-
pletely destroyed by fire during World War II (P. Gutte, in litt.).
The fact that (only) V. e l a t i o r had been known from Funken-
burg was used as base for the interpretation of V. persicifolia
by Reichenbach (1823: 88) and later by Gerstlauer (1943) and
Rauschert (1973). In contrast, Fries (1828: 275) argued that the
Funkenburg violet was V. s t ag n i n a because Ruppius (1726:
233) considered it as not yet described (“nondum descripta”),
whereas V. ela ti or had been repeatedly described and illustrated
by early authors (“planta tum temporis notissima, in quovis li-
bro picta”). Petermann (1838: 192) also concluded that Ruppius
had V. stag nina in mind because of its “frequent” occurrence
in wet meadows.
In our opinion, there is one circumstance neglected before:
Ruppius (1726, 1745), Boehmer (1750) and Schreber (1771) all
recognised only one species of floodplain violets in spite of the
fact that three species grew around the contemporary Leipzig.
From this point of view speculations about which species these
early authors had in mind are less important. Further, there are
reports (Reichenbach, 1839–1840: 46) that plants identified
by Schreber as V. persicifolia are referable both to V. stagnina
and to V. elatior. A specimen of V. elatior collected in 1808
in the botanic garden in Erlangen (ex herb. W.D.J. Koch, now
deposited at ER), where J.C.D. Schreber was director until 1810,
was identified as “V. persicifolia Roth”. However, when we
investigated the material of the Schreber herbarium deposited
at M, we found that all collections identified as V. persicifo-
lia can be considered V. stagnina. This corresponds to what
Mertens & Koch (1826: 266) and Fries (1828: 275) reported.
Also Schweigger (1804: 58), disciple of J.C.D. Schreber, prob-
ably used the name V. persicifolia when referring to V. stag-
nina (see Koch 1843: 93); he accepted the phrase name from
Boehmer (1750) and added: “Pro varietate violae montanae
habetur.” In contrast, the specimens of V. ela tior from the her-
barium Schreber (now at M) were identified as V. montana,
V. canadensis or “V. s i b ir i c a ?”.
The first botanist who clearly linked the name V. persicifo-
lia to V. elatior was Schkuhr (1803: 187 & pl. CCLXIX). He was
later followed by Schultes (1814) and especially Reichenbach
(1823), who published an illustration based on plants from the
Funkenburg site and clearly distinguished between V. persici-
folia (= V. e l a t ior) and V. lactea (= V. pumila and V. stagnina).
These descriptions and plates may be considered informal
emendations and tradition to follow. This point of view was
already presented by Neilreich (1859: 773): “Roth’s Diagnose
passt auf alle hier aufgeführte Varietäten [der V. persicifolia],
obschon er nur die Pflanze Rupp’s d. i. die Var. γ [= V. elatior]
verstanden hat, wie dies aus Reichenb. Icon. I. p. 88 erhellt.”,
though he referred only to Reichenbach. The later note in the
Specimen florae erlangensis (Schweigger, 1804: 58) is less clear
but may be interpreted as indirect emendation in favour of our
V. stag nina.
Under the provision of the Code, no lectotypification is
possible in the absence of any original material or an illustra-
tion. The only way to fix the use of V. persicifolia remains a ne-
otypification with two options (Arts. 7.7, 9.2 & 9.6, McNeill &
al., 2006). Here, in our opinion, a pragmatic choice may be done
between selecting a type referable to either V. e la tior or V. stag-
nina. In the first case the specimen number W 1889/305915
from the Funkenburg site (locus classicus) or a modern speci-
men may be proposed, in the second case a modern specimen
is the only option. Each of the neotypifications would be in
conflict with a part of the protologue, but we believe that the
choice of the well-preserved Funkenburg plant from the her-
barium Reichenbach fil., referable to V. elatior, would be more
evidence-based than the choice of any V. s t a g n i n a specimen.
However, any neotypification is potentially reversible (Art.
9.17, McNeill & al., 2006) if some original material is discov-
ered, and it should not be used to resolve a long-lasting dispute
like this. Conservation of V. persicifolia with a conserved type
(Art. 14.9, McNeill & al., 2006) referable to V. stagnina would
make it possible to retain this name instead of V. s t a g ni n a but
it would bring about an undesirable nomenclatural change in
some national floras (mainly in Central European countries),
which is in conflict with the aim of conservation as stated in
the Code (Art. 14.2, McNeill & al., 2006). Further, we do not
think that it is reasonable to use this option provided by the
ICBN for such a notoriously confused name still in dispute.
For these reasons we decided not to designate a neotype but to
Fig. .
Label of a Viola elatior specimen from the herbarium Reichen-
bach fil. (W1889/305915) collected near Funkenburg in Leipzig.
1875
Danihelka & al. • Reject Viola montana and V. persicifoliaTA XO N 59 (6) • December 2010: 1869–1878
propose the name V. persicifolia for rejection (Art. 56, McNeill
& al., 2006) in a rejection proposal published simultaneously.
Viola persicifolia Schreb., Spic. Fl. Lips.: [163]. 1771, nom.
utique rej. prop. (Van den Hof & al. in Taxon 59: 1900
– this issue). Ind. loc. (Boehmer, 1750: 190): [Germania.
Saxonia, urbs Lipsia.] “In pascuis, auf der Funckenburg
bey Gonnewitz …” Type: not designated.
TYPIFICATION OF VIOLA STAGNINA
The name Viola stagnina was published by Schultes (1814:
426). The original description is brief and poor in diagnostic
characters, and it refers to a plant with developed capsules
and cleistogamous flowers, collected in late spring or early
summer. A comparison with the descriptions of other violet
species described there (see above) makes it possible to link this
description to V. stag n i n a as understood today. The name has
to be cited as Kit. ex Schult. because only the name is ascribed
to Kitaibel but not the diagnosis and description (Art. 46.4,
McNeill & al., 2006); this is different, however, in the case of
e.g., Cerastium eriophorum Kit. (Schultes, 1814: 694). The cor-
responding Viola specimen sent by P. Kitaibel to J.A. Schultes
is still deposited at M as M-0111205. It bears the original label
“Viola stagnina mihi. In Croatiae locis depressis in quibus
aqua stagnat.”, glued on a newer label of the Royal Munich
Herbarium with a note “A Kitaibelio ipso”. There is also a revi-
sion label of L. Gerstlauer on the sheet: “Viola stagnina Kit.,
Originalstück (Cotypus) von Kitaibel selbst. Rev. Gerstlauer,
1941”. The plants (t wo st ems) re present a late spring or su mme r
gathering of V. stag n i n a as understood today, with cleistoga-
mous f lowe rs and capsules. It may b e selected as lect oty pe. As
Kitaibel used to send duplicates also to Willdenow (Z. Bar ina,
in litt.), we searched also in the herbarium Willdenow; however,
Kitaibel’s gatherings found under V. lactea (B-W04916–07) and
V. montana (B-W04915–03) represent V. pumila (see above;
Röpert, 2000–). There is also a sheet of V. s t a g n i n a in the
herbarium Kitaibel (fascicle IX, nr 191) at BP. It is labelled
“stagnina mihi ignota Willdenowio. In pratis humidis ad Bre-
zovicam, integras plagas ita occupat, ut plantas reliquas fere
omnes extendat” (Z. Barina, in litt.; Jávorka, 1936: 115). The
(unmounted) plants were revised by J. Kirschner in 1984. The
gathering consists of two species: the unbranched plant with
large laminae and stipules is referable to V. elat i o r, whereas
the branched small-leaved plants correspond to V. st a g nina.
Kirschner marked one of the V. stagnina specimens as lectotype
but this lectotypification has never been effectively published.
The plants were probably collected during Kitaibel’s journey
to Croatia in 1794 (Z. Barina, in litt.). As reported by Harmatta
(1962: 339), P. Kitaibel collected plants in Brezovica near Za-
greb in Croatia in the second half of May 1794. However, there
is no direct evidence that the plant in M represents the same
gathering as sheet IX/191 in the herbarium Kitaibel, so the lat-
ter should not be considered iso(lecto)type. Curiously, Croatian
floras do not report V. stagnina (cf. Schlosser & Vukotinović,
1869; Domac, 1994).
Viola stagnina Kit. ex Schult., Oestr. Fl., ed. 2, 1: 426. 1814.
Ind. loc.: “In Morästen, in Sümpfen in Kroatien fand sie
Herr Professor Kitaibel.” Lectotype (designated here):
“Viola stagnina mihi. In Croatiae locis depressis in quibus
aqua stagnat.” ([P. Kitaibel] s.a. M 0111205!).
VIOLA HORNEMANNIANA AND
V. S T IPU LACE A
Two a t tempts to g ive Viola elatior a scientific name are
connected with its cultivation in Nordic botanic gardens.
Hornemann (1815: 234–235) initially included in a catalogue
of plants cultivated in Copenhagen three arosulate violets,
V. canina, V. m o n t a n a and V. lactea, the latter with “V. m o n t .
persicifol.” in synonymy and also w ith va r. “β. Ruppii”, b ear ing
white flowers. The cultivated specimens should have differed
from native ones by more erect stems (“caulibus strictiori-
bus”). He returned to the topic in Addenda (p. 958), describing
V. stricta as new species: “Viola stricta mihi: (9–10 [= to be
inserted between V. montana and V. l a c t e a]) caulibus erectis
pedunculisque strictis, foliis ovato-lanceolatis, stipulis utrinque
basi inciso-serratis”. Hornemann further added that the plant,
introduced in 1807, may be cultivated outside and that V. mon-
tana persicifolia of his Enumeratio plantarum horti botanici
hafniensis (Hornemann, 1804: 27, 1809: 14) belongs here. He
also stressed that this new species is different from his V. lactea,
while V. montana from the Copenhagen garden is at most only
a variety of V. ca n in a. In our opinion, the plant cultivated in
the Copenhagen garden may be identified as V. elat i o r. This is
supported both by the opinions of contemporary botanists (e.g.,
Reichenbach, 1832: 708, 1839–1840: 47) and by a specimen of
V. elatior found at C, originating from herbarium Hornemann,
collected in 1800 in the Copenhagen botanic garden and identi-
fied by Hornemann as Viola montana persicifolia (based on the
handwriting, at least the epithet persicifolia may be ascribed
to Hornemann himself; H.Æ. Pedersen, in litt.). The plant was
revised by J.C. Clausen (H.Æ. Pedersen, in litt.) as V. e la t i o r.
This gathering represents plants cultivated in the garden be-
fore “V. str i c t a ” was acquired in 1807; as they are cited in the
protologue (see above), we propose this specimen as lectotype
of the name V. str i c t a Hornem. However, this name is illegiti-
mate due to its homonymy with V. stricta (Vent.) Poir. (1808),
while V. str icta Muhl. (Muhlenberg, 1793: 178; see also Pursh,
1813: 175) is illegitimate as published without any description
or diagnosis.
The later misinterpretation of V. st ricta Hornem. as refer-
ring to V. s tagnina may have been caused by pl. 1812 in Flora
danica (Hornemann, 1825; see also p. 4 for localities) and sup-
ported by the fact that V. e la t ior does not occur in Denmark.
Despite that, both Fries (1828: 275, 278–279) and Becker (1910:
62–63) clearly distinguished between the original meaning
of the name and its later interpretation. As can be observed
in herbaria with historical collections, the illegitimate name
V. stricta was then used for many decades as a “litter bin” for
erect morphotypes of V. c a n i n a , sometimes also for V. stag-
nina and (putative) hybrids of both. This is rather surprising
1876
TA XO N 59 (6) • December 2010: 1869–1878Danihelka & al. • Reject Viola montana and V. persicifolia
because the illegitimate status of V. s tr icta Hornem. was soon
recognised, and the name was replaced by a nomen novum
(Roemer & Schultes, 1819: 370) (Fig. 4), V. hornemanniana
Schult. Schultes completely adopted the text from Hornemann
(1815), adding only a short note about the shape of stipules.
Another attempt to grant Viola elatior a suitable name
was that by Fries. Describing the flora of the Halland prov-
ince of Sweden (Fries, 1817–1819), he recorded three native
species of arosulate violets: Viola canina, V. R u p p i i (“Viola
montana. Vulgo” as “Media inter V. mirabile m & caninam”)
and V. persicifolia, confined to the proximity of water bod-
ies and corresponding to our V. stagnina. Fries regarded the
latter as conspecific with V. montana L. but due to its habitat
requirements he considered the name unsuitable and source of
confusion. Then he drew his attention to a similar, but different
enough species cultivated in the Lund botanic garden, referred
to as “V. persicifolia major”, proposing the name V. st ipularis
for it. The brief description may be unambiguously attributed
to V. e l a t i o r. However, the name is illegitimate due to homon-
ymy (Art. 53.3, McNeill & al., 2006) with the South American
V. stipularis Sw. (Prodr.: 117. 1788).
Immediately after its publication, the illegitimate name
V. stipularis became a source of confusion. The species was
accepted by Hartman (1820: 110), but the spelling of the specific
epithet was changed to “stipulacea”, rather by a writing error
than intentionally. As this altered name, V. s ti pulacea, was also
ascribed to Fries, it was clearly not intended as a replacement
name (avowed substitute; see Art. 7.3, McNeill & al., 2006).
Still, this minor change in the epithet removed the problem of
homonymy as defined by the Code, and the name, accompanied
by a brief description, has to be considered as a legitimate name
of a new species. Though the epithets “stipularis” and “stipula-
cea” are very similar and easy to confuse, as demonstrated by
a series of mistakes by Hartman, Fries and Nikitin described
in this paper, they are not sufficiently similar to be treated as
homonyms (see Art. 53.3, McNeill & al., 2006). In the proto-
logue of V. st ipulacea two elements are cited: a Curtis plate
of V. montana (probably vol. 39, pl. 1595; Sims 1813–1814),
clearly corresponding to V. elat i or, and an Ahlquist specimen
from Öland (“Ö. Ahlquist”). This reference may concern speci-
men UPS 220503, labelled “Viola persicifolia. Rstn 18” (with
later remarks “[Runsten] Ahlqvist” in a different handwriting),
stamped “Herb. Hartman”, eligible as lectotype.
Viola hornemanniana Schult. (1819) and V. stipulacea
Hartm. (1820) both represent earlier legitimate names refer-
ring to the species generally known as V. elat i o r Fr. (1828).
However, their use is undesirable because it would seriously
disturb nomenclatural stability as one of the aims of botanical
nomenclature, and therefore a conservation proposal is pub-
lished simultaneously in the same Taxon issue.
Viola stricta Hornem., Hort. Bot. Hafn.: 958. 1815, nom. il-
leg. (non V. st ricta (Vent.) Poir., Encycl. (Lamarck) 8:
648. 1808). Ind. loc.: [Hortus regius botanicus hafnien-
sis, Hafnia]. Lectotype (designated here): “Viola montana
[manu?] persicifolia [manu Hornemann]; verso folio: “Ex
Horto Botan. Haun. 1800.” [manu?] “HB HORN” [impres-
sum] (Hornemann? C sine no; foto!).
≡ Viola hornemanniana Schult. in Roem. & Schult., Syst. Veg.,
ed. 15 bis, 5: 370. 1819.
= Viola stipularis Fr., Fl. Hall.: 47. 1817, nom. illeg. (non V. stip -
ularis Sw., Prodr.: 117. 1788, nec non V. st i p u l a r i s Cav.,
Icon. 6: 21. 1800, quod est V. c a p i llari s Pers., Syn. Pl. 1:
256. 1805.). Ind. loc.: “Est V. persicifo lia major. Horti Lun-
dens. [= Lund]”. Type: not designated.
= Viola stipulacea Hartm., Handb. Skand. Fl.: 110, 1820. Ind.
loc.: “Ö [= Suecia, ins. Öland]”. Lectotype (designated
here): “Viola persicifolia. Rstn 18” cum adnotatione pos-
teriore “[Runsten] Ahlqvist” manu alia (UPS 220503!).
TYPIFICATION OF VIOLA ELATIOR
Viola elatior was described by Fries (1828: 277) on the
basis of plants from Öland. The diagnosis and description
clearly apply to V. e l a t i o r as understood today. This is also
supported by the fact that Fries at the same time distinguished
V. lacte a (= V. pu mila) and V. persicifolia (= V. s t a g n i n a). He
found V. elat i o r during his visit to Öland in 1818 (cf. p. 276)
and immediately noted the distinctive tall stature of this spe-
cies: “Statura elatiori mox dignoscitur; nomen inde a primo
Clusio sumtum & mihi primo inventionis momento mox in
mentem venit” (p. 278). In the protologue two gatherings are
cited, the first made by Fries himself and the second by A. Ahl-
quist. The corresponding specimens are found at UPS, labelled
“Viola persicifolia. Rstn 18” (with later remarks “[Runsten]
Ahlqvist” in a different handwriting; UPS 220503) and “Viola
elatior. Ölandia ad Allgutsrum 1824. E. Fries scripsit.” (UPS
220505), both stamped “Herb. Hartman”. Also the third speci-
men found at UPS and labelled “Viola elatior Fries. Ölandia.
1818. Haec sunt duo specimina prima in Suecia a me detexta”
(UPS 220509), stamped “Herb. E. Fries”, may be considered
original material. Plants on all three sheets represent V. elatior
as currently understood.
Fig. .
Protologue of Viola
hornemanniana (Roemer &
Schultes, 1819–1820: 370).
1877
Danihelka & al. • Reject Viola montana and V. persicifoliaTA XO N 59 (6) • December 2010: 1869–1878
Becker, W. 1910. Violae europaeae. Dresden-N.: Heinrich.
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Boehmer, G.R. 175 0. Flora Lipsiae indigena. Leipzig: Joh. Gottfried
Dyck.
Borbás, V. 1892. Violarieen. Pp. 161–226 in: Hallier, E. (ed.), Koch’s
Synopsis der deutschen und Schweizer Flora, ed. 3, vol. 1. L eipzi g:
O.R. Reisland.
Chen, Z., Zang, Q., Hideaki, O. & Nik itin, V.V. 2007. Viola Lin naeus,
Sp. Pl. 2: 933. 1753. Pp. 74–111 in: Wu Zhengyi & Raven, P.H.
(ed s.), Flora of China, vol. 13. Beijing: Science Press; St. Louis:
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Dandy, J.E. 1970. Annotated list of the new names published in
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Domac, R. 1994. Flora Hrvatske. Zagreb: Školska kniga.
Eckstein, R.L., Hölzel, N. & Danihelka, J. 2006. Biological Flora of
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Fries, E.[M.] 1817–1819. Flora hallandica. Lund: Typis Berlingianis.
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thorum: ex officina berlingiana.
Gerstlauer, L. 1943. Vorschläge zur Systematik der einheimischen
Vei lc hen . Ber. Bayer. Bot. Ges. 26: 12–55.
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Albers. Stuttgart: Bundesamt für Naturschutz.
Haller, A. 1768. Historia stirpium indigenarum Helvetiae inchoata.
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Sachsens. Dresden: Sächsisches Landesamt für Umwelt und Ge-
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Közlem. 49: 334–345.
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ensis. Havniae.
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horti botanici hafniensis. Havniae.
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floodplain violets in the city and its surroundings. Walter Gutermann
with his unpublished manuscript “Über unnötige und notwendige
Name nsä nderungen de r mitteleuropä ische n Flo ra” provid ed an i mpor-
tant incentive to this study. We are also grateful to both anonymous
reviewers and to John McNeill for their helpful suggestions. The work
done by JD was supported by the Ministry of Education of the Czech
Republic (grants MSM0021622416 and LC06073) and by the long-
term research plan AV0Z60050516 of the Institute of Botany, Czech
Academy of Sciences. The work done by TM was supported by the
Norwegian Research Council (grant 170832: “Allopolyploid evolution
in plants: patterns and processes within the genus Viola”).
LITERATURE CITED
Nikitin (1988) analysed the protologue of V. e lat ior (Fries,
1828: 277–279) and arg ued that this name has to be considered
illegitimate because Fries included in its synonymy V. m o n -
tana L. (cited from the second edition of Species plantarum
in accordance with contemporary practice) without excluding
the type (Art. 52.1, McNeill & al., 2006). However, in the same
work by Fries, V. m o n t a n a served as basionym for V. canina γ
[= var.] montana (L.) Fr. (p. 273); here, V. monta n a was cited
from Flora suecica (Linnaeus, 1755). As already shown by
Kirschner & Skalický (1989), Nikitin’s reasoning is not correct
because Fries (l.c.) excluded the type of V. montana by impli-
cation, as described in Art. 52.2. Ex. 8 (McNeill & al., 2006).
The fact that Fries cited V. montana from different Linnaean
works is unimportant because a name refers to the same type
regardless of the work from which it is cited.
Further, a later lectotypification of V. montana by Niki-
tin (1988) is not retroactive (Art. 52.2. Note 2, McNeill & al.,
2006); i n other words, it cannot make a n ame publ ished in 1828
nomenclaturally superfluous and, consequently, illegitimate.
Nikitin (1995) disagreed and repeated his arguments
against the legitimate status of V. e l a t i o r and added another
reason: in the synonymy of V. e l a t ior (Fries, 1828: 278),
“V. st i p u l a c e a Hartm., 1820 and V. e l a t ior Link, 1821” are in-
cluded, both earlier and validly published, and in consequence
impossible to reject. “Therefore, if somebody does not agree
yet that it is necessary to return to V. montana in its original
sense, he will have to refrain from the use of V. elatior Fr. and
use the earlier name V. stipulacea Hartman instead. The name
V. elatior ascribed to Link, 1821, not to Fries, 1828, should be
included in its synonymy.” (Nikitin 1995: 89; translated from
Russian). However, neither of these statements are entirely
correct. Link (1821: 241) only wrote in comments on his no
2314. V. persicifolia “Roth”: “Differt a V. elata [sic!] Fries foliis
latioribus, ovata basi, non scabris, bracteis minutis sub flore.”
This is by no means a valid publication of a name, as already
noted by Hylander (1945). Further, what Fries actually did was
to include “V. stipularis. Fr. Hall. p. 47. Hartm.! Scand.”, not
V. stipulacea, in the synonymy of V. e l a t i o r (p. 278). These
facts demonstrate that Nikitin’s conclusions are wrong, and that
V. elatior indeed does represent a legitimate name.
Viola elatior Fr., Novit. Fl. Suec. Alt.: 277. 1828, nom. cons.
prop. (Danihelka & al. in Taxon 59: 1902 – this issue). Ind.
loc.: “In Ölandiae tractu silvatico inter Algutsrum & Tveta
uber rime leg i; Ad Runstens Canal rariorem Rev. A hlquist
detexit.” Lectotype (designated here): “Viola elatior. Ölan-
dia ad Allgutsrum 1824. E. Fries scripsit” (UPS 220505!).
ACKNOWLEDGEMENTS
We a re g ratef ul t o Zolt án Bar ina ( BP), H enr ik Æren lun d Pede rsen
(C), Werner Nezadal (ER), Franz Schuhwerk, Hajo Esser (both M) and
Ernst Vitek (W) for their help during our search for nomenclaturally
important herbarium specimens and additional information. Kanchi
Gandhi made us available a copy of Muehlenberg’s paper. Peter Gutte
informed us about the former flora of Leipzig and former localities of
1878
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